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<document id="86987D44921C8A50CBAC79708AD41250" ID-DOI="10.1093/zoolinnean/zlad198" ID-ISSN="0024-4082" ID-Zenodo-Dep="14423193" ID-ZooBank="E0A267D-6195-4EC4-92B1-757DD5956C85" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="carolina" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="carolina" checkinTime="1734029692247" checkinUser="plazi" docAuthor="Hoch, Hannelore, Porter, Megan L., Slav, Christv M., Slav, Michael E., Steck, Mireille &amp; Chong, Rebecca A." docDate="2024" docId="03E3EA7C3150BF011C74FE5DFA75F865" docLanguage="en" docName="zlad198.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (3)" docSource="https://doi.org/10.1093/zoolinnean/zlad198" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Iolania Kirkaldy 1902" docType="treatment" docVersion="1" lastPageNumber="5" masterDocId="FFDA92043154BF051D4CFFB3FFD2FFD2" masterDocTitle="From the dark side of paradise: a new natural replication of cave planthopper evolution from Hawaiian lava tubes (Hemiptera: Fulgoromorpha: Cixiidae)" masterLastPageNumber="13" masterPageNumber="1" pageNumber="5" updateTime="1734447688994" updateUser="carolina">
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<mods:title id="0458AF58AE4F17687E43DA9E27442231">From the dark side of paradise: a new natural replication of cave planthopper evolution from Hawaiian lava tubes (Hemiptera: Fulgoromorpha: Cixiidae)</mods:title>
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<mods:namePart id="534650CF450EE8123A4062A2AA92EF8B">Hoch, Hannelore</mods:namePart>
<mods:affiliation id="4BB48D2650119B4D5CC164200043614D">Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversitv Science, Humboldt-Universität zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germanv</mods:affiliation>
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<mods:namePart id="B481192EB1651B8446F9251A933AA715">Porter, Megan L.</mods:namePart>
<mods:affiliation id="D7BAEE2892023C577B7CD9E2E37F8C2B">School of Life Sciences, Universitv of Hawaiʻi at Mānoa, Honolulu, HI 96822, USA</mods:affiliation>
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<mods:namePart id="B6ABCDFF1CE9F500E0FA87AE03A7C8BE">Slav, Christv M.</mods:namePart>
<mods:affiliation id="70AE7DB5CC18A8F8F572887C04ED7163">Ŋe Sustainabilitv Consortium, Julie Ann Wriglev Global Futures Laboratorv, Arizona State Universitv, Tempe, AZ 85281, USA</mods:affiliation>
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<mods:namePart id="D74CEBEF4DB42835F1C209D14F53C9E2">Slav, Michael E.</mods:namePart>
<mods:affiliation id="1C77199291E54161D6022B203C69B484">Arkansas Field Office, Ŋe Nature Conservancv, Liưle Rock, AR 72201, USA</mods:affiliation>
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<mods:namePart id="5D2B96DD4C687844DD1EECC8DA39F4FB">Steck, Mireille</mods:namePart>
<mods:affiliation id="F0CD935A1B9E46531622B9C149FA9B5A">School of Life Sciences, Universitv of Hawaiʻi at Mānoa, Honolulu, HI 96822, USA</mods:affiliation>
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<mods:namePart id="A677474EB62C0879312CBE1A0B106CAE">Chong, Rebecca A.</mods:namePart>
<mods:affiliation id="D045C1C1DFA2306B06072B376C6BC8D8">School of Life Sciences, Universitv of Hawaiʻi at Mānoa, Honolulu, HI 96822, USA</mods:affiliation>
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<subSubSection id="C35008E13150BF011C74FE5DFD80FDDA" box="[312,594,494,521]" pageId="4" pageNumber="5" type="nomenclature">
<paragraph id="8BF55B6A3150BF011C74FE5DFD80FDDA" blockId="4.[312,594,494,521]" box="[312,594,494,521]" pageId="4" pageNumber="5">
<heading id="D0BDEC063150BF011C74FE5DFD80FDDA" bold="true" box="[312,594,494,521]" centered="true" fontSize="11" level="2" pageId="4" pageNumber="5" reason="2">
<emphasis id="B93E87783150BF011C74FE5DFD80FDDA" bold="true" box="[312,594,494,521]" pageId="4" pageNumber="5">
<taxonomicName id="4C4A20E93150BF011C74FE5DFE33FDDA" ID-CoL="54QG" ID-ENA="2831207" authority="Kirkaldy" authorityName="Kirkaldy" authorityYear="1902" box="[312,481,494,521]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783150BF011C74FE5DFE52FDDB" bold="true" box="[312,384,494,521]" italics="true" pageId="4" pageNumber="5">Iolania</emphasis>
Kirkaldy
</taxonomicName>
19 0 2: 118
</emphasis>
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<subSubSection id="C35008E13150BF011DCDFD93FDBAFDE5" box="[129,616,543,567]" pageId="4" pageNumber="5" type="type_taxon">
<paragraph id="8BF55B6A3150BF011DCDFD93FDBAFDE5" blockId="4.[129,616,543,567]" box="[129,616,543,567]" pageId="4" pageNumber="5">
<emphasis id="B93E87783150BF011DCDFD93FF2EFDE5" box="[129,252,544,567]" italics="true" pageId="4" pageNumber="5">Type species:</emphasis>
<taxonomicName id="4C4A20E93150BF011C4DFDACFDB1FDE5" authority="Kirkaldv 1902: 119" authorityName="Kirkaldv" authorityPageNumber="119" authorityYear="1902" box="[257,611,543,567]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783150BF011C4DFDACFE45FDE5" box="[257,407,543,567]" italics="true" pageId="4" pageNumber="5">Iolania perkinsi</emphasis>
<bibRefCitation id="EFDB269B3150BF011CD0FDACFDB1FDE5" author="Kirkaldv GW" box="[412,611,543,567]" pageId="4" pageNumber="5" pagination="93 - 174" refId="ref9471" refString="Kirkaldv GW. Hemiptera. In: Sharp D (ed.), Fauna Hawaiiensis, Vol. III, 2. Cambridge: Cambridge Universitv Press, 1902; 93 - 174, pls 4, 5." type="journal article" year="1902">Kirkaldv 1902: 119</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C35008E13150BF011C22FDE4FA6DFB7B" pageId="4" pageNumber="5" type="diagnosis">
<paragraph id="8BF55B6A3150BF011C22FDE4FDCEFDA3" blockId="4.[129,778,599,1194]" box="[366,540,599,625]" pageId="4" pageNumber="5">
<emphasis id="B93E87783150BF011C22FDE4FDCEFDA3" bold="true" box="[366,540,599,625]" pageId="4" pageNumber="5">Genus diagnosis</emphasis>
</paragraph>
<paragraph id="8BF55B6A3150BF011DCDFDCEFD1EFBDF" blockId="4.[129,778,599,1194]" pageId="4" pageNumber="5">
Hoch (2006) provided a detailed diagnosis of the genus
<taxonomicName id="4C4A20E93150BF011DCDFD2EFF14FD67" authorityName="Kirkaldy" authorityYear="1902" box="[129,198,669,693]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783150BF011DCDFD2EFF14FD67" box="[129,198,669,693]" italics="true" pageId="4" pageNumber="5">Iolania</emphasis>
</taxonomicName>
, which then contained exclusivelv epigean species. The cavernicolous species
<taxonomicName id="4C4A20E93150BF011C24FD0EFDD0FD06" box="[360,514,700,724]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E87783150BF011C24FD0EFDD0FD06" box="[360,514,700,724]" italics="true" pageId="4" pageNumber="5">I. frankanstonei</emphasis>
</taxonomicName>
from
<collectingRegion id="498E95883150BF011F0AFD0FFD5CFD06" box="[582,654,700,724]" country="United States of America" name="Hawaii" pageId="4" pageNumber="5">Hawaii</collectingRegion>
Island (described below) features characters pertaining to its external morphologv that have evolved in the course of adaptation bv the species to its subterranean habitat (troglomorphies). These include the reduction of compound eves, tegmina and hindwings, and bodilv pigmentation. As has been shown in other cavernicolous planthopper taxa (
<bibRefCitation id="EFDB269B3150BF011CA8FCCBFD2BFC42" author="Hoch H &amp; Howarth FG" box="[484,761,888,912]" pageId="4" pageNumber="5" pagination="377 - 402" refId="ref8822" refString="Hoch H, Howarth FG. Six new cavernicolous cixiid planthoppers in the genus Solonaima from Australia (Homoptera: Fulgoroidea). Systematic Entomology 1989; 14: 377 - 402. hups: // doi. org / 10.1111 / j. 1365 - 3113.1989. tb 00291. x" type="journal article" year="1989">Hoch and Howarth 1989</bibRefCitation>
), these reductions of single traits can influence the configuration of other traits, such as head width, sculpturing (e.g. carination), and general bodv shape. It therefore seems appropriate to provide a separate diagnosis for cavernicolous
<taxonomicName id="4C4A20E93150BF011F7FFC46FD1EFBDF" box="[563,716,1013,1037]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B93E87783150BF011F7FFC46FDAAFBDF" box="[563,632,1013,1037]" italics="true" pageId="4" pageNumber="5">Iolania</emphasis>
species.
</taxonomicName>
</paragraph>
<paragraph id="8BF55B6A3150BF011DD0FBA7FA8EFF15" blockId="4.[129,778,599,1194]" lastBlockId="4.[825,1474,144,199]" pageId="4" pageNumber="5">
All species of the genus
<taxonomicName id="4C4A20E93150BF011CC0FBA7FE03FBFE" authorityName="Kirkaldy" authorityYear="1902" box="[396,465,1044,1068]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783150BF011CC0FBA7FE03FBFE" box="[396,465,1044,1068]" italics="true" pageId="4" pageNumber="5">Iolania</emphasis>
</taxonomicName>
are easilv distinguishable from the other Hawaiian cixiid genus,
<taxonomicName id="4C4A20E93150BF011CA4FB87FDE0FB9E" authorityName="Stal" authorityYear="1862" box="[488,562,1076,1100]" class="Insecta" family="Cixiidae" genus="Oliarus" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783150BF011CA4FB87FDE0FB9E" box="[488,562,1076,1100]" italics="true" pageId="4" pageNumber="5">Oliarus</emphasis>
</taxonomicName>
, bv their tricarinate pronotum (vs. pentecarinate in
<taxonomicName id="4C4A20E93150BF011C91FBE0FDF5FBB9" authorityName="Stal" authorityYear="1862" box="[477,551,1107,1131]" class="Insecta" family="Cixiidae" genus="Oliarus" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783150BF011C91FBE0FDF5FBB9" box="[477,551,1107,1131]" italics="true" pageId="4" pageNumber="5">Oliarus</emphasis>
</taxonomicName>
) and from all other known Cixiini bv the morphological configuration of the male copulatorv organ, the aedeagus (shaft bilaterallv svmmetrical, smooth; flagellum rigid, short, not movable against shaft; phallotreme near base of flagellum, exposed dorsallv).
</paragraph>
<paragraph id="8BF55B6A3150BF011E2BFF5BFB60FEF2" blockId="4.[871,1428,231,288]" pageId="4" pageNumber="5">
<emphasis id="B93E87783150BF011E2BFF5BFB60FEF2" italics="true" pageId="4" pageNumber="5">
Diagnosis of epigean species of
<taxonomicName id="4C4A20E93150BF0119CDFF54FB19FED3" authorityName="Kirkaldy" authorityYear="1902" box="[1153,1227,231,257]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">Iolania</taxonomicName>
(see also Hoch 2006: 303–304)
</emphasis>
</paragraph>
<paragraph id="8BF55B6A3150BF011E75FE9EFAD6FD29" blockId="4.[825,1475,301,763]" pageId="4" pageNumber="5">
Total length (tip of head to distal margin of tegmina): males,
<quantity id="4CB2F68F3150BF0118D9FE9EFC54FEB1" metricMagnitude="-3" metricUnit="m" metricValue="5.55" metricValueMax="6.3" metricValueMin="4.8" pageId="4" pageNumber="5" unit="mm" value="5.55" valueMax="6.3" valueMin="4.8">4.8– 6.3 mm</quantity>
; females,
<quantity id="4CB2F68F3150BF011EA7FEFFFBB6FEB1" box="[1003,1124,332,356]" metricMagnitude="-3" metricUnit="m" metricValue="6.65" metricValueMax="7.1" metricValueMin="6.2" pageId="4" pageNumber="5" unit="mm" value="6.65" valueMax="7.1" valueMin="6.2">6.2–7.1 mm</quantity>
. Moderatelv large cixiids of robust appearance (
<figureCitation id="137147EF3150BF011EF2FED8FBF6FE51" box="[958,1060,363,387]" captionStart="Figure 3" captionStartId="3.[113,178,1836,1860]" captionTargetBox="[121,1451,145,1807]" captionTargetId="figure-6@3.[120,1452,144,1808]" captionTargetPageId="3" captionText="Figure 3. Images of epigean (Iolania perkinsi) and cavernicolous (Iolania Ƒankanstonei) Iolania species, illustrating external morphological features.A, B, Iolania perkinsi Kirkaldv habitus (A) and head and thorax (B), dorsal aspect. C, D, Iolania ssankanstonei habitus (C) and head and thorax (D), dorsal aspect. Specimen from OHS entrance (images bv B. Schurian)." figureDoi="http://doi.org/10.5281/zenodo.14423203" httpUri="https://zenodo.org/record/14423203/files/figure.png" pageId="4" pageNumber="5">Fig. 3A, B</figureCitation>
); tegmina ~2.6× longer than maximallv wide, more or less shallowlv tectiform, surpassing tip of abdomen with approximatelv one-third of total length. Costal veins of tegmina nearlv parallel in dorsal aspect. Tegmina and hindwings translucent to hvaline. Tegmina with colour pattern variable within populations, ranging from hvaline with little recognizable patterning to those with distinct dark brown spots or stripes, especiallv at the tegmen base and in the distal third. Granules on veins usuallv brownish. Bodv coloration inconspicuous; intraspecific variation ranging from vellowish brown to darker brown. Coloration of bodv and tegmina usuallv slightlv darker in females. Vertex short, with lateral margins diverging slightlv posteriorlv. Hind tibiae laterallv with three minute spines.
</paragraph>
<paragraph id="8BF55B6A3150BF011E02FC81FAC0FCB8" blockId="4.[825,1475,817,1193]" pageId="4" pageNumber="5">
<emphasis id="B93E87783150BF011E02FC81FAC0FCB8" italics="true" pageId="4" pageNumber="5">
Diagnosis of cavernicolous species of
<taxonomicName id="4C4A20E93150BF0119D1FC82FB35FC99" authorityName="Kirkaldy" authorityYear="1902" box="[1181,1255,817,843]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">Iolania</taxonomicName>
(
<figureCitation id="137147EF3150BF0119BAFC81FA88FC99" box="[1270,1370,817,843]" captionStart="Figure 3" captionStartId="3.[113,178,1836,1860]" captionTargetBox="[121,1451,145,1807]" captionTargetId="figure-6@3.[120,1452,144,1808]" captionTargetPageId="3" captionText="Figure 3. Images of epigean (Iolania perkinsi) and cavernicolous (Iolania Ƒankanstonei) Iolania species, illustrating external morphological features.A, B, Iolania perkinsi Kirkaldv habitus (A) and head and thorax (B), dorsal aspect. C, D, Iolania ssankanstonei habitus (C) and head and thorax (D), dorsal aspect. Specimen from OHS entrance (images bv B. Schurian)." figureDoi="http://doi.org/10.5281/zenodo.14423203" httpUri="https://zenodo.org/record/14423203/files/figure.png" pageId="4" pageNumber="5">Figs 3C, D</figureCitation>
,
<figureCitation id="137147EF3150BF01182FFC82FAA2FC99" box="[1379,1392,817,843]" captionStart="Figure 4" captionStartId="5.[113,178,1954,1978]" captionTargetBox="[359,1212,820,1927]" captionTargetId="figure-313@5.[356,1216,817,1928]" captionTargetPageId="5" captionText="Figure 4. Iolania Ƒankanstonei, male, wing morphologv. A, right tegmen. B, right hind wing.Scale bar:0.5 mm. Male from CKM entrance." figureDoi="http://doi.org/10.5281/zenodo.14423205" httpUri="https://zenodo.org/record/14423205/files/figure.png" pageId="4" pageNumber="5">4</figureCitation>
,
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), as represented by
<taxonomicName id="4C4A20E93150BF01193DFCE2FAC0FCB8" box="[1137,1298,848,874]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="ssankanstonei">I. frankanstonei</taxonomicName>
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Total length: males,
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; females,
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. Tegmina 2× longer than maximallv wide, shallowlv tectiform, not surpassing tip of anal segment.Costal veins of tegmina in dorsal aspect convex, together creating an oval outline (
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). Vertex with lateral margins stronglv diverging posteriorlv and with posterior margin shallowlv concave; compound eves small, pronotum in dorsal aspect mediallv narrow and laterallv expanded. Compound eves with light red pigmentation, coloration of bodv vellowish to light brown, tegmina with darker spots and stripes, variable among specimens studied. Hind tibiae laterallv with three minute spines.
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Kev to
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species
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(modified from Hoch 2006)
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(1) Tegmina longer than twice their maximal width, well surpassing tip of abdomen; compound eves large and darklv pigmented; vertex with lateral margins slightlv diverging posteriorlv and with posterior margin deeplv concave; pronotum in dorsal aspect narrow throughout; bodv size of males
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, females
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............................................................. 2
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- Tegmina twice as long as maximallv wide, not surpassing tip of anal segment; compound eves small and reddish; vertex with lateral margins stronglv diverging posteriorlv and with posterior margin shallowlv concave; pronotum in dorsal aspect mediallv narrow and laterallv expanded; bodv size of males
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, females
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.................................................... .....................................................................................................................................................................
<emphasis id="B93E87783150BF01193CFA54FA7AF9D3" bold="true" box="[1136,1448,1511,1537]" pageId="4" pageNumber="5">
<taxonomicName id="4C4A20E93150BF01193CFA54FA80F9D2" authority="Fankanstonei" authorityName="Fankanstonei. B" authorityYear="2024" box="[1136,1362,1511,1536]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus" status="sp. nov.">
<emphasis id="B93E87783150BF01193CFA54FA80F9D2" bold="true" box="[1136,1362,1511,1536]" italics="true" pageId="4" pageNumber="5">Iolania frankanstonei</emphasis>
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<taxonomicNameLabel id="A20D3A033150BF01181AFA5AFA7AF9D3" box="[1366,1448,1513,1537]" pageId="4" pageNumber="5" rank="species">sp. nov.</taxonomicNameLabel>
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(2) Male genitalia with aedeagus svmmetrical; anal segment without spinose processes ................................................................... 3
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- Aedeagus with asvmmetrical paired spines; anal segment caudallv bent ventrad, with two distinct ventrolateral spinose processes ........................................................................................................................................................
<taxonomicName id="4C4A20E93150BF01192AF9F4FA75F98C" authority="Hoch, 2006" authorityName="Hoch" authorityYear="2006" box="[1126,1447,1606,1631]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="kraussohana">
<emphasis id="B93E87783150BF01192AF9F4FAFBF98D" box="[1126,1321,1607,1631]" italics="true" pageId="4" pageNumber="5">Iolania kraussohana</emphasis>
Hoch, 2006
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(3) Shaft of aedeagus in lateral aspect curved dorsad; flagellum without sculpturing; gonostvli not ventrallv ridged .................. 4
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- Shaft of aedeagus straight; flagellum sculptured with numerous distinct tuberculous protuberances; gonostvli ventrallv ridged .............................................................................................................................................................................................................. 5 (4) Paired spines of aedeagus foliatelv dilated ...................................................................................................................
<taxonomicName id="4C4A20E93150BF0119B3F977FA75F90E" authorityName="Giffard" authorityYear="1925" box="[1279,1447,1732,1756]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="mauiensis">
<emphasis id="B93E87783150BF0119B3F977FA75F90E" box="[1279,1447,1732,1756]" italics="true" pageId="4" pageNumber="5">Iolania mauiensis</emphasis>
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<keyLead id="30BBF85F3150BF011DD7F957FA75F8C8" pageId="4" pageNumber="5">
- Paired spines of aedeagus not dilated, arm-like, curved dorsad at midlength ....................................................
<taxonomicName id="4C4A20E93150BF0119BFF950FA75F929" authorityName="Giffard" authorityYear="1925" box="[1267,1447,1763,1787]" class="Insecta" family="Cixiidae" genus="Iolanaia" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="lanaiensis">
<emphasis id="B93E87783150BF0119BFF950FA75F929" box="[1267,1447,1763,1787]" italics="true" pageId="4" pageNumber="5">Iolanaia lanaiensis</emphasis>
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(5) Gonostvli with caudal margin triangular; tips of aedeagal spines directed straight basad or mediobasad ................................. 6
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<keyLead id="30BBF85F3150BF011DD7F891FA7AF8AB" pageId="4" pageNumber="5">
- Gonostvli with caudal margin rounded; tips of aedeagal spines directed dorsad ..................................................
<taxonomicName id="4C4A20E93150BF01185DF891FA75F8E8" authorityName="Kirkaldv" authorityYear="1902" box="[1297,1447,1826,1850]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783150BF01185DF891FA75F8E8" box="[1297,1447,1826,1850]" italics="true" pageId="4" pageNumber="5">Iolania perkinsi</emphasis>
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(6) Male anal segment distallv with produced lateroventral corners; shaft of aedeagus in lateral aspect with ventral margin distinctlv narrowing in distal half; tips of aedeagal spines curved mediad .................................................................
<taxonomicName id="4C4A20E93150BF01184CF8D2FA7AF8AB" authorityName="Giffard" authorityYear="1925" box="[1280,1448,1889,1913]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="oahuensis">
<emphasis id="B93E87783150BF01184CF8D2FA7AF8AB" box="[1280,1448,1889,1913]" italics="true" pageId="4" pageNumber="5">Iolania oahuensis</emphasis>
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<keyLead id="30BBF85F3150BF011DD7F833FA75F865" pageId="4" pageNumber="5">
- Male anal segment in caudal aspect ventrallv rounded; shaft of aedeagus in lateral aspect not narrowing towards apex; tips of aedeagal spines directed straight basad .....................................................................................................................
<taxonomicName id="4C4A20E93150BF0119A3F82CFA75F865" authorityName="Giffard" authorityYear="1925" box="[1263,1447,1951,1975]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="koolauensis">
<emphasis id="B93E87783150BF0119A3F82CFA75F865" box="[1263,1447,1951,1975]" italics="true" pageId="4" pageNumber="5">Iolania koolauensis</emphasis>
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<paragraph id="8BF55B6A3151BF001D8FFF3DFDF1FF1B" blockId="5.[195,680,142,201]" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001D8FFF3DFDF1FF1B" bold="true" pageId="5" pageNumber="6">
<taxonomicName id="4C4A20E93151BF001D8FFF3DFD85FF78" ID-CoL="54QG" ID-ENA="2831207" authority="Fankanstonei Hoch &amp; Porter" authorityName="Fankanstonei Hoch &amp; Porter" authorityYear="2024" box="[195,599,142,170]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus" status="sp. nov.">
<emphasis id="B93E87783151BF001D8FFF3DFE67FF78" bold="true" box="[195,437,142,170]" italics="true" pageId="5" pageNumber="6">Iolania frankanstonei</emphasis>
Hoch &amp; Porter
</taxonomicName>
<taxonomicNameLabel id="A20D3A033151BF001F10FF23FD7AFF78" box="[604,680,144,170]" pageId="5" pageNumber="6" rank="species">sp. nov.</taxonomicNameLabel>
(
<figureCitation id="137147EF3151BF001C18FF1CFE70FF1B" box="[340,418,175,201]" captionStart="Figure 1" captionStartId="1.[113,178,1952,1976]" captionTargetBox="[115,1457,1051,1922]" captionTargetId="figure-509@1.[114,1458,1050,1923]" captionTargetPageId="1" captionText="Figure 1. Habitus of closelv related epigean and cavernicolous planthopper species. A, Iolania perkinsi (epigean). B, Iolania Ƒankanstonei (cavernicolous). C, Oliarus cf. filicicola (epigean). D, Oliarus polyphemus (cavernicolous).Images of live specimens bv M.E. Slav." figureDoi="http://doi.org/10.5281/zenodo.14423195" httpUri="https://zenodo.org/record/14423195/files/figure.png" pageId="5" pageNumber="6">Figs 1B</figureCitation>
,
<figureCitation id="137147EF3151BF001CE7FF1CFE31FF1B" box="[427,483,175,201]" captionStart="Figure 3" captionStartId="3.[113,178,1836,1860]" captionTargetBox="[121,1451,145,1807]" captionTargetId="figure-6@3.[120,1452,144,1808]" captionTargetPageId="3" captionText="Figure 3. Images of epigean (Iolania perkinsi) and cavernicolous (Iolania Ƒankanstonei) Iolania species, illustrating external morphological features.A, B, Iolania perkinsi Kirkaldv habitus (A) and head and thorax (B), dorsal aspect. C, D, Iolania ssankanstonei habitus (C) and head and thorax (D), dorsal aspect. Specimen from OHS entrance (images bv B. Schurian)." figureDoi="http://doi.org/10.5281/zenodo.14423203" httpUri="https://zenodo.org/record/14423203/files/figure.png" pageId="5" pageNumber="6">3C D</figureCitation>
,
<figureCitation id="137147EF3151BF001CA1FF1CFDCAFF1B" box="[493,536,175,201]" captionStart-0="Figure 4" captionStart-1="Figure 5" captionStartId-0="5.[113,178,1954,1978]" captionStartId-1="7.[113,178,1768,1792]" captionTargetBox-0="[359,1212,820,1927]" captionTargetBox-1="[115,1456,146,1738]" captionTargetId-0="figure-313@5.[356,1216,817,1928]" captionTargetId-1="figure-6@7.[112,1459,144,1740]" captionTargetPageId-0="5" captionTargetPageId-1="7" captionText-0="Figure 4. Iolania Ƒankanstonei, male, wing morphologv. A, right tegmen. B, right hind wing.Scale bar:0.5 mm. Male from CKM entrance." captionText-1="Figure 5. Iolania Ƒankanstonei, male genitalia.A, genital segment, less lateral aspect. B, same, caudal aspect. C, anal segment, right lateral aspect.D, same, caudal aspect. E, less gonostvle, maximal aspect.F, aedeagus, dorsal aspect.G, same, less lateral aspect. Scale bars: 0.1 mm. Male from CKM entrance, same as in Figure 4." figureDoi-0="http://doi.org/10.5281/zenodo.14423205" figureDoi-1="http://doi.org/10.5281/zenodo.14423207" httpUri-0="https://zenodo.org/record/14423205/files/figure.png" httpUri-1="https://zenodo.org/record/14423207/files/figure.png" pageId="5" pageNumber="6">4–5</figureCitation>
)
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C35008E13151BF031CCCFF69FA6DFE97" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6" type="description">
<paragraph id="8BF55B6A3151BF001CCCFF69FE3EFF21" blockId="5.[384,492,218,243]" box="[384,492,218,243]" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001CCCFF69FE3EFF21" box="[384,492,218,243]" italics="true" pageId="5" pageNumber="6">Description</emphasis>
</paragraph>
<paragraph id="8BF55B6A3151BF001D3DFEB3FDDEFE66" blockId="5.[113,763,256,436]" pageId="5" pageNumber="6">
Slightlv troglomorphic species with compound eves present, but small; tegmina covering most of the abdomen, barelv attaining but not surpassing tip of abdomen; hind wings well developed but slightlv reduced in length in comparison to epigean
<taxonomicName id="4C4A20E93151BF001FFAFEEDFF6FFE47" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001FFAFEEDFD29FEA4" box="[694,763,350,374]" italics="true" pageId="5" pageNumber="6">Iolania</emphasis>
species
</taxonomicName>
; coloration generallv lighter than in epigean species, with darker portions (see ‘
<emphasis id="B93E87783151BF001C06FE2FFE60FE66" box="[330,434,412,436]" italics="true" pageId="5" pageNumber="6">Coloration</emphasis>
’ below).
</paragraph>
<paragraph id="8BF55B6A3151BF001D3DFE65FF1EFDDF" blockId="5.[113,764,469,525]" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001D3DFE65FF20FE3C" box="[113,242,470,494]" italics="true" pageId="5" pageNumber="6">Body length:</emphasis>
Male,
<quantity id="4CB2F68F3151BF001C0CFE66FE69FE3F" box="[320,443,469,493]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" metricValueMax="4.5" metricValueMin="3.5" pageId="5" pageNumber="6" unit="mm" value="4.0" valueMax="4.5" valueMin="3.5">3.5–4.5 mm</quantity>
(
<emphasis id="B93E87783151BF001C9DFE65FE36FE3F" box="[465,484,470,493]" italics="true" pageId="5" pageNumber="6">N</emphasis>
= 6); female,
<quantity id="4CB2F68F3151BF001FCDFE65FD2EFE3F" box="[641,764,470,494]" metricMagnitude="-3" metricUnit="m" metricValue="4.5" metricValueMax="4.6" metricValueMin="4.4" pageId="5" pageNumber="6" unit="mm" value="4.5" valueMax="4.6" valueMin="4.4">4.4–4.6 mm</quantity>
(
<emphasis id="B93E87783151BF001D32FE45FF43FDDF" box="[126,145,502,525]" italics="true" pageId="5" pageNumber="6">N</emphasis>
= 2).
</paragraph>
<paragraph id="8BF55B6A3151BF001D3DFD81FBA7FEF7" blockId="5.[113,763,562,742]" lastBlockId="5.[810,1459,144,293]" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001D3DFD81FF33FD98" box="[113,225,562,586]" italics="true" pageId="5" pageNumber="6">Coloration:</emphasis>
Bodv and legs vellowish. Compound eves reddish; antennae vellowish-white. Head and mesothorax laterallv sordid light brown. Tergites and sternites of abdominal segments slightlv darker brown. Tegmina translucent, light vellowish, with irregularlv defined brownish portions distallv of nodal line and faint brown transverse stripes across postcostal cell; brownish markings also accompanving median branch of Y-vein (= Pcu) and common stem of Y-vein (= Pcu * A1), in dorsal aspect creating a median ‘V’ across the tegmina. Coloration similar in both sexes; the brownish portions appear to be darker in females, creating a more contrasting pattern.
</paragraph>
<paragraph id="8BF55B6A3151BF031E66FEFCFF27FEB1" blockId="5.[810,1460,335,735]" lastBlockId="6.[129,778,144,356]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001E66FEFCFCB5FEB5" box="[810,871,335,359]" italics="true" pageId="5" pageNumber="6">Head:</emphasis>
Vertex short, ~3.6× wider posteriorlv than mediallv long, broadlv triangular, concave; posterior margin shallowlv incised mediallv, lateral margins ridged; transverse carina mediallv forming an obtuse angle. Areolets (or ‘fossette’
<emphasis id="B93E87783151BF001860FE1DFA8DFE17" box="[1324,1375,430,453]" italics="true" pageId="5" pageNumber="6">sensu</emphasis>
<bibRefCitation id="EFDB269B3151BF001826FE1EFC8CFE36" author="Giffard WM" pageId="5" pageNumber="6" pagination="1 - 121" refId="ref8694" refString="Giffard WM. A review of the Hawaiian Cixiidae with descriptions of species (Homoptera). Proceedings of the Hawaiian Entomological Society 1925; 6: 1 - 121." type="journal article" year="1925">Giffard 1925</bibRefCitation>
) indistinctlv divided mediallv bv a short obtuse carina. Frons separated from anterior margins of areolets bv an obtuse transverse carina, and with a median longitudinal carina that is distinctlv ridged and running from apical transverse carina to frontoclvpeal suture. Frons widest at base of antennae, 0.8× shorter mediallv than maximallv wide. Lateral margins of frons in ventral portion foliatelv ridged. Postclvpeus almost smooth, with a verv faint obtuse median carina in upper portion; anteclvpeus with a median faint carina; post- and anteclvpeus together mediallv ~1.6× longer than frons. Frontoclvpeal suture highlv vaulted. Compound eves small; lateral ocelli present, vestigial; median frontal ocellus absent. First antennal joint (scape) short, ring-like; second antennal joint (pedicel) cvlindrical, slightlv longer than wide, with sensorv plaque organs distinctlv recognizable. Rostrum slightlv surpassing posterior margin of hind coxae.
</paragraph>
<caption id="DF350BE23151BF001D3DF811FA42F868" ID-DOI="http://doi.org/10.5281/zenodo.14423205" ID-Zenodo-Dep="14423205" box="[113,1424,1954,1983]" httpUri="https://zenodo.org/record/14423205/files/figure.png" pageId="5" pageNumber="6" startId="5.[113,178,1954,1978]" targetBox="[359,1212,820,1927]" targetPageId="5" targetType="figure">
<paragraph id="8BF55B6A3151BF001D3DF811FA42F868" blockId="5.[113,1424,1954,1983]" box="[113,1424,1954,1983]" pageId="5" pageNumber="6">
<emphasis id="B93E87783151BF001D3DF811FE4DF86D" bold="true" box="[113,415,1954,1983]" pageId="5" pageNumber="6">
Figure 4.
<taxonomicName id="4C4A20E93151BF001D82F815FE4DF86D" authority="Fankanstonei" authorityName="Fankanstonei. B" authorityYear="2024" box="[206,415,1958,1983]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783151BF001D82F815FE4DF86D" bold="true" box="[206,415,1958,1983]" italics="true" pageId="5" pageNumber="6">Iolania frankanstonei</emphasis>
</taxonomicName>
</emphasis>
, male, wing morphologv. A, right tegmen. B, right hind wing. Scale bar: 0.5 mm. Male from CKM entrance.
</paragraph>
</caption>
<paragraph id="8BF55B6A3152BF031DCDFE3DFE20FB67" blockId="6.[128,779,398,1205]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031DCDFE3DFF1DFE77" box="[129,207,398,421]" italics="true" pageId="6" pageNumber="7">Thorax:</emphasis>
Pronotum short, ~15× wider than mediallv long, and 1.6× wider than maximum width of head including eves; posteriorlv deeplv incised, with obtuse median and two sharplv ridged lateral carinae. Mesonotum tricarinate; carinae distinctlv ridged; lateral carinae slightlv diverging posteriorlv. Mesonotum mediallv slightlv longer than maximallv wide, with lateral portions oblique. Tegulae small, without carina. Tegmina short, ~2× longer than maximal width, not surpassing tip of abdomen; in males with contracted abdomen attaining cephal margin of genital segment; in males with expanded abdomen attaining cephal margin of abdominal tergite VII; in females not surpassing tip of ovipositor; distal margin rounded, pterostigma indistinct, veins denselv covered with bases of setae. Venation with proximal branching pattern as in epigean
<taxonomicName id="4C4A20E93152BF031C9FFC96FDB9FCEF" box="[467,619,805,829]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031C9FFC96FDCAFCEF" box="[467,536,805,829]" italics="true" pageId="6" pageNumber="7">Iolania</emphasis>
species
</taxonomicName>
, but postcostal cell relativelv wider; venation distallv of nodal line varving among individuals, with variation concerning the branching pattern of posterior radius and media and that of anterior cubitus (
<figureCitation id="137147EF3152BF031DC7FC10FF16FC69" box="[139,196,931,955]" captionStart="Figure 4" captionStartId="5.[113,178,1954,1978]" captionTargetBox="[359,1212,820,1927]" captionTargetId="figure-313@5.[356,1216,817,1928]" captionTargetPageId="5" captionText="Figure 4. Iolania Ƒankanstonei, male, wing morphologv. A, right tegmen. B, right hind wing.Scale bar:0.5 mm. Male from CKM entrance." figureDoi="http://doi.org/10.5281/zenodo.14423205" httpUri="https://zenodo.org/record/14423205/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
). Claval vein (CuP) and Y-vein (Pcu, A1 and Pcu * A1) as in epigean
<taxonomicName id="4C4A20E93152BF031C49FC71FE4EFC08" box="[261,412,962,986]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B93E87783152BF031C49FC71FE98FC08" box="[261,330,962,986]" italics="true" pageId="6" pageNumber="7">Iolania</emphasis>
species.
</taxonomicName>
Hind wings well developed (
<figureCitation id="137147EF3152BF031F8CFC71FD25FC08" box="[704,759,962,986]" captionStart="Figure 4" captionStartId="5.[113,178,1954,1978]" captionTargetBox="[359,1212,820,1927]" captionTargetId="figure-313@5.[356,1216,817,1928]" captionTargetPageId="5" captionText="Figure 4. Iolania Ƒankanstonei, male, wing morphologv. A, right tegmen. B, right hind wing.Scale bar:0.5 mm. Male from CKM entrance." figureDoi="http://doi.org/10.5281/zenodo.14423205" httpUri="https://zenodo.org/record/14423205/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
); in comparison to epigean
<taxonomicName id="4C4A20E93152BF031CCBFC52FDCEFC2B" box="[391,540,993,1017]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031CCBFC52FE1EFC2B" box="[391,460,993,1017]" italics="true" pageId="6" pageNumber="7">Iolania</emphasis>
species
</taxonomicName>
, relativelv shorter. Legs moderatelv long; metatibiae laterallv with three minute spines, and five to seven strong apical spines, bilaterallv and individuallv variable (5/6; 5/7; 6/6). Metabasitarsus elongate, approximatelv as long as second and third tarsomere together, with five apical spines. Second tarsomere with four or five apical spines. Pretarsal claws small, arolia pad-like.
</paragraph>
<paragraph id="8BF55B6A3152BF031DCCFB64FDD9F8F0" blockId="6.[128,778,1239,1826]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031DCCFB64FEC6FB3D" box="[128,276,1239,1263]" italics="true" pageId="6" pageNumber="7">Male genitalia:</emphasis>
Genital segment in caudal aspect slightlv higher than wide, laterodorsal margin rounded, medioventral process triangular. Anal segment distallv of anal stvle on both sides broadlv rounded, ventrocaudal margin mediallv straight; anal segment in dorsal aspect with lateral margins more or less parallel, in lateral aspect straight, distallv of anal stvle not bent ventrallv. Gonostvli proximallv slender, ventrallv ridged, ridge finelv striate, distal portion slightlv dilated, slender, produced dorsallv, caudal margin rounded, dorsal margin slightlv concave. Aedeagus bilaterallv svmmetrical; shaft with enhanced sclerotization in basal half, in lateral aspect distinctlv narrowing in apical third (
<figureCitation id="137147EF3152BF031C6EF99CFEBCF995" box="[290,366,1583,1607]" captionStart="Figure 5" captionStartId="7.[113,178,1768,1792]" captionTargetBox="[115,1456,146,1738]" captionTargetId="figure-6@7.[112,1459,144,1740]" captionTargetPageId="7" captionText="Figure 5. Iolania Ƒankanstonei, male genitalia.A, genital segment, less lateral aspect. B, same, caudal aspect. C, anal segment, right lateral aspect.D, same, caudal aspect. E, less gonostvle, maximal aspect.F, aedeagus, dorsal aspect.G, same, less lateral aspect. Scale bars: 0.1 mm. Male from CKM entrance, same as in Figure 4." figureDoi="http://doi.org/10.5281/zenodo.14423207" httpUri="https://zenodo.org/record/14423207/files/figure.png" pageId="6" pageNumber="7">Fig. 5G</figureCitation>
, arrow); shaft ventrallv in basal portion finelv rugose; flagellum short, with surface distinctlv sculptured, dorsal surface mediallv concave, with conspicuous groove distallv of phallotreme; with two movable spinose processes near apex, processes arm-like, proximallv taeniform, distallv terete and tapering in an acute tip, in repose directed mediobasallv, not attaining basal (proximal) third of shaft. Gonoduct with conspicuous protuberance at apex of shaft.
</paragraph>
<paragraph id="8BF55B6A3152BF031DCCF8F7FC70FF15" blockId="6.[128,778,1860,1978]" lastBlockId="6.[825,1474,144,199]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031DCCF8F7FEF9F88E" box="[128,299,1860,1884]" italics="true" pageId="6" pageNumber="7">Female genitalia:</emphasis>
External structures of female genitalia as in epigean
<taxonomicName id="4C4A20E93152BF031D95F8D0FEA7F8A9" box="[217,373,1891,1915]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031D95F8D0FECCF8A9" box="[217,286,1891,1915]" italics="true" pageId="6" pageNumber="7">Iolania</emphasis>
species
</taxonomicName>
: ovipositor ensiform, caudallv slightlv surpassing anal segment. Tergite IX caudallv truncate, verv slightlv concave, with wax-secreting area inconspicuouslv delimited. Anal segment slender, ventral margin caudallv slightlv produced.
</paragraph>
<paragraph id="8BF55B6A3152BF031943FF54FB3EFED3" blockId="6.[825,1474,231,325]" box="[1039,1260,231,257]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031943FF54FB3EFED3" box="[1039,1260,231,257]" italics="true" pageId="6" pageNumber="7">Molecular identification</emphasis>
</paragraph>
<paragraph id="8BF55B6A3152BF031E75FEBEFA6DFE97" blockId="6.[825,1474,231,325]" pageId="6" pageNumber="7">
Mitochondrial
<emphasis id="B93E87783152BF031E9FFEBDFC2DFEF7" box="[979,1023,270,293]" italics="true" pageId="6" pageNumber="7">COI</emphasis>
sequences of
<taxonomicName id="4C4A20E93152BF0319C0FEBDFAF0FEF7" box="[1164,1314,269,293]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E87783152BF0319C0FEBDFAF0FEF7" box="[1164,1314,269,293]" italics="true" pageId="6" pageNumber="7">I. frankanstonei</emphasis>
</taxonomicName>
were deposited in GenBank under accession numbers MZ048276–MZ048281.
</paragraph>
</subSubSection>
<subSubSection id="C35008E13152BF03197CFED6FC44FC4A" pageId="6" pageNumber="7" type="diagnosis">
<paragraph id="8BF55B6A3152BF03197CFED6FB1EFEAC" blockId="6.[825,1475,356,920]" box="[1072,1228,356,383]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF03197CFED6FB1EFEAC" box="[1072,1228,356,383]" italics="true" pageId="6" pageNumber="7">Species diagnosis</emphasis>
</paragraph>
<paragraph id="8BF55B6A3152BF031E75FE38FC44FC4A" blockId="6.[825,1475,356,920]" pageId="6" pageNumber="7">
<taxonomicName id="4C4A20E93152BF031E75FE38FBD8FE71" box="[825,1034,395,419]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E87783152BF031E75FE38FBD8FE71" box="[825,1034,395,419]" italics="true" pageId="6" pageNumber="7">Iolania frankanstonei</emphasis>
</taxonomicName>
differs from all epigean
<taxonomicName id="4C4A20E93152BF031843FE38FA77FE71" box="[1295,1445,395,419]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031843FE38FA86FE71" box="[1295,1364,395,419]" italics="true" pageId="6" pageNumber="7">Iolania</emphasis>
species
</taxonomicName>
in external characters related to cave adaptation (reduction of compound eves, configuration of head carination and proportions, length and venation of tegmina and wings, pigmentation pattern; see above). Concerning the male genital structures,
<taxonomicName id="4C4A20E93152BF0318F8FDBAFC6EFDED" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E87783152BF0318F8FDBAFC6EFDED" italics="true" pageId="6" pageNumber="7">I. frankanstonei</emphasis>
</taxonomicName>
is most similar to
<taxonomicName id="4C4A20E93152BF03193FFD9BFB00FDED" authorityName="Kirkaldv" authorityYear="1902" box="[1139,1234,551,575]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF03193FFD9BFB00FDED" box="[1139,1234,551,575]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
, but differs in the shape of the anal segment (distal portion straight vs. bent ventrallv in
<taxonomicName id="4C4A20E93152BF0318F8FDF4FC57FDAC" authorityName="Kirkaldv" authorityYear="1902" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF0318F8FDF4FC57FDAC" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
) and of the aedeagus (shaft with enhanced sclerotization in basal half vs. in basal third in
<taxonomicName id="4C4A20E93152BF031932FD35FB0CFD4F" authorityName="Kirkaldv" authorityYear="1902" box="[1150,1246,645,669]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF031932FD35FB0CFD4F" box="[1150,1246,645,669]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
; enhanced sclerotized part ventrallv rugose vs. smooth in
<taxonomicName id="4C4A20E93152BF0319EEFD16FAD0FD6F" authorityName="Kirkaldv" authorityYear="1902" box="[1186,1282,677,701]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF0319EEFD16FAD0FD6F" box="[1186,1282,677,701]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
; shaft in lateral aspect distinctlv narrowing in apical third vs. straight in
<taxonomicName id="4C4A20E93152BF031817FD76FA69FD0E" authorityName="Kirkaldv" authorityYear="1902" box="[1371,1467,708,732]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF031817FD76FA69FD0E" box="[1371,1467,708,732]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
; arm-like paired processes at transition of shaft and flagellum taeniform throughout vs. distallv terete in
<taxonomicName id="4C4A20E93152BF0319B8FCB0FA84FCC8" authorityName="Kirkaldv" authorityYear="1902" box="[1268,1366,770,794]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF0319B8FCB0FA84FCC8" box="[1268,1366,770,794]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
, and their tips in repose directed straight basomediallv vs. pointed dorsallv in
<taxonomicName id="4C4A20E93152BF031E18FCF1FC67FC8B" authorityName="Kirkaldv" authorityYear="1902" box="[852,949,833,857]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF031E18FCF1FC67FC8B" box="[852,949,833,857]" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
; dorsal surface of flagellum with phallotreme wide but without longitudinal conspicuous groove distal of it as in
<taxonomicName id="4C4A20E93152BF0318F8FCD2FC57FC4A" authorityName="Kirkaldv" authorityYear="1902" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E87783152BF0318F8FCD2FC57FC4A" italics="true" pageId="6" pageNumber="7">I. perkinsi</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="C35008E13152BF03191FFC04FA63FA5F" pageId="6" pageNumber="7" type="discussion">
<paragraph id="8BF55B6A3152BF03191FFC04FB75FC03" blockId="6.[1107,1191,951,977]" box="[1107,1191,951,977]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF03191FFC04FB75FC03" box="[1107,1191,951,977]" italics="true" pageId="6" pageNumber="7">Remarks</emphasis>
</paragraph>
<paragraph id="8BF55B6A3152BF031E75FC6DFA63FA5F" blockId="6.[825,1474,990,1421]" pageId="6" pageNumber="7">
The description of
<taxonomicName id="4C4A20E93152BF031EB5FC6CFB5DFC24" box="[1017,1167,990,1014]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E87783152BF031EB5FC6CFB5DFC24" box="[1017,1167,990,1014]" italics="true" pageId="6" pageNumber="7">I. frankanstonei</emphasis>
</taxonomicName>
is based on comparativelv few specimens (
<specimenCount id="9D4C90E33152BF031EFFFC4EFBC6FBC7" box="[947,1044,1021,1045]" count="6" pageId="6" pageNumber="7" type="male">six males</specimenCount>
and
<specimenCount id="9D4C90E33152BF031905FC4DFB19FBC7" box="[1097,1227,1021,1045]" count="2" pageId="6" pageNumber="7" type="female">two females</specimenCount>
). The range of variation within and between populations from different passages of the Kipuka Kanohina lava tube svstem can therefore not be assessed fullv. Variation exists, however slight, in the size of the compound eves, relative length and venation of the tegmen, and male genital characters (e.g. the shape of gonostvli) within and among specimens from different populations; therefore, it cannot be excluded that the specimens here, hvpothesized to present a single species, might belong to separate reproductive units. We therefore decided to assign the holotvpe and paratvpes from a single population and chose the original collection location, the CKM entrance, as the tvpe localitv, because it was the first location in the Kipuka Kanohina svstem where specimens were collected.
</paragraph>
</subSubSection>
<subSubSection id="C35008E13152BF0D1968FA1EFE8CFDED" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7" type="materials_examined">
<paragraph id="8BF55B6A3152BF031968FA1EFB05FA15" blockId="6.[1060,1239,1453,1479]" box="[1060,1239,1453,1479]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031968FA1EFB05FA15" box="[1060,1239,1453,1479]" italics="true" pageId="6" pageNumber="7">Material examined</emphasis>
</paragraph>
<paragraph id="8BF55B6A3152BF031E75FA67FAFCF998" blockId="6.[825,1474,1492,1610]" pageId="6" pageNumber="7">
<materialsCitation id="3B2251373152BF031E75FA67FAFCF998" collectingDate="2016-11-24" collectionCode="BPBM" collectorName="M. E. Slav &amp; C. A. M. Slav &amp; M. L. Porter" country="United States of America" elevation="1096" location="Male" municipality="Kipuka Kanohina" pageId="6" pageNumber="7" specimenCount="1" stateProvince="Hawaii" typeStatus="holotype">
<emphasis id="B93E87783152BF031E75FA67FC4BFA3E" box="[825,921,1492,1516]" italics="true" pageId="6" pageNumber="7">
<typeStatus id="54F1E5C83152BF031E75FA67FC46FA3E" box="[825,916,1492,1516]" pageId="6" pageNumber="7" type="holotype">Holotype</typeStatus>
:
</emphasis>
<location id="8E950DB13152BF031ED2FA67FC01FA3E" LSID="urn:lsid:plazi:treatment:03E3EA7C3151BF0D1D8FFF3DFB23FDEA:8E950DB13152BF031ED2FA67FC01FA3E" box="[926,979,1492,1516]" country="United States of America" municipality="Kipuka Kanohina" name="Male" pageId="6" pageNumber="7" stateProvince="Hawaii">Male</location>
,
<collectingRegion id="498E95883152BF031E97FA67FBFCFA3E" box="[987,1070,1492,1516]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
/
<collectingCountry id="F35D1BFA3152BF03197AFA67FBBEFA3E" box="[1078,1132,1492,1516]" name="United States of America" pageId="6" pageNumber="7">USA</collectingCountry>
:
<collectingRegion id="498E95883152BF031938FA67FB11FA3E" box="[1140,1219,1492,1516]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
Island,
<collectingMunicipality id="6B91C1103152BF031843FA67FA10FA3E" box="[1295,1474,1492,1516]" pageId="6" pageNumber="7">Kipuka Kanohina</collectingMunicipality>
lava tube svstem, CKM entrance (HI 00153),
<elevation id="0067BC593152BF031855FA40FA4EF9D9" box="[1305,1436,1523,1547]" metricMagnitude="3" metricUnit="m" metricValue="1.0959999999999999" pageId="6" pageNumber="7" unit="m" value="1096.0">
<quantity id="4CB2F68F3152BF031855FA40FABBF9D9" box="[1305,1385,1523,1547]" metricMagnitude="3" metricUnit="m" metricValue="1.0959999999999999" pageId="6" pageNumber="7" unit="m" value="1096.0">1096 m</quantity>
a.s.l.
</elevation>
,
<date id="FFF47DAA3152BF0318E4FA40FC36F9F9" pageId="6" pageNumber="7" value="2016-11-24">
<collectingDate id="EFB084423152BF0318E4FA40FC36F9F9" pageId="6" pageNumber="7" value="2016-11-24">24 November 2016</collectingDate>
</date>
,
<collectorName id="26BF3EBC3152BF031EA2F9A0FB83F9F9" box="[1006,1105,1555,1579]" pageId="6" pageNumber="7">M.E. Slav</collectorName>
,
<collectorName id="26BF3EBC3152BF031917F9A0FB08F9F9" box="[1115,1242,1555,1579]" pageId="6" pageNumber="7">C.A.M. Slav</collectorName>
, and
<collectorName id="26BF3EBC3152BF03185DF9A0FA59F9F8" box="[1297,1419,1555,1578]" pageId="6" pageNumber="7">M.L. Porter</collectorName>
coll.,
<collectionCode id="ED5BC3AF3152BF031E75F981FCACF99B" box="[825,894,1586,1609]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34697" lsid="urn:lsid:biocol.org:col:34697" name="Bishop Museum" pageId="6" pageNumber="7" type="Museum">BPBM</collectionCode>
(tvpe number:
<emphasis id="B93E87783152BF031953F980FACFF998" box="[1055,1309,1586,1610]" italics="true" pageId="6" pageNumber="7">BPBMENT 2008035007</emphasis>
).
</materialsCitation>
</paragraph>
<paragraph id="8BF55B6A3152BF031E75F9DFFA87F951" blockId="6.[825,1365,1643,1667]" box="[825,1365,1643,1667]" pageId="6" pageNumber="7">
<materialsCitation id="3B2251373152BF031E75F9DFFA83F951" box="[825,1361,1643,1667]" collectionCode="BPBM" pageId="6" pageNumber="7" specimenCount="2" specimenCount-male="2" typeStatus="paratype">
<emphasis id="B93E87783152BF031E75F9DFFC72F951" box="[825,928,1644,1667]" italics="true" pageId="6" pageNumber="7">
<typeStatus id="54F1E5C83152BF031E75F9DFFC49F951" box="[825,923,1644,1667]" pageId="6" pageNumber="7" type="paratype">Paratypes</typeStatus>
:
</emphasis>
<specimenCount id="9D4C90E33152BF031EE9F9DFFBC7F951" box="[933,1045,1643,1667]" count="2" pageId="6" pageNumber="7" type="male">Two males</specimenCount>
, same data as holotvpe,
<collectionCode id="ED5BC3AF3152BF031845F9DFFA83F951" box="[1289,1361,1644,1667]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34697" lsid="urn:lsid:biocol.org:col:34697" name="Bishop Museum" pageId="6" pageNumber="7" type="Museum">BPBM</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BF55B6A3152BF031E75F916FC57F8EB" blockId="6.[825,1475,1701,1975]" pageId="6" pageNumber="7">
<emphasis id="B93E87783152BF031E75F916FBDBF96F" box="[825,1033,1701,1725]" italics="true" pageId="6" pageNumber="7">Additional material:</emphasis>
<materialsCitation id="3B2251373152BF03195BF916FC53F8EB" collectingDate="2018-11-24" collectionCode="BPBM" collectorName="M. E. Slav &amp; C. A. M. Slav &amp; A. Katz &amp; J. Gunter &amp; K. Yelverton &amp; P. Bosted" country="United States of America" elevation="730" location="Kipuka Kanohina" pageId="6" pageNumber="7" specimenCount="3" specimenCount-female="2" specimenCount-male="1" stateProvince="Hawaii">
<specimenCount id="9D4C90E33152BF03195BF916FB54F96F" box="[1047,1158,1701,1725]" count="1" pageId="6" pageNumber="7" type="male">One male</specimenCount>
,
<specimenCount id="9D4C90E33152BF0319DBF916FAC9F96F" box="[1175,1307,1701,1725]" count="2" pageId="6" pageNumber="7" type="female">two females</specimenCount>
,
<collectingRegion id="498E95883152BF031861F916FA52F96F" box="[1325,1408,1701,1725]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
/
<collectingCountry id="F35D1BFA3152BF0318C5F916FA6DF96F" box="[1417,1471,1701,1725]" name="United States of America" pageId="6" pageNumber="7">USA</collectingCountry>
:
<collectingRegion id="498E95883152BF031E75F977FC5AF90E" box="[825,904,1732,1756]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
Island,
<location id="8E950DB13152BF031EACF977FB4BF90E" LSID="urn:lsid:plazi:treatment:03E3EA7C3151BF0D1D8FFF3DFB23FDEA:8E950DB13152BF031EACF977FB4BF90E" box="[992,1177,1732,1756]" country="United States of America" name="Kipuka Kanohina" pageId="6" pageNumber="7" stateProvince="Hawaii">Kipuka Kanohina</location>
lava tube svstem, OHS entrance (HI 00809),
<elevation id="0067BC593152BF03194BF950FBAEF929" box="[1031,1148,1763,1787]" metricMagnitude="2" metricUnit="m" metricValue="7.3" pageId="6" pageNumber="7" unit="m" value="730.0">
<quantity id="4CB2F68F3152BF03194BF950FB9BF929" box="[1031,1097,1763,1787]" metricMagnitude="2" metricUnit="m" metricValue="7.3" pageId="6" pageNumber="7" unit="m" value="730.0">730 m</quantity>
a.s.l.
</elevation>
,
<date id="FFF47DAA3152BF0319CBF950FA80F929" box="[1159,1362,1763,1787]" pageId="6" pageNumber="7" value="2018-11-24">
<collectingDate id="EFB084423152BF0319CBF950FA80F929" box="[1159,1362,1763,1787]" pageId="6" pageNumber="7" value="2018-11-24">24 November 2018</collectingDate>
</date>
,
<collectorName id="26BF3EBC3152BF031810F957FA6DF929" box="[1372,1471,1763,1787]" pageId="6" pageNumber="7">M.E. Slav</collectorName>
,
<collectorName id="26BF3EBC3152BF031E77F8B0FC65F8C9" box="[827,951,1795,1819]" pageId="6" pageNumber="7">C.A.M. Slav</collectorName>
,
<collectorName id="26BF3EBC3152BF031EF3F8B0FBDFF8C8" box="[959,1037,1795,1819]" pageId="6" pageNumber="7">A. Katz</collectorName>
,
<collectorName id="26BF3EBC3152BF03195AF8B0FBA1F8C9" box="[1046,1139,1795,1819]" pageId="6" pageNumber="7">J. Gunter</collectorName>
,
<collectorName id="26BF3EBC3152BF031930F8B0FB2EF8C9" box="[1148,1276,1795,1819]" pageId="6" pageNumber="7">K. Yelverton</collectorName>
, and
<collectorName id="26BF3EBC3152BF03187CF8B0FA5CF8C9" box="[1328,1422,1795,1819]" pageId="6" pageNumber="7">P. Bosted</collectorName>
coll.,
<collectionCode id="ED5BC3AF3152BF031E75F891FC53F8EB" box="[825,897,1826,1849]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34697" lsid="urn:lsid:biocol.org:col:34697" name="Bishop Museum" pageId="6" pageNumber="7" type="Museum">BPBM</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BF55B6A3152BF031E19F8F2FBCDF865" blockId="6.[825,1475,1701,1975]" pageId="6" pageNumber="7">
<materialsCitation id="3B2251373152BF031E19F8F2FAC9F845" collectingDate="2018-03-25" collectionCode="V" collectorName="M. E. Slav &amp; A. S. Engel &amp; S. Engel" country="United States of America" elevation="539" location="Kipuka Kanohina" pageId="6" pageNumber="7" specimenCount="1" specimenCount-male="1" stateProvince="Hawaii">
<specimenCount id="9D4C90E33152BF031E19F8F2FC6FF88B" box="[853,957,1857,1881]" count="1" pageId="6" pageNumber="7" type="male">One male</specimenCount>
,
<collectingRegion id="498E95883152BF031E85F8F2FBCEF88B" box="[969,1052,1857,1881]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
/
<collectingCountry id="F35D1BFA3152BF031969F8F2FB89F88B" box="[1061,1115,1857,1881]" name="United States of America" pageId="6" pageNumber="7">USA</collectingCountry>
:
<collectingRegion id="498E95883152BF03192BF8F2FB64F88B" box="[1127,1206,1857,1881]" country="United States of America" name="Hawaii" pageId="6" pageNumber="7">Hawaiʻi</collectingRegion>
Island,
<location id="8E950DB13152BF031847F8F2FA10F88B" LSID="urn:lsid:plazi:treatment:03E3EA7C3151BF0D1D8FFF3DFB23FDEA:8E950DB13152BF031847F8F2FA10F88B" box="[1291,1474,1857,1881]" country="United States of America" name="Kipuka Kanohina" pageId="6" pageNumber="7" stateProvince="Hawaii">Kipuka Kanohina</location>
lava tube svstem, HLC entrance (HI 00323),
<elevation id="0067BC593152BF03186DF8D3FA49F8AB" box="[1313,1435,1888,1913]" metricMagnitude="2" metricUnit="m" metricValue="5.39" pageId="6" pageNumber="7" unit="m" value="539.0">
<quantity id="4CB2F68F3152BF03186DF8D3FAB7F8AA" box="[1313,1381,1888,1912]" metricMagnitude="2" metricUnit="m" metricValue="5.39" pageId="6" pageNumber="7" unit="m" value="539.0">539 m</quantity>
a.s.l.
</elevation>
,
<date id="FFF47DAA3152BF0318E4F8D3FC68F84A" pageId="6" pageNumber="7" value="2018-03-25">
<collectingDate id="EFB084423152BF0318E4F8D3FC68F84A" pageId="6" pageNumber="7" value="2018-03-25">25 March 2018</collectingDate>
</date>
,
<collectorName id="26BF3EBC3152BF031E88F833FBFAF84A" box="[964,1064,1920,1944]" pageId="6" pageNumber="7">M.E. Slav</collectorName>
,
<collectorName id="26BF3EBC3152BF03197EF833FB4DF84A" box="[1074,1183,1920,1944]" pageId="6" pageNumber="7">A.S. Engel</collectorName>
,
<collectorName id="26BF3EBC3152BF0319E6F833FB2DF84A" box="[1194,1279,1920,1944]" pageId="6" pageNumber="7">S. Engel</collectorName>
,
<collectionCode id="ED5BC3AF3152BF031846F833FAC9F845" box="[1290,1307,1920,1943]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="6" pageNumber="7" type="Museum">V</collectionCode>
</materialsCitation>
.
<materialsCitation id="3B2251373152BF031869F833FBC9F865" collectionCode="T, BPBM" collectorName="Gracanin" country="United States of America" location="Hackell" pageId="6" pageNumber="7" specimenCount="1">
<location id="8E950DB13152BF031869F833FAA7F84A" LSID="urn:lsid:plazi:treatment:03E3EA7C3151BF0D1D8FFF3DFB23FDEA:8E950DB13152BF031869F833FAA7F84A" box="[1317,1397,1920,1944]" country="United States of America" name="Hackell" pageId="6" pageNumber="7">Hackell</location>
, and
<collectionCode id="ED5BC3AF3152BF0318E2F833FA6DF845" box="[1454,1471,1920,1943]" name="Tavera, Department of Geology and Geophysics" pageId="6" pageNumber="7">T</collectionCode>
.
<collectorName id="26BF3EBC3152BF031E75F82CFC4AF865" box="[825,920,1951,1975]" pageId="6" pageNumber="7">Gracanin</collectorName>
coll.,
<collectionCode id="ED5BC3AF3152BF031E9FF813FBC9F865" box="[979,1051,1952,1975]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34697" lsid="urn:lsid:biocol.org:col:34697" name="Bishop Museum" pageId="6" pageNumber="7" type="Museum">BPBM</collectionCode>
</materialsCitation>
.
</paragraph>
<caption id="DF350BE23153BF021D3DF95BFE20F8E5" ID-DOI="http://doi.org/10.5281/zenodo.14423207" ID-Zenodo-Dep="14423207" httpUri="https://zenodo.org/record/14423207/files/figure.png" pageId="7" pageNumber="8" startId="7.[113,178,1768,1792]" targetBox="[115,1456,146,1738]" targetPageId="7" targetType="figure">
<paragraph id="8BF55B6A3153BF021D3DF95BFE20F8E5" blockId="7.[113,1458,1768,1848]" pageId="7" pageNumber="8">
<emphasis id="B93E87783153BF021D3DF95BFE4CF8D6" bold="true" box="[113,414,1768,1796]" pageId="7" pageNumber="8">
Figure 5.
<taxonomicName id="4C4A20E93153BF021D81F958FE4CF8D6" authority="Fankanstonei" authorityName="Fankanstonei. B" authorityYear="2024" box="[205,414,1771,1796]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis id="B93E87783153BF021D81F958FE4CF8D6" bold="true" box="[205,414,1771,1796]" italics="true" pageId="7" pageNumber="8">Iolania frankanstonei</emphasis>
</taxonomicName>
</emphasis>
, male genitalia. A, genital segment, left lateral aspect. B, same, caudal aspect. C, anal segment, right lateral aspect. D, same, caudal aspect. E, left gonostvle, maximal aspect. F, aedeagus, dorsal aspect. G, same, left lateral aspect. Scale bars: 0.1 mm. Male from CKM entrance, same as in Figure 4.
</paragraph>
</caption>
<paragraph id="8BF55B6A315CBF0D1DD0FF23FF1FFED7" blockId="8.[129,778,144,575]" pageId="8" pageNumber="9">
<materialsCitation id="3B225137315CBF0D1DD0FF23FF1BFED7" collectingDate="2018-11-19" collectionCode="R, BPBM" collectorName="A. Chong &amp; A. S. Engel &amp; M. L. Porter" country="United States of America" elevation="539" location="Kipuka Kanohina" pageId="8" pageNumber="9" specimenCount="1" specimenCount-male="1" stateProvince="Hawaii">
<specimenCount id="9D4C90E3315CBF0D1DD0FF23FED7FF7A" box="[156,261,144,168]" count="1" pageId="8" pageNumber="9" type="male">One male</specimenCount>
,
<collectingRegion id="498E9588315CBF0D1C5CFF23FEB1FF7A" box="[272,355,144,168]" country="United States of America" name="Hawaii" pageId="8" pageNumber="9">Hawaiʻi</collectingRegion>
/
<collectingCountry id="F35D1BFA315CBF0D1C20FF23FE70FF7A" box="[364,418,144,168]" name="United States of America" pageId="8" pageNumber="9">USA</collectingCountry>
:
<collectingRegion id="498E9588315CBF0D1CE2FF23FE2FFF7A" box="[430,509,144,168]" country="United States of America" name="Hawaii" pageId="8" pageNumber="9">Hawaiʻi</collectingRegion>
Island,
<location id="8E950DB1315CBF0D1F1EFF23FCDBFF7A" LSID="urn:lsid:plazi:treatment:03E3EA7C3151BF0D1D8FFF3DFB23FDEA:8E950DB1315CBF0D1F1EFF23FCDBFF7A" box="[594,777,144,168]" country="United States of America" name="Kipuka Kanohina" pageId="8" pageNumber="9" stateProvince="Hawaii">Kipuka Kanohina</location>
lava tube svstem, HLC entrance (HI 00524),
<elevation id="0067BC59315CBF0D1F24FF1CFD30FF15" box="[616,738,175,199]" metricMagnitude="2" metricUnit="m" metricValue="5.39" pageId="8" pageNumber="9" unit="m" value="539.0">
<quantity id="4CB2F68F315CBF0D1F24FF1CFD7FFF15" box="[616,685,175,199]" metricMagnitude="2" metricUnit="m" metricValue="5.39" pageId="8" pageNumber="9" unit="m" value="539.0">539 m</quantity>
a.s.l.
</elevation>
,
<date id="FFF47DAA315CBF0D1FA3FF1CFEFBFF35" pageId="8" pageNumber="9" value="2018-11-19">
<collectingDate id="EFB08442315CBF0D1FA3FF1CFEFBFF35" pageId="8" pageNumber="9" value="2018-11-19">19 November 2018</collectingDate>
</date>
,
<collectionCode id="ED5BC3AF315CBF0D1C7FFF7CFE95FF35" box="[307,327,207,231]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="8" pageNumber="9">R</collectionCode>
.
<collectorName id="26BF3EBC315CBF0D1C06FF7CFE60FF35" box="[330,434,207,231]" pageId="8" pageNumber="9">A. Chong</collectorName>
,
<collectorName id="26BF3EBC315CBF0D1CF7FF7CFDF4FF35" box="[443,550,207,231]" pageId="8" pageNumber="9">A.S. Engel</collectorName>
, and
<collectorName id="26BF3EBC315CBF0D1F17FF7CFD06FF34" box="[603,724,207,230]" pageId="8" pageNumber="9">M.L. Porter</collectorName>
coll.,
<collectionCode id="ED5BC3AF315CBF0D1DCDFF5DFF1BFED7" box="[129,201,238,261]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34697" lsid="urn:lsid:biocol.org:col:34697" name="Bishop Museum" pageId="8" pageNumber="9" type="Museum">BPBM</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BF55B6A315CBF0D1DD0FEBEFE8CFDED" blockId="8.[129,778,144,575]" pageId="8" pageNumber="9">
Cixiid nvmphs belonging to
<taxonomicName id="4C4A20E9315CBF0D1C8FFEBEFDDAFEF7" authorityName="Fankanstonei. B" authorityYear="2024" box="[451,520,269,293]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="B93E8778315CBF0D1C8FFEBEFDDAFEF7" box="[451,520,269,293]" italics="true" pageId="8" pageNumber="9">Iolania</emphasis>
</taxonomicName>
were also collected from cave sections within the Kipuka Kanohina lava tube svstem in close proximitv to adult specimens used for this description. However, nvmphs of epigean
<taxonomicName id="4C4A20E9315CBF0D1C8BFED8FDDEFE51" authorityName="Fankanstonei. B" authorityYear="2024" box="[455,524,363,387]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="B93E8778315CBF0D1C8BFED8FDDEFE51" box="[455,524,363,387]" italics="true" pageId="8" pageNumber="9">Iolania</emphasis>
</taxonomicName>
are ‘hitherto unknown – thev most likelv live (like the nvmphs of all cixiids) close to or inside the soil, and are not documented in collections’ (Hoch 2006: 317). Based on morphologv alone, we therefore cannot assign the nvmphs with certaintv to either
<taxonomicName id="4C4A20E9315CBF0D1FC7FE5AFD3EFDD3" authorityName="Kirkaldv" authorityYear="1902" box="[651,748,489,513]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E8778315CBF0D1FC7FE5AFD3EFDD3" box="[651,748,489,513]" italics="true" pageId="8" pageNumber="9">I. perkinsi</emphasis>
</taxonomicName>
or
<taxonomicName id="4C4A20E9315CBF0D1DCDFDBAFECBFDF2" box="[129,281,520,544]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D1DCDFDBAFECBFDF2" box="[129,281,520,544]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
, hence we have opted to refrain from accessing these specimens here.
</paragraph>
</subSubSection>
<subSubSection id="C35008E1315CBF0D1CDEFDECFCD8FC24" pageId="8" pageNumber="9" type="etymology">
<paragraph id="8BF55B6A315CBF0D1CDEFDECFE2AFDAB" blockId="8.[129,778,607,1014]" box="[402,504,607,633]" pageId="8" pageNumber="9">
<emphasis id="B93E8778315CBF0D1CDEFDECFE2AFDAB" box="[402,504,607,633]" italics="true" pageId="8" pageNumber="9">Etymology</emphasis>
</paragraph>
<paragraph id="8BF55B6A315CBF0D1DCDFD35FCD8FC24" blockId="8.[129,778,607,1014]" pageId="8" pageNumber="9">The species epithet is chosen to honour the friendship and collaboration between the pioneers of Hawaiian cave biologv, the late Fred D. Stone (1938–2018) and Frank G. Howarth, Bishop Museum. Over the course of nearlv six decades, thev explored hundreds of caves, discovered dozens of new species, and inspired and supported countless people (students in addition to novice and senior scientists). Known to fellow cavers as ‘Frank ’n Stone’, thev are held in high esteem as reliable buddies in the field and valued discussion partners. Their scientific expertise, combined with personal integritv and their notorious sense of (occasionallv dark) humour, renders them unforgettable to anvone who has had the good fortune to meet and spend time underground with them.</paragraph>
</subSubSection>
<subSubSection id="C35008E1315CBF0D1CC7FBA6FE1BF95A" pageId="8" pageNumber="9" type="distribution">
<paragraph id="8BF55B6A315CBF0D1CC7FBA6FE2CFBFD" blockId="8.[395,510,1045,1071]" box="[395,510,1045,1071]" pageId="8" pageNumber="9">
<emphasis id="B93E8778315CBF0D1CC7FBA6FE2CFBFD" box="[395,510,1045,1071]" italics="true" pageId="8" pageNumber="9">Distribution</emphasis>
</paragraph>
<paragraph id="8BF55B6A315CBF0D1DCDFB8FFE1BF95A" blockId="8.[129,778,1084,1672]" pageId="8" pageNumber="9">
Specimens of
<taxonomicName id="4C4A20E9315CBF0D1C53FB8EFE6CFB86" box="[287,446,1084,1108]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D1C53FB8EFE6CFB86" box="[287,446,1084,1108]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
have been documented from multiple lava tubes across a range of elevations in the Kipuka Kanohina Cave Svstem (
<bibRefCitation id="EFDB269B315CBF0D1CC5FBC8FDDCFB41" author="Bosted P" box="[393,526,1147,1171]" pageId="8" pageNumber="9" pagination="4 - 11" refId="ref8365" refString="Bosted P. Update on the Kipuka Kanohina cave svstem in Hawai'i. NSS News 2017; 75: 4 - 11." type="journal article" year="2017">Bosted 2017</bibRefCitation>
), on the south-western side of Mauna Loa volcano in 750- to 1500-vear-old lava flows. It is a highlv braided and mazv svstem, with multiple levels and interconnected passages, located in the desert of Kaʻu near the communitv of Ocean View, ~
<quantity id="4CB2F68F315CBF0D1CFEFB4BFE24FAC2" box="[434,502,1272,1296]" metricMagnitude="4" metricUnit="m" metricValue="7.2" pageId="8" pageNumber="9" unit="km" value="72.0">72 km</quantity>
south of Kailua-Kona and ~
<quantity id="4CB2F68F315CBF0D1DC2FAA4FF01FAFD" box="[142,211,1303,1327]" metricMagnitude="4" metricUnit="m" metricValue="4.8" pageId="8" pageNumber="9" unit="km" value="48.0">48 km</quantity>
north of South Point (the southernmost point in the
<collectingCountry id="F35D1BFA315CBF0D1DCDFA84FF6AFA9D" box="[129,184,1335,1359]" name="United States of America" pageId="8" pageNumber="9">USA</collectingCountry>
). So far,&gt;
<quantity id="4CB2F68F315CBF0D1C57FA84FE89FA9D" box="[283,347,1335,1359]" metricMagnitude="4" metricUnit="m" metricValue="2.9" pageId="8" pageNumber="9" unit="km" value="29.0">29 km</quantity>
of continuous cave passages have been explored within the svstem, with&gt;
<quantity id="4CB2F68F315CBF0D1C98FAE5FDC5FABC" box="[468,535,1366,1390]" metricMagnitude="4" metricUnit="m" metricValue="4.2" pageId="8" pageNumber="9" unit="km" value="42.0">42 km</quantity>
in the entire svstem. It has a vertical extent of&gt;
<quantity id="4CB2F68F315CBF0D1C3DFAC6FE12FA5F" box="[369,448,1397,1421]" metricMagnitude="3" metricUnit="m" metricValue="1.3" pageId="8" pageNumber="9" unit="m" value="1300.0">1300 m</quantity>
, with passages at almost sea level up to&gt;
<quantity id="4CB2F68F315CBF0D1D83FA26FEF3FA7E" box="[207,289,1429,1453]" metricMagnitude="3" metricUnit="m" metricValue="1.4" pageId="8" pageNumber="9" unit="m" value="1400.0">1400 m</quantity>
a.s.l. The cave svstem is currentlv listed as the third longest lava tube in the world, one of onlv three to surpass
<quantity id="4CB2F68F315CBF0D1DCDFA60FF16FA39" box="[129,196,1491,1515]" metricMagnitude="4" metricUnit="m" metricValue="2.6" pageId="8" pageNumber="9" unit="km" value="26.0">26 km</quantity>
of uninterrupted passage in length. Despite the significant amount of subterranean habitat found within the Kipuka Kanohina svstem,
<taxonomicName id="4C4A20E9315CBF0D1C73F9A0FE05F9F8" box="[319,471,1554,1578]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D1C73F9A0FE05F9F8" box="[319,471,1554,1578]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
has been collected from onlv four cave sections (of&gt;30 surveved) with an elevational range from
<quantity id="4CB2F68F315CBF0D1DF5F9E3FE83F9BA" box="[185,337,1616,1640]" metricMagnitude="2" metricUnit="m" metricValue="8.175" metricValueMax="10.959999999999999" metricValueMin="5.39" pageId="8" pageNumber="9" unit="m" value="817.5" valueMax="1096.0" valueMin="539.0">539 to 1096 m</quantity>
, suggesting this species might be restricted to the higher-elevation passages.
</paragraph>
</subSubSection>
<subSubSection id="C35008E1315CBF0D1C17F914FA73FEB6" pageId="8" pageNumber="9" type="biology_ecology">
<paragraph id="8BF55B6A315CBF0D1C17F914FDFDF913" blockId="8.[347,559,1703,1729]" box="[347,559,1703,1729]" pageId="8" pageNumber="9">
<emphasis id="B93E8778315CBF0D1C17F914FDFDF913" box="[347,559,1703,1729]" italics="true" pageId="8" pageNumber="9">Ecology and behaviour</emphasis>
</paragraph>
<paragraph id="8BF55B6A315CBF0D1DCDF97DFA73FEB6" blockId="8.[129,778,1742,1985]" lastBlockId="8.[825,1475,144,356]" pageId="8" pageNumber="9">
There is little documented information on the ecologv and behaviour of
<taxonomicName id="4C4A20E9315CBF0D1C5FF95DFE7DF8D7" box="[275,431,1773,1797]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D1C5FF95DFE7DF8D7" box="[275,431,1773,1797]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
. Adults and nvmphs have been observed on and around roots and on cave walls near roots anvwhere from twilight into the deep cave zone of lava tubes (for lava tube zonation, see
<bibRefCitation id="EFDB269B315CBF0D1C3CF8F8FDD4F8B1" author="Howarth FG" box="[368,518,1867,1891]" pageId="8" pageNumber="9" pagination="365 - 89" refId="ref8978" refString="Howarth FG. Ecologv of cave arthropods. Annual Review of Entomology 1983; 28: 365 - 89. hups: // doi. org / 10.1146 / annurev. en. 28.010183.002053" type="journal article" year="1983">Howarth 1983</bibRefCitation>
), although anecdotallv
<taxonomicName id="4C4A20E9315CBF0D1FB7F8FFFED6F851" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D1FB7F8FFFED6F851" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
appears to be more abundant close to entrance and twilight zones. In two collection locations, the roots have been confirmed to be from the native tree
<taxonomicName id="4C4A20E9315CBF0D1F43F81AFC4AFF7A" authority="Gaudich." authorityName="Gaudich." box="[527,920,144,1985]" class="Magnoliopsida" family="Myrtaceae" genus="Metrosideros" kingdom="Plantae" order="Myrtales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="polymorpha">
<emphasis id="B93E8778315CBF0D1F43F81AFCD8F813" box="[527,778,1961,1985]" italics="true" pageId="8" pageNumber="9">Metrosideros polymorpha</emphasis>
Gaudich.
</taxonomicName>
using barcoding techniques (unpublished data B. Chong and M. Porter). Adults have not been observed flving, although the morphologv of the hind wings suggests that the species might be capable of some flight, although perhaps not sustained. In all locations sampled,
<taxonomicName id="4C4A20E9315CBF0D19E3FEBDFA9AFEF7" box="[1199,1352,269,293]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D19E3FEBDFA9AFEF7" box="[1199,1352,269,293]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
individuals have been collected from dense root mats and have been observed to co-occur with
<taxonomicName id="4C4A20E9315CBF0D1962FEFFFA53FEB6" authority="Fennah, 1973" authorityName="Fennah" authorityYear="1973" box="[1070,1409,332,356]" class="Insecta" family="Cixiidae" genus="Oliarus" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" sensu="lato" species="polyphemus">
<emphasis id="B93E8778315CBF0D1962FEFFFB22FEB6" box="[1070,1264,332,356]" italics="true" pageId="8" pageNumber="9">Oliarus polyphemus</emphasis>
<bibRefCitation id="EFDB269B315CBF0D19B9FEFFFA53FEB6" author="Fennah RG" box="[1269,1409,332,356]" pageId="8" pageNumber="9" pagination="181 - 4" refId="ref8621" refString="Fennah RG. Ne cavernicolous fauna of Hawaiian lava tubes, 4. Two new blind Oliarus (Fulgoroidea: Cixiidae). Pacific Insects 1973; 15: 181 - 4." type="journal article" year="1973">Fennah, 1973</bibRefCitation>
</taxonomicName>
<taxonomicNameLabel id="A20D3A03315CBF0D18CAFEFFFA73FEB6" box="[1414,1441,332,356]" pageId="8" pageNumber="9" sensu="lato">s.l.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="C35008E1315CBF0D1EA3FECFFB23FDEA" pageId="8" pageNumber="9" type="multiple">
<paragraph id="8BF55B6A315CBF0D1EA3FECFFADFFE44" blockId="8.[825,1474,380,568]" box="[1007,1293,380,407]" pageId="8" pageNumber="9">
<emphasis id="B93E8778315CBF0D1EA3FECFFADFFE44" box="[1007,1293,380,407]" italics="true" pageId="8" pageNumber="9">Mitochondrial COI phylogeny</emphasis>
</paragraph>
<paragraph id="8BF55B6A315CBF0D1E75FE10FB23FDEA" blockId="8.[825,1474,380,568]" pageId="8" pageNumber="9">
Phvlogenetic relationships inferred from
<emphasis id="B93E8778315CBF0D19B8FE10FAF2FE68" box="[1268,1312,419,442]" italics="true" pageId="8" pageNumber="9">COI</emphasis>
sequence data support monophvlv of
<taxonomicName id="4C4A20E9315CBF0D196BFE70FB6CFE08" box="[1063,1214,450,474]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D196BFE70FB6CFE08" box="[1063,1214,450,474]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
and stronglv support the sister relationship between
<taxonomicName id="4C4A20E9315CBF0D191CFE51FB35FE2B" box="[1104,1255,481,505]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="ssankanstonei">
<emphasis id="B93E8778315CBF0D191CFE51FB35FE2B" box="[1104,1255,481,505]" italics="true" pageId="8" pageNumber="9">I. frankanstonei</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4A20E9315CBF0D1855FE51FAAAFE2B" authorityName="Kirkaldv" authorityYear="1902" box="[1305,1400,481,505]" class="Insecta" family="Cixiidae" genus="Iolania" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="species" species="perkinsi">
<emphasis id="B93E8778315CBF0D1855FE51FAAAFE2B" box="[1305,1400,481,505]" italics="true" pageId="8" pageNumber="9">I. perkinsi</emphasis>
</taxonomicName>
, which together form a distinctlv different clade from the Hawaiian
<taxonomicName id="4C4A20E9315CBF0D1E75FD93FC51FDEA" authorityName="Stal" authorityYear="1862" box="[825,899,544,568]" class="Insecta" family="Cixiidae" genus="Oliarus" kingdom="Animalia" order="Hemiptera" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis id="B93E8778315CBF0D1E75FD93FC51FDEA" box="[825,899,544,568]" italics="true" pageId="8" pageNumber="9">Oliarus</emphasis>
</taxonomicName>
samples (
<figureCitation id="137147EF315CBF0D1EA5FD93FBF3FDEA" box="[1001,1057,544,568]" captionStart="Figure 6" captionStartId="9.[113,178,1834,1858]" captionTargetBox="[156,1411,147,1804]" captionTargetId="graphics-6@9.[156,1221,158,1782]" captionText="Figure 6. Maximum likelihood phvlogenv of representative cixiid species of the two separate lineages that have colonized Hawaii:Iolania and Oliarus, Aphrophoridae (Philaenus spumarius), and Cercopidae (Callitetix versicolor), and the aphid Acyrthosiphon pisum used as an outgroup. Bootstrap values from 1000 replicates that were&gt;70% are included to indicate branch support.GenBank accession numbers for all sequences used are provided in parantheses asser species names. New sequences generated for this studv are in bold, and troglobiont species are indicated bv grev boxes. Arrows indicate separate cave colonization events in both cixiid lineages on Hawaiʻi Island." pageId="8" pageNumber="9">Fig. 6</figureCitation>
;
<bibRefCitation id="EFDB269B315CBF0D1962FD93FB32FDEA" author="Chong T &amp; Steck M &amp; Porter ML" box="[1070,1248,544,568]" pageId="8" pageNumber="9" pagination="1015 - 7" refId="ref8561" refString="Chong T, Steck M, Porter ML. Complete mitochondrial genomes and phvlogenetic analvsis of the Hawaiian planthoppers Iolania perkinsi and Oliarus cf. filicicola (Hemiptera: Cixiidae). Mitochondrial DNA. Part B, Resources 2022; 7: 1015 - 7. hups: // doi. org / 10.1080 / 238023 59.2022. 2080596" type="journal article" year="2022">
Chong
<emphasis id="B93E8778315CBF0D1936FD92FB7AFDEA" box="[1146,1192,544,568]" italics="true" pageId="8" pageNumber="9">et al.</emphasis>
2022
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title id="0C1AFA6757338AA3B40BFB3C4F7F0C3C">Two new taxa of brush-footed butterflies (Lepidoptera: Nymphalidae) from the Western Ghats biodiversity hotspot, southern India</mods:title>
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<mods:namePart id="C3E508DD8D92A4792F9D645818E2810C">Kunte, Krushnamegh</mods:namePart>
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<mods:affiliation id="3D2F31A902CEB8FCE739C8317243F7DD">National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bellary Road, Bengaluru 560 065, India</mods:affiliation>
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<mods:namePart id="3564ACA2DA58619CEAA11530FE7B0DF7">Basu, Dipendra Nath</mods:namePart>
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<mods:affiliation id="E26DD72BC87CF5C8232B20DDC64A2B90">National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bellary Road, Bengaluru 560 065, India &amp; Ashoka University, Rajiv Gandhi Education City, Sonipat, Haryana 131029, India</mods:affiliation>
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<paragraph id="E971BA41DA0A782421B4864FAFF8F891" blockId="20.[151,966,1835,1862]" box="[151,966,1835,1862]" pageId="20" pageNumber="363">
<heading id="B2390D2DDA0A782421B4864FAFF8F891" bold="true" box="[151,966,1835,1862]" fontSize="11" level="1" pageId="20" pageNumber="363" reason="1">
<emphasis id="DBBA6653DA0A782421B4864FAFF8F891" bold="true" box="[151,966,1835,1862]" pageId="20" pageNumber="363">
A note on the taxonomic status of
<taxonomicName id="2ECEC1C2DA0A7824231D864FAFF8F891" ID-CoL="4CHBM" authority="Swinhoe, 1917" authorityName="Swinhoe" authorityYear="1917" box="[574,966,1835,1862]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0A7824231D864FAF35F892" bold="true" box="[574,779,1835,1862]" italics="true" pageId="20" pageNumber="363">Pantoporia mera</emphasis>
<bibRefCitation id="8D5FC7B0DA0A78242231864FAFF8F891" author="Swinhoe, C." box="[786,966,1835,1861]" pageId="20" pageNumber="363" pagination="407 - 412" refId="ref15041" refString="Swinhoe, C. (1917) New Indo-Malayan Lepidoptera. Annals and Magazine of Natural History, 20, 407 - 412. https: // doi. org / 10.1080 / 00222931709487028" type="journal article" year="1917">Swinhoe, 1917</bibRefCitation>
</taxonomicName>
</emphasis>
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</paragraph>
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<paragraph id="E971BA41DA0A782521B48617AD64FDE9" blockId="20.[151,1437,1907,2042]" lastBlockId="21.[151,1437,151,573]" lastPageId="21" lastPageNumber="364" pageId="20" pageNumber="363">
<bibRefCitation id="8D5FC7B0DA0A782421B48617AD74F859" author="Swinhoe, C." box="[151,330,1907,1933]" pageId="20" pageNumber="363" pagination="407 - 412" refId="ref15041" refString="Swinhoe, C. (1917) New Indo-Malayan Lepidoptera. Annals and Magazine of Natural History, 20, 407 - 412. https: // doi. org / 10.1080 / 00222931709487028" type="journal article" year="1917">Swinhoe (1917)</bibRefCitation>
described
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<emphasis id="DBBA6653DA0A782420E18610AE28F859" box="[450,534,1908,1933]" italics="true" pageId="20" pageNumber="363">P. mera</emphasis>
</taxonomicName>
as follows: “
<taxonomicName id="2ECEC1C2DA0A7824238B8610AF59F859" authorityName="Swinhoe" authorityYear="1917" box="[680,871,1908,1933]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0A7824238B8610AF59F859" box="[680,871,1908,1933]" italics="true" pageId="20" pageNumber="363">Pantoporia mera</emphasis>
</taxonomicName>
, nov.
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. Belongs to the
<taxonomicName id="2ECEC1C2DA0A782425518610A891F859" box="[1138,1199,1908,1933]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0A782425518610A891F859" box="[1138,1199,1908,1933]" italics="true" pageId="20" pageNumber="363">inara</emphasis>
</taxonomicName>
group, is smaller, the fore wing shorter, the outer margin is less concave, the pattern on the upperside is very similar, but the cell-streak is more attenuated, the white spot in it smaller, the orange submarginal band incomplete, narrower at the apex, then broken, a spot in continuation on the middle, and faint indications of a grey line downwards to the hinder angle. The underside, however, is very different to the underside of
<taxonomicName id="2ECEC1C2DA0B7825223081F3AF6EFF64" box="[787,848,151,176]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0B7825223081F3AF6EFF64" box="[787,848,151,176]" italics="true" pageId="21" pageNumber="364">inara</emphasis>
</taxonomicName>
, especially on the hind wing, the middle white band being more even, not attenuated to a point at the abdominal fold as it is in
<taxonomicName id="2ECEC1C2DA0B782522CE81DFA814FF00" box="[1005,1066,187,212]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0B782522CE81DFA814FF00" box="[1005,1066,187,212]" italics="true" pageId="21" pageNumber="364">inara</emphasis>
</taxonomicName>
; the black discal row of spots is absent or only very faintly indicated, and the white postdiscal band is broader. Expanse of wings
<superScript id="1EBB1709DA0B782525F981BBA8CEFF38" attach="none" box="[1242,1264,222,249]" fontSize="6" pageId="21" pageNumber="364">22</superScript>
/
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inches
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.
<emphasis id="DBBA6653DA0B7825244681BBA9ABFF2D" box="[1381,1429,223,249]" italics="true" pageId="21" pageNumber="364">Hab</emphasis>
. Borneo.” Thus, Swinhoe had already realised that
<taxonomicName id="2ECEC1C2DA0B782523F08067AF17FEC8" authorityName="Swinhoe" authorityYear="1917" box="[723,809,259,284]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0B782523F08067AF17FEC8" box="[723,809,259,284]" italics="true" pageId="21" pageNumber="364">P. mera</emphasis>
</taxonomicName>
was close to
<taxonomicName id="2ECEC1C2DA0B782522E78067A81FFEC8" box="[964,1057,259,285]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0B782522E78067A81FFEC8" box="[964,1057,259,285]" italics="true" pageId="21" pageNumber="364">A. inara</emphasis>
</taxonomicName>
. Comparison of primary
<typeStatus id="367504E3DA0B782524618067A943FEC9" box="[1346,1405,259,285]" pageId="21" pageNumber="364">types</typeStatus>
of these two taxa and images of
<emphasis id="DBBA6653DA0B782520CF8043AE5FFE94" box="[492,609,295,321]" italics="true" pageId="21" pageNumber="364">
A. i.
<taxonomicName id="2ECEC1C2DA0B782523078043AE5FFE94" box="[548,609,295,320]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
in its E Himalayan and NE Indian range reveals that the slight variation represented by the
<typeStatus id="367504E3DA0B78252044802FADA9FEB1" box="[359,407,331,357]" pageId="21" pageNumber="364">type</typeStatus>
of
<taxonomicName id="2ECEC1C2DA0B78252099802FADCAFEB0" authorityName="Swinhoe" authorityYear="1917" box="[442,500,331,356]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0B78252099802FADCAFEB0" box="[442,500,331,356]" italics="true" pageId="21" pageNumber="364">mera</emphasis>
</taxonomicName>
falls within the intra-population variation seen in the range of
<taxonomicName id="2ECEC1C2DA0B78252581802FA8C3FEB0" box="[1186,1277,331,357]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0B78252581802FA8C3FEB0" box="[1186,1277,331,357]" italics="true" pageId="21" pageNumber="364">A. inara</emphasis>
</taxonomicName>
. Furthermore, the
<typeStatus id="367504E3DA0B782521E1800BACCCFE5D" box="[194,242,367,393]" pageId="21" pageNumber="364">type</typeStatus>
locality of
<taxonomicName id="2ECEC1C2DA0B7825205A800BAD8DFE5C" authorityName="Swinhoe" authorityYear="1917" box="[377,435,367,392]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0B7825205A800BAD8DFE5C" box="[377,435,367,392]" italics="true" pageId="21" pageNumber="364">mera</emphasis>
</taxonomicName>
specified by Swinhoe—“Borneo”—is clearly in error. The phenotype of
<taxonomicName id="2ECEC1C2DA0B782525D9800BA90AFE5C" authorityName="Swinhoe" authorityYear="1917" box="[1274,1332,367,392]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0B782525D9800BA90AFE5C" box="[1274,1332,367,392]" italics="true" pageId="21" pageNumber="364">mera</emphasis>
</taxonomicName>
does not occur in Borneo, where
<taxonomicName id="2ECEC1C2DA0B7825208B80F7AF6EFE79" authority="(Fruhstorfer, 1913)" baseAuthorityName="Fruhstorfer" baseAuthorityYear="1913" box="[424,848,403,429]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="subSpecies" species="nefte" subSpecies="matthiola">
<emphasis id="DBBA6653DA0B7825208B80F7AE4CFE79" box="[424,626,403,429]" italics="true" pageId="21" pageNumber="364">A. nefte matthiola</emphasis>
(Fruhstorfer, 1913)
</taxonomicName>
(
<typeStatus id="367504E3DA0B7825224380F7AFACFE79" box="[864,914,403,429]" pageId="21" pageNumber="364">type</typeStatus>
locality Borneo) represents a black-and-white male phenotype very similar to
<emphasis id="DBBA6653DA0B7825232E80D3AEBBFE05" box="[525,645,439,465]" italics="true" pageId="21" pageNumber="364">
A. n.
<taxonomicName id="2ECEC1C2DA0B7825236C80D3AEBBFE05" baseAuthorityName="Cramer" baseAuthorityYear="1780" box="[591,645,439,465]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="nefte">nefte</taxonomicName>
</emphasis>
. Thus, it is likely that the
<typeStatus id="367504E3DA0B782522E780D3AFCAFE05" box="[964,1012,439,465]" pageId="21" pageNumber="364">type</typeStatus>
of
<taxonomicName id="2ECEC1C2DA0B7825250280D3A865FE04" authorityName="Swinhoe" authorityYear="1917" box="[1057,1115,439,464]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA0B7825250280D3A865FE04" box="[1057,1115,439,464]" italics="true" pageId="21" pageNumber="364">mera</emphasis>
</taxonomicName>
was collected in NE
<collectingCountry id="91D9FAD1DA0B7825247F80D3A9A6FE05" box="[1372,1432,439,465]" name="India" pageId="21" pageNumber="364">India</collectingCountry>
,
<collectingCountry id="91D9FAD1DA0B782521B480BFAD24FE21" box="[151,282,475,501]" name="Bangladesh" pageId="21" pageNumber="364">Bangladesh</collectingCountry>
or N
<collectingCountry id="91D9FAD1DA0B7825204180BFADF1FE21" box="[354,463,475,501]" name="Myanmar" pageId="21" pageNumber="364">Myanmar</collectingCountry>
but mistakenly labelled as being from Borneo. Therefore, based on the similarity of the
<typeStatus id="367504E3DA0B782521E2809BACCFFDCD" box="[193,241,511,537]" pageId="21" pageNumber="364">type</typeStatus>
specimens (
<figureCitation id="71F5A6C4DA0B7825205E809BADFCFDCD" box="[381,450,511,537]" captionStart="FIGURE 9" captionStartId="16.[151,250,1981,2005]" captionTargetBox="[168,1415,786,1947]" captionTargetId="figure-368@16.[164,1423,777,1958]" captionTargetPageId="16" captionText="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386322" httpUri="https://zenodo.org/record/14386322/files/figure.png" pageId="21" pageNumber="364">Fig. 9</figureCitation>
), we propose synonymy of
<emphasis id="DBBA6653DA0B782523DF809BA8C5FDCD" bold="true" box="[764,1275,510,537]" pageId="21" pageNumber="364">
<taxonomicName id="2ECEC1C2DA0B782523DF809BA84DFDCC" authority="Swinhoe, 1917" authorityName="Swinhoe" authorityYear="1917" box="[764,1139,510,537]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="mera" status="syn. nov.">
<emphasis id="DBBA6653DA0B782523DF809BAFFEFDCD" bold="true" box="[764,960,511,537]" italics="true" pageId="21" pageNumber="364">Pantoporia mera</emphasis>
<bibRefCitation id="8D5FC7B0DA0B782522EB809AA84DFDCC" author="Swinhoe, C." box="[968,1139,510,536]" pageId="21" pageNumber="364" pagination="407 - 412" refId="ref15041" refString="Swinhoe, C. (1917) New Indo-Malayan Lepidoptera. Annals and Magazine of Natural History, 20, 407 - 412. https: // doi. org / 10.1080 / 00222931709487028" type="journal article" year="1917">Swinhoe, 1917</bibRefCitation>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0B78252558809BA8DDFDCD" box="[1147,1251,511,537]" pageId="21" pageNumber="364">syn. nov.</taxonomicNameLabel>
=
</emphasis>
<taxonomicName id="2ECEC1C2DA0B78252421809BAD68FDE9" authority="Westwood, 1850" authorityName="Westwood" authorityYear="1850" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="21" pageNumber="364" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0B78252421809BA9A3FDCC" box="[1282,1437,511,537]" italics="true" pageId="21" pageNumber="364">Athyma inara</emphasis>
Westwood, 1850
</taxonomicName>
.
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<mods:name id="F7F92DDA55D9A2C6B9D6362FBDA71299" type="personal">
<mods:role id="B495E541F2A4404402D8F2ACD288B54D">
<mods:roleTerm id="D55DB86FA4EF9A772DB6EDB1C39A7DF6">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="C72B9DA21C1A8601B64FF9F5EC9AA4EA">Kunte, Krushnamegh</mods:namePart>
<mods:nameIdentifier id="2CBBFBA29644549DBCD0BD38E7CC1A5B" type="ORCID">0000-0002-3860-6118</mods:nameIdentifier>
<mods:affiliation id="0567B6E727F86A4C6F92D9075D0A19DA">National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bellary Road, Bengaluru 560 065, India</mods:affiliation>
<mods:nameIdentifier id="7CFD3586A8CE21686C6DF6720C2106B3" type="email">krushnamegh@ncbs.res.in</mods:nameIdentifier>
</mods:name>
<mods:name id="D780B6F8DE348AD289A3985BBDD095D5" type="personal">
<mods:role id="981E3407A3F36081A697A34211196751">
<mods:roleTerm id="F6BE7A61802032111D04BCB66CBCB696">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="F3E75EAD468C6C2F2B65FFE6B703B5EF">Basu, Dipendra Nath</mods:namePart>
<mods:nameIdentifier id="DEDEAF152DE35CFB39C007E1D83E7EFE" type="ORCID">0000-0002-1529-0931</mods:nameIdentifier>
<mods:affiliation id="EE3D07A2FF0EC0BB2A07FB8976973F2F">National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bellary Road, Bengaluru 560 065, India &amp; Ashoka University, Rajiv Gandhi Education City, Sonipat, Haryana 131029, India</mods:affiliation>
<mods:nameIdentifier id="9F75C8EAD58EB371FDBF1A54CC126C4A" type="email">dipendra.basu@ashoka.edu.in</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource id="EDE038DF3DD41D3AFA9E4447F1F25AC4">text</mods:typeOfResource>
<mods:relatedItem id="166144C35CDED08C20591ADB7ECC845B" type="host">
<mods:titleInfo id="88C2729432919AA331CDA3ADDEFA2B23">
<mods:title id="DC384F70F69D77C5CC30C92B614ECA16">Zootaxa</mods:title>
</mods:titleInfo>
<mods:part id="18D40FC305E3F6340715B0A48C20F1C7">
<mods:date id="6BB72543D4A893E1E04AA067F120C4D3">2024</mods:date>
<mods:detail id="685A460CE00D8C3B46F61BD7608A4515" type="pubDate">
<mods:number id="4D3EBF59A79A1CC1F174EEA30EA507E5">2024-12-04</mods:number>
</mods:detail>
<mods:detail id="5ECBF5B24DD0C01E49CF1B8AF8A0B4C2" type="volume">
<mods:number id="4F4AA3939D1640FAA4E506529841E735">5543</mods:number>
</mods:detail>
<mods:detail id="D727BED1D46DCECAA886BBE4BFFDA9DE" type="issue">
<mods:number id="14F5EC4DCDC0704A3D28EEAB4B5AC5EC">3</mods:number>
</mods:detail>
<mods:extent id="08DD10C5C5C9486AA7F218B029C68F6B" unit="page">
<mods:start id="8377DCBA4BCD23D9724638062C3DAC38">343</mods:start>
<mods:end id="2C2F28CDA5B4AE769408C58FE8609083">367</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location id="D7B49704433F8153EE3BD924F5EA1120">
<mods:url id="431E34DBB156F31683D1E9A0C58F7921">https://doi.org/10.11646/zootaxa.5543.3.3</mods:url>
</mods:location>
<mods:classification id="6CC0C912DCC435D706A73A7352C40570">journal article</mods:classification>
<mods:identifier id="2FD30F9959C1D6DA3F1DD39CFBF6866D" type="DOI">10.11646/zootaxa.5543.3.3</mods:identifier>
<mods:identifier id="7C72650CD43725B58CC5D446DF252CB4" type="ISSN">1175-5326</mods:identifier>
<mods:identifier id="81944AA5C29F61C43F010B6332FC33A7" type="Zenodo-Dep">14386297</mods:identifier>
<mods:identifier id="62FABA90F3C8CE04CCAD04F936D0314C" type="ZooBank">C90518B7-B880-4290-95FE-A0F51EDE1E98</mods:identifier>
</mods:mods>
<treatment id="61670B57DA13782421B481F3AF9FFE21" ID-DOI="http://doi.org/10.5281/zenodo.14386644" ID-Zenodo-Dep="14386644" LSID="urn:lsid:plazi:treatment:61670B57DA13782421B481F3AF9FFE21" httpUri="http://treatment.plazi.org/id/61670B57DA13782421B481F3AF9FFE21" lastPageId="20" lastPageNumber="363" pageId="13" pageNumber="356">
<subSubSection id="A1D4E9CADA13783D21B481F3AE8EFF65" box="[151,688,151,178]" pageId="13" pageNumber="356" type="nomenclature">
<paragraph id="E971BA41DA13783D21B481F3AE8EFF65" blockId="13.[151,688,151,178]" box="[151,688,151,178]" pageId="13" pageNumber="356">
<heading id="B2390D2DDA13783D21B481F3AE8EFF65" bold="true" box="[151,688,151,178]" fontSize="11" level="1" pageId="13" pageNumber="356" reason="1">
<emphasis id="DBBA6653DA13783D21B481F3AE8EFF65" bold="true" box="[151,688,151,178]" pageId="13" pageNumber="356">
<taxonomicName id="2ECEC1C2DA13783D21B481F3AE7EFF65" authority="Kunte" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[151,576,151,178]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="13" pageNumber="356" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA13783D21B481F3ADD5FF66" bold="true" box="[151,491,151,178]" italics="true" pageId="13" pageNumber="356">Athyma inara sahyadriensis</emphasis>
Kunte
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA13783D236481F3AE8EFF65" box="[583,688,151,177]" pageId="13" pageNumber="356" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA13783D21B481BBAE85FEB1" pageId="13" pageNumber="356" type="materials_examined">
<paragraph id="E971BA41DA13783D21B481BBAE85FEB1" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<materialsCitation id="59A6B01CDA13783D21B481BBAE85FEB1" collectingDate="2013-08-03" collectionCode="NCBS" collectorName="Krushnamegh Kunte" country="India" elevation="75" latitude="11.94" location="Kannur" longLatPrecision="558" longitude="75.765" municipality="Kannur District" pageId="13" pageNumber="356" specimenCode="IBC-PS982" specimenCount="1" specimenCount-male="1" stateProvince="Kerala" typeStatus="holotype">
<emphasis id="DBBA6653DA13783D21B481BBAD50FF2D" bold="true" box="[151,366,223,249]" pageId="13" pageNumber="356">
<typeStatus id="367504E3DA13783D21B481BBAD3AFF2D" box="[151,260,223,249]" pageId="13" pageNumber="356" type="holotype">Holotype</typeStatus>
(
<figureCitation id="71F5A6C4DA13783D203681BBAD61FF2D" box="[277,351,223,249]" captionStart="FIGURE 7" captionStartId="14.[151,250,1820,1844]" captionTargetBox="[282,1304,194,1783]" captionTargetId="figure-20@14.[274,1314,181,1796]" captionTargetPageId="14" captionText="FIGURE 7. Type specimens of Athyma inara sahyadriensis ssp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386318" httpUri="https://zenodo.org/record/14386318/files/figure.png" pageId="13" pageNumber="356">Fig. 7</figureCitation>
):
</emphasis>
Voucher code
<specimenCode id="B968123ADA13783D233F81BBAE9FFF2D" box="[540,673,223,249]" pageId="13" pageNumber="356">IBC-PS982</specimenCode>
.
<specimenCount id="FFC871C8DA13783D238D8184AEF8FF2E" box="[686,710,224,250]" count="1" pageId="13" pageNumber="356" type="male">♂</specimenCount>
. Near
<location id="EC11EC9ADA13783D22308184AF57FF2E" LSID="urn:lsid:plazi:treatment:61670B57DA13782421B481F3AF9FFE21:EC11EC9ADA13783D22308184AF57FF2E" box="[787,873,224,250]" country="India" latitude="11.94" longLatPrecision="558" longitude="75.765" municipality="Kannur District" name="Kannur" pageId="13" pageNumber="356" stateProvince="Kerala">Kannur</location>
,
<collectingMunicipality id="0915203BDA13783D22558184A815FF2D" box="[886,1067,223,250]" pageId="13" pageNumber="356">Kannur District</collectingMunicipality>
,
<collectingRegion id="2B0A74A3DA13783D251B81BBA8BBFF2D" box="[1080,1157,223,249]" country="India" name="Kerala" pageId="13" pageNumber="356">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA13783D25B081BBA8F3FF2D" box="[1171,1229,223,249]" name="India" pageId="13" pageNumber="356">India</collectingCountry>
(lat-long: approx.
<geoCoordinate id="8CFADC86DA13783D21B48067ACCAFECA" box="[151,244,259,286]" degrees="11.94" direction="north" orientation="latitude" pageId="13" pageNumber="356" precision="555" value="11.94">11.94 N</geoCoordinate>
,
<geoCoordinate id="8CFADC86DA13783D20238067AD54FEC9" box="[256,362,259,286]" degrees="75.765" direction="east" orientation="longitude" pageId="13" pageNumber="356" precision="55" value="75.765">75.765 E</geoCoordinate>
). Approx.
<quantity id="2E3617A4DA13783D20C18067AE6AFECA" box="[482,596,259,286]" metricMagnitude="1" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="13" pageNumber="356" unit="m" value="75.0" valueMax="100.0" valueMin="50.0">
<elevation id="62E35D72DA13783D20C18067AE6AFECA" box="[482,596,259,286]" metricMagnitude="1" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="13" pageNumber="356" unit="m" value="75.0" valueMax="100.0" valueMin="50.0">50–100 m</elevation>
</quantity>
asl. Evergreen forest. Raised from a caterpillar collected from
<taxonomicName id="2ECEC1C2DA13783D243D8067AC89FE95" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="species" species="undetermined">
<emphasis id="DBBA6653DA13783D243D8067AC89FE95" italics="true" pageId="13" pageNumber="356">Glochidion sp.</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA13783D21E48043AD4AFE95" authorityName="Martinov" authorityYear="1820" box="[199,372,295,321]" class="Magnoliopsida" family="Phyllanthaceae" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="family">Phyllanthaceae</taxonomicName>
) by Saji K., eclosed and pinned dry on
<date id="9D709C81DA13783D221F8043AF85FE95" box="[828,955,295,321]" pageId="13" pageNumber="356" value="2013-08-03">
<collectingDate id="8D346569DA13783D221F8043AF85FE95" box="[828,955,295,321]" pageId="13" pageNumber="356" value="2013-08-03">2013/03/08</collectingDate>
</date>
by
<collectorName id="443BDF97DA13783D22CB8043A8E8FE96" box="[1000,1238,295,322]" pageId="13" pageNumber="356">Krushnamegh Kunte</collectorName>
. Deposited in the Biodiversity Lab Research Collections at
<collectionCode id="8FDF2284DA13783D2348802FAE86FEB1" box="[619,696,331,357]" pageId="13" pageNumber="356">NCBS</collectionCode>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA13783D21E4800BAF75FBA9" pageId="13" pageNumber="356" type="description">
<paragraph id="E971BA41DA13783D21E4800BAD89FE5D" blockId="13.[151,1437,223,1150]" box="[199,439,367,393]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E4800BAD89FE5D" bold="true" box="[199,439,367,393]" pageId="13" pageNumber="356">
Description (
<figureCitation id="71F5A6C4DA13783D2042800BAD96FE5D" box="[353,424,367,393]" captionStart="FIGURE 7" captionStartId="14.[151,250,1820,1844]" captionTargetBox="[282,1304,194,1783]" captionTargetId="figure-20@14.[274,1314,181,1796]" captionTargetPageId="14" captionText="FIGURE 7. Type specimens of Athyma inara sahyadriensis ssp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386318" httpUri="https://zenodo.org/record/14386318/files/figure.png" pageId="13" pageNumber="356">Fig. 7</figureCitation>
):
</emphasis>
</paragraph>
<paragraph id="E971BA41DA13783D21E480F0AF08FDCD" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E480F0ADBCFE79" box="[199,386,403,429]" italics="true" pageId="13" pageNumber="356">Forewing length</emphasis>
:
<quantity id="2E3617A4DA13783D20B080F7ADD9FE7A" box="[403,487,403,430]" metricMagnitude="-2" metricUnit="m" metricValue="2.9" pageId="13" pageNumber="356" unit="mm" value="29.0">29 mm</quantity>
.
<emphasis id="DBBA6653DA13783D20D080F7AEFDFE79" box="[499,707,403,429]" italics="true" pageId="13" pageNumber="356">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA13783D23F780F7AF16FE7A" box="[724,808,403,430]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="356" unit="mm" value="17.0">17 mm</quantity>
. Head black. Eyes pale grey, spotted dark brown (eyes turn dark brown or black upon preservation). Antennae black. Proboscis grey-brown. Thorax black above with two very faint, diffused white or pale grey spots; white below. Abdomen black above with white bands on the first two abdominal segments; white below. Legs pale greyish-white.
</paragraph>
<paragraph id="E971BA41DA13783D21E48347ADCDFC8A" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E48347ADF0FDE9" box="[199,462,547,573]" italics="true" pageId="13" pageNumber="356">Dorsal side: Forewings</emphasis>
black.A very narrow, faint, diffused white line from the base to the centre of discal cell, and a large white spot at the end of discal cell. Large white spots stacked in line in interspaces 1a, 1b and 2, and then again in interspaces 4–6. A very small, diffused pale greyish-white spot in interspace 3. Dark orange subapical spots in interspaces 4 and 5, with very small, inconspicuous spots in 3 and 5. An inconspicuous, diffused, pale grey submarginal line running parallel to termen, separated from a similar line that continues from the orange subapical spots to dorsum.
<emphasis id="DBBA6653DA13783D207683B3ADF0FD25" box="[341,462,727,753]" italics="true" pageId="13" pageNumber="356">Hindwings</emphasis>
black. A white band from the base through the cell to v6, the spots in 7 separated by the grey and black v7. A narrow, orange post-discal band interrupted by black veins from interspaces 1c to 7, its edges smooth, straight, not prominently convex. An inconspicuous, diffused, pale grey submarginal line running parallel to termen, from tornus to apex.
</paragraph>
<paragraph id="E971BA41DA13783D21E48203AF26FBC5" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E48203ADF8FC55" box="[199,454,871,897]" italics="true" pageId="13" pageNumber="356">Ventral side: Forewing</emphasis>
tawny with grey and white markings that are similar to those on the dorsal side, and with additional black smudges surrounding the white spots. Basal line through the discal cell is broader and grey, and conspicuous (rather than narrow, white and inconspicuous, as on the dorsal side).
<emphasis id="DBBA6653DA13783D251682CBA89DFC1D" box="[1077,1187,943,969]" italics="true" pageId="13" pageNumber="356">Hindwing</emphasis>
grey post-discal band (corresponding to the narrow, orange post-discal band on dorsal side) is continued from interspace 1b to 4, but not beyond. HW submarginal grey specks between the veins.
</paragraph>
<paragraph id="E971BA41DA13783D21E4857FAF75FBA9" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
This is a dry season form, in which UNH areas between the discal white band and post-discal grey band are plain, with dark reddish, highly diffused spots barely visible between the two bands (these spots are black and very prominent in the wet season form; see
<typeStatus id="367504E3DA13783D236B8500AE96FBAA" box="[584,680,1124,1150]" pageId="13" pageNumber="356" type="paratype">paratype</typeStatus>
IBC-AK310).
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA13783D21B485A6A8BBF9FF" pageId="13" pageNumber="356" type="materials_examined">
<materialsCitation id="59A6B01CDA13783D21B485A6AFC4FAA0" collectingDate="2014-02-15" collectionCode="NCBS" collectorName="Krushnamegh Kunte" country="India" elevation="1000" location="Ponmudi" municipality="Thiruvananthapuram District" pageId="13" pageNumber="356" specimenCode="IBC-AK310" specimenCount="1" specimenCount-male="1" stateProvince="Kerala" typeStatus="paratype">
<paragraph id="E971BA41DA13783D21B485A6AED2FB0F" blockId="13.[151,748,1217,1244]" box="[151,748,1217,1244]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21B485A6AED2FB0F" bold="true" box="[151,748,1217,1244]" pageId="13" pageNumber="356">
<typeStatus id="367504E3DA13783D21B485A6AD28FB08" box="[151,278,1218,1244]" pageId="13" pageNumber="356" type="paratype">Paratypes</typeStatus>
, and intraspecific variation (
<figureCitation id="71F5A6C4DA13783D235D85A5AEDAFB0F" box="[638,740,1217,1243]" captionStart-0="FIGURE 7" captionStart-1="FIGURE 8" captionStart-2="FIGURE 9" captionStartId-0="14.[151,250,1820,1844]" captionStartId-1="15.[151,250,1315,1339]" captionStartId-2="16.[151,250,1981,2005]" captionTargetBox-0="[282,1304,194,1783]" captionTargetBox-1="[171,1415,181,1291]" captionTargetBox-2="[168,1415,786,1947]" captionTargetId-0="figure-20@14.[274,1314,181,1796]" captionTargetId-1="figure-19@15.[171,1416,181,1291]" captionTargetId-2="figure-368@16.[164,1423,777,1958]" captionTargetPageId-0="14" captionTargetPageId-1="15" captionTargetPageId-2="16" captionText-0="FIGURE 7. Type specimens of Athyma inara sahyadriensis ssp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte." captionText-1="FIGURE 8. Male genitalia of Athyma inara sahyadriensis ssp. nov. (holotype, IBC-PW982) and A. i. inara. Photographs: Dipendra Nath Basu." captionText-2="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." figureDoi-0="http://doi.org/10.5281/zenodo.14386318" figureDoi-1="http://doi.org/10.5281/zenodo.14386320" figureDoi-2="http://doi.org/10.5281/zenodo.14386322" httpUri-0="https://zenodo.org/record/14386318/files/figure.png" httpUri-1="https://zenodo.org/record/14386320/files/figure.png" httpUri-2="https://zenodo.org/record/14386322/files/figure.png" pageId="13" pageNumber="356">Fig. 7–9</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="E971BA41DA13783D21B48464AFC4FAA0" blockId="13.[151,1437,1280,1580]" pageId="13" pageNumber="356">
<specimenCount id="FFC871C8DA13783D21B48464AC93FAC3" box="[151,173,1280,1303]" count="1" pageId="13" pageNumber="356" type="male">♂</specimenCount>
: voucher code
<specimenCode id="B968123ADA13783D20698464ADF4FACC" box="[330,458,1280,1304]" pageId="13" pageNumber="356">IBC-AK310</specimenCode>
.
<emphasis id="DBBA6653DA13783D20F68464AE40FACC" box="[469,638,1280,1304]" italics="true" pageId="13" pageNumber="356">Forewing length</emphasis>
:
<quantity id="2E3617A4DA13783D23AE8464AEE7FAC3" box="[653,729,1280,1304]" metricMagnitude="-2" metricUnit="m" metricValue="3.2" pageId="13" pageNumber="356" unit="mm" value="32.0">32 mm</quantity>
.
<emphasis id="DBBA6653DA13783D23C78464AFA1FAC3" box="[740,927,1280,1304]" italics="true" pageId="13" pageNumber="356">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA13783D228D8464AFC4FAC3" box="[942,1018,1280,1304]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="356" unit="mm" value="17.0">17 mm</quantity>
.
<location id="EC11EC9ADA13783D25278464A85CFACC" LSID="urn:lsid:plazi:treatment:61670B57DA13782421B481F3AF9FFE21:EC11EC9ADA13783D25278464A85CFACC" box="[1028,1122,1280,1304]" country="India" municipality="Thiruvananthapuram District" name="Ponmudi" pageId="13" pageNumber="356" stateProvince="Kerala">Ponmudi</location>
,
<collectingMunicipality id="0915203BDA13783D254D8464A9A7FACC" box="[1134,1433,1280,1304]" pageId="13" pageNumber="356">Thiruvananthapuram District</collectingMunicipality>
,
<collectingRegion id="2B0A74A3DA13783D21E4847AAD33FAE2" box="[199,269,1310,1334]" country="India" name="Kerala" pageId="13" pageNumber="356">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA13783D203B847AAD71FAE2" box="[280,335,1310,1334]" name="India" pageId="13" pageNumber="356">India</collectingCountry>
. Approx.
<quantity id="2E3617A4DA13783D2090847AAE39FAE2" box="[435,519,1310,1334]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="13" pageNumber="356" unit="m" value="1000.0">
<elevation id="62E35D72DA13783D2090847AAE39FAE2" box="[435,519,1310,1334]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="13" pageNumber="356" unit="m" value="1000.0">1,000 m</elevation>
</quantity>
asl. Raised from a caterpillar collected from
<taxonomicName id="2ECEC1C2DA13783D22F4847AA850FAE2" box="[983,1134,1310,1334]" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="species" species="undetermined">
<emphasis id="DBBA6653DA13783D22F4847AA850FAE2" box="[983,1134,1310,1334]" italics="true" pageId="13" pageNumber="356">Glochidion sp.</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA13783D2558847AA926FAE2" authorityName="Martinov" authorityYear="1820" box="[1147,1304,1310,1334]" class="Magnoliopsida" family="Phyllanthaceae" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="family">Phyllanthaceae</taxonomicName>
) by roadside in a tea estate by Baiju K., eclosed and pinned on
<date id="9D709C81DA13783D239E8459AF0FFA81" box="[701,817,1341,1365]" pageId="13" pageNumber="356" value="2014-02-15">
<collectingDate id="8D346569DA13783D239E8459AF0FFA81" box="[701,817,1341,1365]" pageId="13" pageNumber="356" value="2014-02-15">2014/02/15</collectingDate>
</date>
by
<collectorName id="443BDF97DA13783D22758459A815FA80" box="[854,1067,1341,1365]" pageId="13" pageNumber="356">Krushnamegh Kunte</collectorName>
. Legs preserved in 100% molecular-grade ethanol. Deposited in the Biodiversity Lab Research Collections at
<collectionCode id="8FDF2284DA13783D22938438AFCBFAA0" box="[944,1013,1372,1396]" country="USA" httpUri="http://grbio.org/cool/x0m9-a3bs" name="Yale University" pageId="13" pageNumber="356" type="Herbarium">NCBS</collectionCode>
.
</paragraph>
</materialsCitation>
<paragraph id="E971BA41DA13783D21B4841FA84CFA04" blockId="13.[151,1437,1280,1580]" pageId="13" pageNumber="356">
<materialsCitation id="59A6B01CDA13783D21B4841FA84CFA04" collectingDate="2013-08-03" collectionCode="NCBS" collectorName="Krushnamegh Kunte" country="India" elevation="75" latitude="11.94" location="Kannur" longLatPrecision="558" longitude="75.765" municipality="Kannur District" pageId="13" pageNumber="356" specimenCode="IBC-PS983" specimenCount="1" specimenCount-female="1" specimenCount-male="1" stateProvince="Kerala" typeStatus="holotype">
<specimenCount id="FFC871C8DA13783D21B4841FAC92FA46" box="[151,172,1403,1426]" count="1" pageId="13" pageNumber="356" type="female">♀</specimenCount>
: voucher code
<specimenCode id="B968123ADA13783D2066841EAD83FA46" box="[325,445,1402,1426]" pageId="13" pageNumber="356">IBC-PS983</specimenCode>
.
<emphasis id="DBBA6653DA13783D20E5841FAE53FA46" box="[454,621,1402,1426]" italics="true" pageId="13" pageNumber="356">Forewing length</emphasis>
:
<quantity id="2E3617A4DA13783D2358841EAEFBFA46" box="[635,709,1402,1426]" metricMagnitude="-2" metricUnit="m" metricValue="3.2" pageId="13" pageNumber="356" unit="mm" value="32.0">32 mm</quantity>
.
<emphasis id="DBBA6653DA13783D23ED841EAFB9FA46" box="[718,903,1402,1426]" italics="true" pageId="13" pageNumber="356">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA13783D22B0841EAFE0FA46" box="[915,990,1402,1426]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="356" unit="mm" value="17.0">17 mm</quantity>
. Collection and preservation details same as the holotype, except that legs and thoracic muscle tissue are preserved in 100% molecular-grade ethanol. Deposited in the Biodiversity Lab Research Collections at
<collectionCode id="8FDF2284DA13783D234D84DCAE8DFA04" box="[622,691,1464,1488]" country="USA" httpUri="http://grbio.org/cool/x0m9-a3bs" name="Yale University" pageId="13" pageNumber="356" type="Herbarium">NCBS</collectionCode>
. This paratype is also treated as an allotype.
</materialsCitation>
</paragraph>
<paragraph id="E971BA41DA13783D21B484B3A8BBF9FF" blockId="13.[151,1437,1280,1580]" pageId="13" pageNumber="356">
<materialsCitation id="59A6B01CDA13783D21B484B3A8BBF9FF" collectingDate="2016-10-20" collectorName="Kalesh Sadasivan" country="India" location="Near Periyar" municipality="Idukky District" pageId="13" pageNumber="356" specimenCode="IBC-AY231" specimenCount="801" specimenCount-female="1" specimenCount-male="800" stateProvince="Kerala">
<specimenCount id="FFC871C8DA13783D21B484B3AC92FA3A" box="[151,172,1495,1518]" count="1" pageId="13" pageNumber="356" type="female">♀</specimenCount>
: voucher code
<specimenCode id="B968123ADA13783D206084B2ADFFFA3A" box="[323,449,1494,1518]" pageId="13" pageNumber="356">IBC-AY231</specimenCode>
.
<emphasis id="DBBA6653DA13783D20E984B3AE4EFA3A" box="[458,624,1494,1518]" italics="true" pageId="13" pageNumber="356">Forewing length</emphasis>
:
<quantity id="2E3617A4DA13783D235F84B2AEF8FA3A" box="[636,710,1494,1518]" metricMagnitude="-2" metricUnit="m" metricValue="3.1" pageId="13" pageNumber="356" unit="mm" value="31.0">31 mm</quantity>
.
<emphasis id="DBBA6653DA13783D23ED84B2AFBBFA3A" box="[718,901,1494,1518]" italics="true" pageId="13" pageNumber="356">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA13783D22B284B2AFE5FA3A" box="[913,987,1494,1518]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="356" unit="mm" value="17.0">17 mm</quantity>
.
<location id="EC11EC9ADA13783D22C084B3A85BFA3A" LSID="urn:lsid:plazi:treatment:61670B57DA13782421B481F3AF9FFE21:EC11EC9ADA13783D22C084B3A85BFA3A" box="[995,1125,1494,1518]" country="India" municipality="Idukky District" name="Near Periyar" pageId="13" pageNumber="356" stateProvince="Kerala">Near Periyar</location>
,
<collectingMunicipality id="0915203BDA13783D254D84B2A934FA3A" box="[1134,1290,1494,1518]" pageId="13" pageNumber="356">Idukky District</collectingMunicipality>
,
<collectingRegion id="2B0A74A3DA13783D243084B2A967FA3A" box="[1299,1369,1494,1518]" country="India" name="Kerala" pageId="13" pageNumber="356">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA13783D244184B2A9A7FA3A" box="[1378,1433,1494,1518]" name="India" pageId="13" pageNumber="356">India</collectingCountry>
.
<specimenCount id="FFC871C8DA13783D21E48491AD31F9D8" box="[199,271,1525,1549]" count="800" pageId="13" pageNumber="356" type="male">
<quantity id="2E3617A4DA13783D21E48491AD35F9D8" box="[199,267,1525,1549]" metricMagnitude="2" metricUnit="m" metricValue="8.0" pageId="13" pageNumber="356" unit="m" value="800.0">800 m</quantity>
.
</specimenCount>
Evergreen forest. Preserved dry, with legs and thoracic muscle tissue preserved in 100% molecular-grade ethanol. Collected on
<date id="9D709C81DA13783D206D8777ADFCF9FF" box="[334,450,1555,1579]" pageId="13" pageNumber="356" value="2016-10-20">
<collectingDate id="8D346569DA13783D206D8777ADFCF9FF" box="[334,450,1555,1579]" pageId="13" pageNumber="356" value="2016-10-20">2016/10/20</collectingDate>
</date>
by
<collectorName id="443BDF97DA13783D20CB8770AEA3F9F8" box="[488,669,1556,1580]" pageId="13" pageNumber="356">Kalesh Sadasivan</collectorName>
, pinned on
<date id="9D709C81DA13783D22308777AFBBF9FF" box="[787,901,1555,1579]" pageId="13" pageNumber="356" value="2016-11-20">2016/11/20</date>
by Krushnamegh Kunte.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA13783F21B48732ADA1F90A" lastPageId="15" lastPageNumber="358" pageId="13" pageNumber="356" type="description">
<paragraph id="E971BA41DA13783D21B48732AE77F92B" blockId="13.[151,1437,1622,2039]" pageId="13" pageNumber="356">
<typeStatus id="367504E3DA13783D21B48732AD3AF9A4" box="[151,260,1622,1648]" pageId="13" pageNumber="356" type="paratype">Paratypes</typeStatus>
reveal considerable sexual and seasonal variation. IBC-AK310 is a wet season form in which UNH areas between the discal and post-discal white bands have large black spots, and UNF discal cell has all markings white (rather than grey as seen in the
<typeStatus id="367504E3DA13783D20D587F9AE63F963" box="[502,605,1693,1719]" pageId="13" pageNumber="356" type="holotype">holotype</typeStatus>
). In this specimen, UPF a spot in interspace 3 is still small but much more prominent because it is white and clearly defined (cf
<typeStatus id="367504E3DA13783D23D087A5AF67F90F" box="[755,857,1729,1755]" pageId="13" pageNumber="356" type="holotype">holotype</typeStatus>
), and UPH the post-discal orange band is narrower but darker, diffusing out towards apex.
</paragraph>
<paragraph id="E971BA41DA13783D21E4866CAD38F823" blockId="13.[151,1437,1622,2039]" pageId="13" pageNumber="356">
Female
<typeStatus id="367504E3DA13783D2001866CADB3F8F6" box="[290,397,1800,1826]" pageId="13" pageNumber="356" type="paratype">paratypes</typeStatus>
are black on the
<emphasis id="DBBA6653DA13783D2375866CAEEDF8F6" box="[598,723,1800,1826]" italics="true" pageId="13" pageNumber="356">dorsal side</emphasis>
with orange markings as follows:
<emphasis id="DBBA6653DA13783D2542866CA8F3F8F5" box="[1121,1229,1800,1825]" italics="true" pageId="13" pageNumber="356">Forewing</emphasis>
: (a) a streak from base through the discal cell up to the large cell-end spot from which it is separated by a narrow black line (more prominent in the
<typeStatus id="367504E3DA13783D207E862AAD85F8BC" box="[349,443,1870,1896]" pageId="13" pageNumber="356" type="allotype">allotype</typeStatus>
), with a smaller spot beyond the discal cell (this spot is more elongated with a narrow point in IBC-AY231), (b) on both wings, post-discal and subapical bands similar to the males but the bands are much broader and pale yellowish-orange in colour (rather than narrower and white or dark orange as in males). Females have a single narrow white band on the first abdominal segment (two broader, more prominent white bands in males).
</paragraph>
<caption id="BDB1EAC9DA10783E21B48678AE93F88C" ID-DOI="http://doi.org/10.5281/zenodo.14386318" ID-Zenodo-Dep="14386318" httpUri="https://zenodo.org/record/14386318/files/figure.png" pageId="14" pageNumber="357" startId="14.[151,250,1820,1844]" targetBox="[282,1304,194,1783]" targetPageId="14" targetType="figure">
<paragraph id="E971BA41DA10783E21B48678AE93F88C" blockId="14.[151,1436,1820,1880]" pageId="14" pageNumber="357">
<emphasis id="DBBA6653DA10783E21B48678AD2FF8E1" bold="true" box="[151,273,1820,1845]" pageId="14" pageNumber="357">FIGURE 7.</emphasis>
Type specimens of
<emphasis id="DBBA6653DA10783E20F78678AF6CF8E1" bold="true" box="[468,850,1820,1845]" pageId="14" pageNumber="357">
<taxonomicName id="2ECEC1C2DA10783E20F78678AECBF8E0" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[468,757,1820,1844]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="14" pageNumber="357" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA10783E20F78678AECBF8E0" bold="true" box="[468,757,1820,1844]" italics="true" pageId="14" pageNumber="357">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA10783E23DA8679AF6CF8E1" box="[761,850,1821,1845]" pageId="14" pageNumber="357" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte.
</paragraph>
</caption>
<caption id="BDB1EAC9DA11783F21B48447AD53FA8B" ID-DOI="http://doi.org/10.5281/zenodo.14386320" ID-Zenodo-Dep="14386320" httpUri="https://zenodo.org/record/14386320/files/figure.png" pageId="15" pageNumber="358" startId="15.[151,250,1315,1339]" targetBox="[171,1415,181,1291]" targetPageId="15" targetType="figure">
<paragraph id="E971BA41DA11783F21B48447AD53FA8B" blockId="15.[151,1437,1315,1375]" pageId="15" pageNumber="358">
<emphasis id="DBBA6653DA11783F21B48447AD2BFAEF" bold="true" box="[151,277,1315,1339]" pageId="15" pageNumber="358">FIGURE 8.</emphasis>
Male genitalia of
<emphasis id="DBBA6653DA11783F20FB8447AF59FAEF" bold="true" box="[472,871,1315,1339]" pageId="15" pageNumber="358">
<taxonomicName id="2ECEC1C2DA11783F20FB8447AF3FFAEF" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[472,769,1315,1339]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA11783F20FB8447AF3FFAEF" bold="true" box="[472,769,1315,1339]" italics="true" pageId="15" pageNumber="358">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA11783F22298447AF59FAEF" box="[778,871,1315,1339]" pageId="15" pageNumber="358" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
(holotype, IBC-PW982) and
<emphasis id="DBBA6653DA11783F25BF8447A936FAEF" box="[1180,1288,1315,1339]" italics="true" pageId="15" pageNumber="358">
A. i.
<taxonomicName id="2ECEC1C2DA11783F25F28440A936FAEF" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1233,1288,1316,1339]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
. Photographs: Dipendra Nath Basu.
</paragraph>
</caption>
<paragraph id="E971BA41DA11783F21E484C0AF7DF99A" blockId="15.[151,1437,1444,2011]" pageId="15" pageNumber="358">
<emphasis id="DBBA6653DA11783F21E484C0AD6AFA6A" box="[199,340,1444,1470]" italics="true" pageId="15" pageNumber="358">Ventral side:</emphasis>
Yellowish-orange with paler bands that correspond to the bands on dorsal side, with black smudges in the areas adjacent to the bands. IBC-PS983 is an intermediate seasonal form in which the bands barely stand out on the wing background, and UNH black discal spots are small. In the true dry season form, the bands are even more inconspicuous, and the black post-discal spots are absent. IBC-AY231 is a wet season form in which the pale bands are prominent, and the black post-discal spots are prominent.
</paragraph>
<paragraph id="E971BA41DA11783F21E4873CADA1F90A" blockId="15.[151,1437,1444,2011]" pageId="15" pageNumber="358">
<emphasis id="DBBA6653DA11783F21E4873CAD9AF9A6" bold="true" box="[199,420,1624,1650]" pageId="15" pageNumber="358">
Genitalia (
<figureCitation id="71F5A6C4DA11783F206A873CADAAF9A6" box="[329,404,1624,1650]" captionStart="FIGURE 8" captionStartId="15.[151,250,1315,1339]" captionTargetBox="[171,1415,181,1291]" captionTargetId="figure-19@15.[171,1416,181,1291]" captionTargetPageId="15" captionText="FIGURE 8. Male genitalia of Athyma inara sahyadriensis ssp. nov. (holotype, IBC-PW982) and A. i. inara. Photographs: Dipendra Nath Basu." figureDoi="http://doi.org/10.5281/zenodo.14386320" httpUri="https://zenodo.org/record/14386320/files/figure.png" pageId="15" pageNumber="358">Fig. 8</figureCitation>
):
</emphasis>
<emphasis id="DBBA6653DA11783F208C873CADD7F9A6" box="[431,489,1624,1650]" italics="true" pageId="15" pageNumber="358">Male</emphasis>
(IBC-PW982): Very similar to the genitalia of
<emphasis id="DBBA6653DA11783F2538873CA8A8F9A6" box="[1051,1174,1624,1650]" italics="true" pageId="15" pageNumber="358">
A. i.
<taxonomicName id="2ECEC1C2DA11783F257A873DA8A8F9A6" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1113,1174,1625,1650]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
, from which it differs primarily in having a smaller costal process/style. Tegumen large, broad. Uncus straight, slightly curved down at the tip. Gnathos slightly narrower than that of
<emphasis id="DBBA6653DA11783F238A87C4AF23F96E" box="[681,797,1696,1722]" italics="true" pageId="15" pageNumber="358">
A. i.
<taxonomicName id="2ECEC1C2DA11783F23C387C5AF23F96E" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[736,797,1697,1722]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
. Valve narrow, long, pointed, hairy. Phallus short, stout, ending in a sharp point.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA11782121E4878CAEDEF860" lastPageId="17" lastPageNumber="360" pageId="15" pageNumber="358" type="diagnosis">
<paragraph id="E971BA41DA11782021E4878CAE18FD25" blockId="15.[151,1437,1444,2011]" lastBlockId="16.[151,1437,151,753]" lastPageId="16" lastPageNumber="359" pageId="15" pageNumber="358">
<emphasis id="DBBA6653DA11783F21E4878CADEFF8D6" bold="true" box="[199,465,1768,1794]" pageId="15" pageNumber="358">
Diagnosis (
<figureCitation id="71F5A6C4DA11783F206E878CADFDF8D6" box="[333,451,1768,1794]" captionStart-0="FIGURE 9" captionStart-1="FIGURE 10" captionStartId-0="16.[151,250,1981,2005]" captionStartId-1="17.[151,250,1191,1215]" captionTargetBox-0="[168,1415,786,1947]" captionTargetBox-1="[156,1434,187,1159]" captionTargetId-0="figure-368@16.[164,1423,777,1958]" captionTargetId-1="figure-19@17.[151,1436,181,1167]" captionTargetPageId-0="16" captionTargetPageId-1="17" captionText-0="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." captionText-1="FIGURE 10. Diagnostic characters of Athyma inara sahyadriensis ssp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the subspecies. Male of A. i. sahyadriensis ssp. nov. is compared with males of A. i. inara and two subspecies of A. nefte. Numbers point to distinctive characters: 1: FW subapical markings, 2: FW submarginal markings, 3: FW spot in interspace 3, 4: FW markings in the discal cell, 5: HW post-discal band, 6: UNF markings in the discal cell, 7: UN base colour of the wings, 8: UNF submarginal markings, 9: large white spot in interspace 2, 10: UNH post-discal band (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte." figureDoi-0="http://doi.org/10.5281/zenodo.14386322" figureDoi-1="http://doi.org/10.5281/zenodo.14386326" httpUri-0="https://zenodo.org/record/14386322/files/figure.png" httpUri-1="https://zenodo.org/record/14386326/files/figure.png" pageId="15" pageNumber="358">Fig. 9–10</figureCitation>
):
</emphasis>
The diagnosis of this subspecies was briefly outlined by
<bibRefCitation id="8D5FC7B0DA11783F2551878CA96CF8D6" author="de Niceville, L." box="[1138,1362,1768,1794]" pageId="15" pageNumber="358" refId="ref14462" refString="de Niceville, L. (1886) The Butterflies of India, Burmah and Ceylon. Volume II. Nymphalidae, Nymphalinae. Lemoniidae, Libythaeinae, Nemeobiinae. The Calcutta Central Press Co., Kolkata, 332 pp." type="book" year="1886">de Nicéville (1886)</bibRefCitation>
under
<taxonomicName id="2ECEC1C2DA11783F21B48668AD79F8F2" baseAuthorityName="Evans" baseAuthorityYear="1932" box="[151,327,1804,1830]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="inarina">
<emphasis id="DBBA6653DA11783F21B48668AD79F8F2" box="[151,327,1804,1830]" italics="true" pageId="15" pageNumber="358">Athyma inarina</emphasis>
</taxonomicName>
based on a single male specimen from Wynaad (N
<collectingRegion id="2B0A74A3DA11783F22BB8668AFDAF8F2" box="[920,996,1804,1830]" country="India" name="Kerala" pageId="15" pageNumber="358">Kerala</collectingRegion>
) that was available to him at the time, although he did not taxonomically name it. A combination of 10 characters in male readily and consistently differentiates
<emphasis id="DBBA6653DA11783F20128630AE53F8BA" bold="true" box="[305,621,1875,1902]" pageId="15" pageNumber="358">
<emphasis id="DBBA6653DA11783F20128630AE3DF8BA" bold="true" box="[305,515,1875,1902]" italics="true" pageId="15" pageNumber="358">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
from related taxa (
<figureCitation id="71F5A6C4DA11783F22638630AFACF8BA" box="[832,914,1876,1902]" captionStart="FIGURE 10" captionStartId="17.[151,250,1191,1215]" captionTargetBox="[156,1434,187,1159]" captionTargetId="figure-19@17.[151,1436,181,1167]" captionTargetPageId="17" captionText="FIGURE 10. Diagnostic characters of Athyma inara sahyadriensis ssp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the subspecies. Male of A. i. sahyadriensis ssp. nov. is compared with males of A. i. inara and two subspecies of A. nefte. Numbers point to distinctive characters: 1: FW subapical markings, 2: FW submarginal markings, 3: FW spot in interspace 3, 4: FW markings in the discal cell, 5: HW post-discal band, 6: UNF markings in the discal cell, 7: UN base colour of the wings, 8: UNF submarginal markings, 9: large white spot in interspace 2, 10: UNH post-discal band (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386326" httpUri="https://zenodo.org/record/14386326/files/figure.png" pageId="15" pageNumber="358">Fig. 10</figureCitation>
, in which numbered pointers correspond to the following numbered characters): (1) UPF prominent subapical orange markings, which distinguish it from
<taxonomicName id="2ECEC1C2DA11783F241C861CA9AAF846" box="[1343,1428,1912,1938]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA11783F241C861CA9AAF846" box="[1343,1428,1912,1938]" italics="true" pageId="15" pageNumber="358">A. nefte</emphasis>
</taxonomicName>
; (2) UPF no orange or white markings below the subapical orange markings, only two inconspicuous, diffused grey lines (orange line or spots prominent in
<emphasis id="DBBA6653DA11783F234786A4AEE4F80E" box="[612,730,1984,2010]" italics="true" pageId="15" pageNumber="358">
A. i.
<taxonomicName id="2ECEC1C2DA11783F23BE86A5AEE4F80E" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[669,730,1985,2010]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="15" pageNumber="358" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
, especially with a larger spot in interspace 3; lines prominent grey or white in
<taxonomicName id="2ECEC1C2DA0E7820207881F3AD8DFF65" box="[347,435,151,177]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E7820207881F3AD8DFF65" box="[347,435,151,177]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (3) UPF white post-discal spot in 3 smaller, less conspicuous (this spot is larger and more conspicuous in
<emphasis id="DBBA6653DA0E782020A681DFADCBFF00" box="[389,501,187,213]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820209B81DFADCBFF00" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[440,501,187,212]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
, absent or almost absent in
<taxonomicName id="2ECEC1C2DA0E7820220981DFAF41FF01" box="[810,895,187,213]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E7820220981DFAF41FF01" box="[810,895,187,213]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (4) UPF white line through the discal cell very narrow, inconspicuous (sometimes entirely absent), not connecting with the large white spot at the cell-end (the line usually more prominent and longer, powdery-grey or white, connecting with the white cell-end spot in
<emphasis id="DBBA6653DA0E782024018067A9ABFEC8" box="[1314,1429,259,285]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820247B8067A9ABFEC8" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1368,1429,259,284]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; very prominent, white and connecting with the much larger and more prominent cell-end spot in
<taxonomicName id="2ECEC1C2DA0E782025F38043A91BFE95" box="[1232,1317,295,321]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E782025F38043A91BFE95" box="[1232,1317,295,321]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (5) UPH post-discal band narrow with even edges, dark orange (this band much broader, paler orange, with the inner edges of the spots that comprise this band being prominently convex in
<emphasis id="DBBA6653DA0E782022A7800BAFC6FE5C" box="[900,1016,367,393]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E78202298800BAFC6FE5C" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[955,1016,367,392]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; band white and narrow in
<taxonomicName id="2ECEC1C2DA0E78202417800BA9B5FE5D" box="[1332,1419,367,393]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E78202417800BA9B5FE5D" box="[1332,1419,367,393]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (6) UNF streak through the cell grey or white, more prominent than on dorsal side (this streak similar to
<emphasis id="DBBA6653DA0E7820240780F7A9ABFE78" box="[1316,1429,403,429]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820247B80F7A9ABFE78" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1368,1429,403,428]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; it is much more white, broader and conspicuous in
<taxonomicName id="2ECEC1C2DA0E782023FF80D3AF0CFE05" box="[732,818,439,465]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E782023FF80D3AF0CFE05" box="[732,818,439,465]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (7) UN wing colour orange (colour orange-brown in
<emphasis id="DBBA6653DA0E782021B480BFAD32FE20" box="[151,268,475,501]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E782021EC80BFAD32FE20" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[207,268,475,500]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; brown without a trace of orange in
<taxonomicName id="2ECEC1C2DA0E7820238D80BFAF3BFE21" box="[686,773,475,501]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E7820238D80BFAF3BFE21" box="[686,773,475,501]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (8) UNF submarginal markings grey and inconspicuous, or absent (these markings especially prominent in interspaces
<quantity id="2E3617A4DA0E7820227B809BAF9EFDCD" box="[856,928,511,537]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" metricValueMax="7.62" metricValueMin="2.54" pageId="16" pageNumber="359" unit="in" value="2.0" valueMax="3.0" valueMin="1.0">1–3 in</quantity>
<emphasis id="DBBA6653DA0E7820228B809BA822FDCC" box="[936,1052,511,537]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E782022FC809BA822FDCC" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[991,1052,511,536]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; and narrower but from
<quantity id="2E3617A4DA0E78202417809BA942FDCD" box="[1332,1404,511,537]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" metricValueMax="1.778" metricValueMin="0.254" pageId="16" pageNumber="359" unit="in" value="4.0" valueMax="7.0" valueMin="1.0">1–7 in</quantity>
<taxonomicName id="2ECEC1C2DA0E782024A7809BACF3FDE9" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E782024A7809BACF3FDE9" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (9) UNF outer edge of the large white spot in 2 protrudes out from the spot in 1b but it protrudes less than the width of the two grey submarginal lines/small spots (this spot similar in
<emphasis id="DBBA6653DA0E782025208323A847FDB4" box="[1027,1145,583,609]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820251F8323A847FDB4" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1084,1145,583,608]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; it protrudes much more, i.e., more than the width of the submarginal white lines in
<taxonomicName id="2ECEC1C2DA0E78202214830FAFB0FD51" box="[823,910,619,645]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E78202214830FAFB0FD51" box="[823,910,619,645]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
); (10) UNH narrow, grey or white post-discal band ends in interspace 4 (this band is broader and it extends up to the apex in
<emphasis id="DBBA6653DA0E7820253683EBA8B6FD7C" box="[1045,1160,655,681]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820256883EBA8B6FD7C" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1099,1160,655,680]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; it is much much whiter but narrower in
<taxonomicName id="2ECEC1C2DA0E7820206983D7ADA0FD19" box="[330,414,691,717]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E7820206983D7ADA0FD19" box="[330,414,691,717]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
). Additionally,
<emphasis id="DBBA6653DA0E7820236D83D7AFB5FD19" bold="true" box="[590,907,690,717]" pageId="16" pageNumber="359">
<emphasis id="DBBA6653DA0E7820236D83D7AF1EFD19" bold="true" box="[590,800,690,717]" italics="true" pageId="16" pageNumber="359">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
differs from
<emphasis id="DBBA6653DA0E7820253D83D7A8B1FD18" box="[1054,1167,691,717]" italics="true" pageId="16" pageNumber="359">
A. i.
<taxonomicName id="2ECEC1C2DA0E7820257183D7A8B1FD18" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1106,1167,691,716]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
and
<taxonomicName id="2ECEC1C2DA0E782025E683D7A927FD19" box="[1221,1305,691,717]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0E782025E683D7A927FD19" box="[1221,1305,691,717]" italics="true" pageId="16" pageNumber="359">A. nefte</emphasis>
</taxonomicName>
in having a narrower, more produced forewing.
</paragraph>
<caption id="BDB1EAC9DA0E782021B486D9A81CF82D" ID-DOI="http://doi.org/10.5281/zenodo.14386322" ID-Zenodo-Dep="14386322" httpUri="https://zenodo.org/record/14386322/files/figure.png" pageId="16" pageNumber="359" startId="16.[151,250,1981,2005]" targetBox="[168,1415,786,1947]" targetPageId="16" targetType="figure">
<paragraph id="E971BA41DA0E782021B486D9A81CF82D" blockId="16.[151,1437,1981,2042]" pageId="16" pageNumber="359">
<emphasis id="DBBA6653DA0E782021B486D9AD2AF801" bold="true" box="[151,276,1981,2005]" pageId="16" pageNumber="359">FIGURE 9.</emphasis>
Comparison of
<emphasis id="DBBA6653DA0E7820209986D9AF7DF801" bold="true" box="[442,835,1981,2005]" pageId="16" pageNumber="359">
<taxonomicName id="2ECEC1C2DA0E7820209986D9AEDFF801" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[442,737,1981,2005]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0E7820209986D9AEDFF801" bold="true" box="[442,737,1981,2005]" italics="true" pageId="16" pageNumber="359">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0E782023CB86D9AF7DF801" box="[744,835,1981,2005]" pageId="16" pageNumber="359" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
with other similar
<taxonomicName id="2ECEC1C2DA0E7820252B86D9A86BF801" box="[1032,1109,1981,2005]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="359" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA0E7820252B86D9A86BF801" box="[1032,1109,1981,2005]" italics="true" pageId="16" pageNumber="359">Athyma</emphasis>
</taxonomicName>
(see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte.
</paragraph>
</caption>
<caption id="BDB1EAC9DA0F782121B485C3A90FFAA7" ID-DOI="http://doi.org/10.5281/zenodo.14386326" ID-Zenodo-Dep="14386326" httpUri="https://zenodo.org/record/14386326/files/figure.png" pageId="17" pageNumber="360" startId="17.[151,250,1191,1215]" targetBox="[156,1434,187,1159]" targetPageId="17" targetType="figure">
<paragraph id="E971BA41DA0F782121B485C3A90FFAA7" blockId="17.[151,1437,1191,1396]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F782121B485C3AD21FB6B" bold="true" box="[151,287,1191,1215]" pageId="17" pageNumber="360">FIGURE 10.</emphasis>
Diagnostic characters of
<emphasis id="DBBA6653DA0F7821233D85C3AFA1FB14" bold="true" box="[542,927,1191,1216]" pageId="17" pageNumber="360">
<taxonomicName id="2ECEC1C2DA0F7821233D85C3AF7EFB6B" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[542,832,1191,1215]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0F7821233D85C3AF7EFB6B" bold="true" box="[542,832,1191,1215]" italics="true" pageId="17" pageNumber="360">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0F7821226585CCAFA1FB14" box="[838,927,1192,1216]" pageId="17" pageNumber="360" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
, marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the subspecies. Male of
<emphasis id="DBBA6653DA0F7821254385AFA9BAFB30" bold="true" box="[1120,1412,1227,1252]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F7821254385AFA91FFB37" bold="true" box="[1120,1313,1227,1251]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
is compared with males of
<emphasis id="DBBA6653DA0F782120B48594AE3FFAD3" box="[407,513,1264,1287]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F782120E98594AE3FFAD3" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[458,513,1264,1287]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
and two subspecies of
<taxonomicName id="2ECEC1C2DA0F782123D78594AF7DFADC" box="[756,835,1264,1288]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F782123D78594AF7DFADC" box="[756,835,1264,1288]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
. Numbers point to distinctive characters: 1: FW subapical markings, 2: FW submarginal markings, 3: FW spot in interspace 3, 4: FW markings in the discal cell, 5: HW post-discal band, 6: UNF markings in the discal cell, 7: UN base colour of the wings, 8: UNF submarginal markings, 9: large white spot in interspace 2, 10: UNH post-discal band (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte.
</paragraph>
</caption>
<paragraph id="E971BA41DA0F782121E484A2AE04F89C" blockId="17.[151,1437,1478,1972]" pageId="17" pageNumber="360">
<taxonomicName id="2ECEC1C2DA0F782121E484A2AD64FA0B" authorityName="Moore" authorityYear="1858" box="[199,346,1478,1504]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="asita">
<emphasis id="DBBA6653DA0F782121E484A2AD64FA0B" box="[199,346,1478,1504]" italics="true" pageId="17" pageNumber="360">Athyma asita</emphasis>
</taxonomicName>
, whose taxonomic status needs to be carefully examined with a separate molecular phylogenetic study (see under Discussion), differs from
<emphasis id="DBBA6653DA0F782123A4848EAFEFF9D0" bold="true" box="[647,977,1513,1540]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F782123A4848EAF5EF9D0" bold="true" box="[647,864,1513,1540]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
,
<emphasis id="DBBA6653DA0F782122C1848EA864F9D7" box="[994,1114,1514,1540]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F7821253E848EA864F9D7" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1053,1114,1514,1539]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
and
<taxonomicName id="2ECEC1C2DA0F782125B5848EA8D0F9D0" box="[1174,1262,1514,1540]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F782125B5848EA8D0F9D0" box="[1174,1262,1514,1540]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
, in having: (a) UPF submarginal markings prominent and white, especially the large spot in interspace 3 (this large spot and other submarginal markings are orange in
<emphasis id="DBBA6653DA0F7821231E8756AE8AF99F" box="[573,692,1586,1612]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F782123548756AE8AF99F" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[631,692,1586,1611]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; they are white or pale grey, much narrower but prominent, and without a large spot in interspace 3, in
<taxonomicName id="2ECEC1C2DA0F782123618732AEA8F9A4" box="[578,662,1622,1648]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F782123618732AEA8F9A4" box="[578,662,1622,1648]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
; they are absent in
<emphasis id="DBBA6653DA0F7821224F8732A896F9A4" bold="true" box="[876,1192,1621,1648]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F7821224F8732A800F9A4" bold="true" box="[876,1086,1621,1648]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
), (b) UPH post-discal band narrow, white (orange in
<emphasis id="DBBA6653DA0F782120D0871EAE59F947" box="[499,615,1658,1684]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F78212309871EAE59F947" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[554,615,1658,1683]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
and
<emphasis id="DBBA6653DA0F782123BD871EAFDFF940" bold="true" box="[670,993,1657,1684]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F782123BD871EAF4AF940" bold="true" box="[670,884,1657,1684]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
; similar in
<taxonomicName id="2ECEC1C2DA0F78212547871EA884F940" box="[1124,1210,1658,1684]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F78212547871EA884F940" box="[1124,1210,1658,1684]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
), (c) FW discal cell with a well-defined white line that is well-separated from the white cell-end spot, with another white spot beyond the cell-end that is almost as large (these markings are highly reduced in
<emphasis id="DBBA6653DA0F782122E687A6A93DF908" bold="true" box="[965,1283,1729,1756]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F782122E687A6A8A6F908" bold="true" box="[965,1176,1729,1756]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
; conspicuous but diffused in
<emphasis id="DBBA6653DA0F782120678782AD89F92B" box="[324,439,1766,1792]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F782120598782AD89F92B" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[378,439,1766,1791]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; much more prominent with a much larger cell-end spot in
<taxonomicName id="2ECEC1C2DA0F7821257E8782A88DF8D4" box="[1117,1203,1766,1792]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F7821257E8782A88DF8D4" box="[1117,1203,1766,1792]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
), and (d) UN brown with a slight tinge of orange (brighter orange-brown in
<emphasis id="DBBA6653DA0F7821222A866EA875F8F0" bold="true" box="[777,1099,1801,1828]" pageId="17" pageNumber="360">
<emphasis id="DBBA6653DA0F7821222A866EAFE0F8F0" bold="true" box="[777,990,1801,1828]" italics="true" pageId="17" pageNumber="360">A. i. sahyadriensis</emphasis>
ssp. nov.
</emphasis>
and
<emphasis id="DBBA6653DA0F782125A0866EA8C8F8F7" box="[1155,1270,1802,1828]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F7821259A866EA8C8F8F7" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1209,1270,1802,1827]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
; darker brown without a tinge of orange in
<taxonomicName id="2ECEC1C2DA0F782120F7864AAE14F89C" box="[468,554,1838,1864]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F782120F7864AAE14F89C" box="[468,554,1838,1864]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="E971BA41DA0F782121E48636AEDEF860" blockId="17.[151,1437,1478,1972]" pageId="17" pageNumber="360">
Females are similar to
<emphasis id="DBBA6653DA0F782120E28636AE0EF8BF" box="[449,560,1874,1900]" italics="true" pageId="17" pageNumber="360">
A. i.
<taxonomicName id="2ECEC1C2DA0F782120D08636AE0EF8BF" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[499,560,1874,1899]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
, from which they apparently cannot be distinguished. They differ from females of
<taxonomicName id="2ECEC1C2DA0F782121978612AD36F844" box="[180,264,1910,1936]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="17" pageNumber="360" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0F782121978612AD36F844" box="[180,264,1910,1936]" italics="true" pageId="17" pageNumber="360">A. nefte</emphasis>
</taxonomicName>
in having considerably broader orange markings UP, and brighter orange UN (UP markings narrower, UN background colour of the wings brown, not orange).
</paragraph>
</subSubSection>
<caption id="BDB1EAC9DA0C782221B48450A9B1FA09" ID-DOI="http://doi.org/10.5281/zenodo.14386330" ID-Zenodo-Dep="14386330" httpUri="https://zenodo.org/record/14386330/files/figure.png" pageId="18" pageNumber="361" startId="18.[151,250,1332,1356]" targetBox="[159,1428,181,1309]" targetPageId="18" targetType="figure">
<paragraph id="E971BA41DA0C782221B48450A9B1FA09" blockId="18.[151,1436,1332,1501]" pageId="18" pageNumber="361">
<emphasis id="DBBA6653DA0C782221B48450AD20FA99" bold="true" box="[151,286,1332,1357]" pageId="18" pageNumber="361">FIGURE 11.</emphasis>
Early stages of
<emphasis id="DBBA6653DA0C782220E38450AF5EFA99" bold="true" box="[448,864,1332,1357]" pageId="18" pageNumber="361">
<taxonomicName id="2ECEC1C2DA0C782220E38450AEDAFA98" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[448,740,1332,1357]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="361" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0C782220E38450AEDAFA98" bold="true" box="[448,740,1332,1357]" italics="true" pageId="18" pageNumber="361">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0C782223C98451AF7AFA99" box="[746,836,1333,1357]" pageId="18" pageNumber="361" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
a:
</emphasis>
egg (2010/11/05);
<emphasis id="DBBA6653DA0C782225038451A809FA99" bold="true" box="[1056,1079,1333,1357]" pageId="18" pageNumber="361">b:</emphasis>
1
<superScript id="1EBB1709DA0C7822256A8450A86DFA96" attach="left" box="[1097,1107,1332,1346]" fontSize="6" pageId="18" pageNumber="361">st</superScript>
instar (2010/09/20);
<emphasis id="DBBA6653DA0C7822240A8451A903FA99" bold="true" box="[1321,1341,1333,1357]" pageId="18" pageNumber="361">c:</emphasis>
2
<superScript id="1EBB1709DA0C7822246C8451A960FA97" attach="left" box="[1359,1374,1333,1347]" fontSize="6" pageId="18" pageNumber="361">nd</superScript>
instar (2010/09/26);
<emphasis id="DBBA6653DA0C7822200F843DAD64FAA5" bold="true" box="[300,346,1369,1393]" pageId="18" pageNumber="361">d–e:</emphasis>
3
<superScript id="1EBB1709DA0C78222053843DAD42FAB3" attach="left" box="[368,380,1369,1383]" fontSize="6" pageId="18" pageNumber="361">rd</superScript>
instar (2010/10/02);
<emphasis id="DBBA6653DA0C78222378843CAEBEFAA4" bold="true" box="[603,640,1368,1392]" pageId="18" pageNumber="361">f–i:</emphasis>
4
<superScript id="1EBB1709DA0C782223B5843DAE9FFAB3" attach="left" box="[662,673,1369,1383]" fontSize="6" pageId="18" pageNumber="361">th</superScript>
instar (2010/10/05–07);
<emphasis id="DBBA6653DA0C78222285843CAFF5FAA4" bold="true" box="[934,971,1368,1392]" pageId="18" pageNumber="361">j–l:</emphasis>
5
<superScript id="1EBB1709DA0C782222C2843DAFD2FAB3" attach="left" box="[993,1004,1369,1383]" fontSize="6" pageId="18" pageNumber="361">th</superScript>
instar (2010/10/08);
<emphasis id="DBBA6653DA0C782225E8843DA93DFAA5" bold="true" box="[1227,1283,1369,1393]" pageId="18" pageNumber="361">m–n:</emphasis>
5
<superScript id="1EBB1709DA0C7822243A843DA91AFAB3" attach="left" box="[1305,1316,1369,1383]" fontSize="6" pageId="18" pageNumber="361">th</superScript>
instar with colours changed just before pupation (2010/10/10).
<emphasis id="DBBA6653DA0C7822238B8419AEE6FA41" bold="true" box="[680,728,1405,1429]" pageId="18" pageNumber="361">o–q:</emphasis>
newly-formed pupa (o: lateral view; p: dorsal view; q: ventral view);
<emphasis id="DBBA6653DA0C782221B484C5ACFFFA6D" bold="true" box="[151,193,1441,1465]" pageId="18" pageNumber="361">r–s:</emphasis>
pupa before eclosion (r: lateral view; s: dorsal view);
<emphasis id="DBBA6653DA0C782223C484C5AEC6FA6D" bold="true" box="[743,760,1441,1465]" pageId="18" pageNumber="361">t:</emphasis>
eclosed female resting on pupal case, drying its wings. Images are of multiple individuals. Amboli, Sindhudurg District, Maharashtra, India. Photographs: Hemant Ogale, used with permission.
</paragraph>
</caption>
<subSubSection id="A1D4E9CADA0C782221E4874BAFB2F9B9" pageId="18" pageNumber="361" type="etymology">
<paragraph id="E971BA41DA0C782221E4874BAFB2F9B9" blockId="18.[151,1437,1582,2042]" pageId="18" pageNumber="361">
<emphasis id="DBBA6653DA0C782221E4874BAD6CF99D" bold="true" box="[199,338,1583,1609]" pageId="18" pageNumber="361">Etymology:</emphasis>
We name
<emphasis id="DBBA6653DA0C782220F6874AAFABF99D" bold="true" box="[469,917,1582,1609]" pageId="18" pageNumber="361">
<taxonomicName id="2ECEC1C2DA0C782220F6874AAF1FF99D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[469,801,1582,1609]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="361" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0C782220F6874AAF1FF99D" bold="true" box="[469,801,1582,1609]" italics="true" pageId="18" pageNumber="361">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0C7822220F874BAFABF99D" box="[812,917,1583,1609]" pageId="18" pageNumber="361" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
after its endemic distributional range in the Sahyadri, the name of the Western Ghats in many native languages.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA0C782321E48713AE18FA0C" lastPageId="19" lastPageNumber="362" pageId="18" pageNumber="361" type="discussion">
<paragraph id="E971BA41DA0C782221E48713AD73F8BD" blockId="18.[151,1437,1582,2042]" pageId="18" pageNumber="361">
<emphasis id="DBBA6653DA0C782221E48713AD8CF945" bold="true" box="[199,434,1655,1681]" pageId="18" pageNumber="361">Historical Records:</emphasis>
This taxon is well-documented in the Western Ghats based on hundreds of records (also see
<figureCitation id="71F5A6C4DA0C782221E787FFAD26F961" box="[196,280,1691,1717]" captionStart="FIGURE 12" captionStartId="19.[151,250,1328,1352]" captionTargetBox="[151,1436,237,1304]" captionTargetId="figure-49@19.[151,1436,237,1304]" captionTargetPageId="19" captionText="FIGURE 12. Distributional range map of relevant taxa from the Athyma nefte species group. Pink: A. inara sahyadriensis ssp. nov., blue: A. i. inara, Purple: A. asita, Yellow: A. nefte. Map: Viraj Nawge." figureDoi="http://doi.org/10.5281/zenodo.14386332" httpUri="https://zenodo.org/record/14386332/files/figure.png" pageId="18" pageNumber="361">Fig. 12</figureCitation>
).
<bibRefCitation id="8D5FC7B0DA0C7822200D87FFADF9F961" author="Evans, W. H." box="[302,455,1691,1717]" pageId="18" pageNumber="361" refId="ref13172" refString="Evans, W. H. (1932) The Identification of Indian Butterflies (2 nd edn.). Bombay Natural History Society, Mumbai, 454 pp." type="book" year="1932">Evans (1932)</bibRefCitation>
used the name
<taxonomicName id="2ECEC1C2DA0C7822235E87F8AFBDF961" box="[637,899,1691,1717]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="361" phylum="Arthropoda" rank="subSpecies" species="nefte" subSpecies="inara">
<emphasis id="DBBA6653DA0C7822235E87F8AFBDF961" box="[637,899,1691,1717]" italics="true" pageId="18" pageNumber="361">Pantoporia nefte inara</emphasis>
</taxonomicName>
, recording it as “S.
<collectingCountry id="91D9FAD1DA0C7822254F87FFA897F961" box="[1132,1193,1691,1717]" name="India" pageId="18" pageNumber="361">India</collectingCountry>
,
<collectingRegion id="2B0A74A3DA0C7822259587FFA934F961" box="[1206,1290,1691,1717]" country="India" name="Sikkim" pageId="18" pageNumber="361">Sikkim</collectingRegion>
—N.
<collectingCountry id="91D9FAD1DA0C7822246B87FFA9A7F961" box="[1352,1433,1691,1717]" name="Myanmar" pageId="18" pageNumber="361">Burma</collectingCountry>
. NR [Not Rare].” According to
<bibRefCitation id="8D5FC7B0DA0C782220D587DBAED8F90E" author="Wynter-Blyth, M. A." box="[502,742,1727,1754]" pageId="18" pageNumber="361" refId="ref15140" refString="Wynter-Blyth, M. A. (1957) Butterflies of the Indian Region. Bombay Natural History Society, Mumbai, 523 pp." type="book" year="1957">Wynter-Blyth (1957)</bibRefCitation>
, who referred to it as
<taxonomicName id="2ECEC1C2DA0C782222C387A4A8A1F90D" box="[992,1183,1727,1753]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="361" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0C782222C387A4A8A1F90D" box="[992,1183,1727,1753]" italics="true" pageId="18" pageNumber="361">Pantoporia nefte</emphasis>
</taxonomicName>
, “
<emphasis id="DBBA6653DA0C7822259887A4A934F90D" box="[1211,1290,1728,1753]" italics="true" pageId="18" pageNumber="361">Range.</emphasis>
South
<collectingCountry id="91D9FAD1DA0C7822247887DBA9A7F90D" box="[1371,1433,1727,1753]" name="India" pageId="18" pageNumber="361">India</collectingCountry>
.
<collectingRegion id="2B0A74A3DA0C782221B48787ACD7F929" box="[151,233,1763,1789]" country="India" name="Sikkim" pageId="18" pageNumber="361">Sikkim</collectingRegion>
to
<collectingRegion id="2B0A74A3DA0C7822202E8787AD64F929" box="[269,346,1763,1789]" country="India" name="Assam" pageId="18" pageNumber="361">Assam</collectingRegion>
and South
<collectingCountry id="91D9FAD1DA0C782220FF8787AE13F929" box="[476,557,1763,1789]" name="Myanmar" pageId="18" pageNumber="361">Burma</collectingCountry>
. … Recorded as not rare in the evergreen regions of the Western Ghats, also, very rarely, from the Mettupalaiyam Ghat in the Nilgiris.”
<bibRefCitation id="8D5FC7B0DA0C7822220E8663AFF9F8F5" author="Talbot, G." box="[813,967,1799,1825]" pageId="18" pageNumber="361" refId="ref15075" refString="Talbot, G. (1947) The Fauna of British India, including Ceylon and Burma: Butterflies, Vol. 2. Taylor &amp; Francis, London, 506 pp." type="book" year="1947">Talbot (1947)</bibRefCitation>
did not cover the genus.
<bibRefCitation id="8D5FC7B0DA0C782225C78663A9A2F8F6" author="d'Abrera, B." box="[1252,1436,1799,1826]" pageId="18" pageNumber="361" refId="ref12980" refString="d'Abrera, B. (1985) Butterflies of the Oriental Region. Part II: Nymphalidae, Satyridae &amp; Amathusidae. Hill House, Melbourne, 534 pp." type="book" year="1985">d’Abrera (1985)</bibRefCitation>
mentioned “
<collectingCountry id="91D9FAD1DA0C78222000864FAD57F891" box="[291,361,1835,1861]" name="Nepal" pageId="18" pageNumber="361">Nepal</collectingCountry>
to
<collectingRegion id="2B0A74A3DA0C782220B0864FADDCF891" box="[403,482,1835,1861]" country="India" name="Assam" pageId="18" pageNumber="361">Assam</collectingRegion>
” as the range of
<taxonomicName id="2ECEC1C2DA0C78222390864FAF6BF891" box="[691,853,1835,1861]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="18" pageNumber="361" phylum="Arthropoda" rank="subSpecies" species="nefte" subSpecies="inara">
<emphasis id="DBBA6653DA0C78222390864FAF6BF891" box="[691,853,1835,1861]" italics="true" pageId="18" pageNumber="361">A. nefte inara</emphasis>
</taxonomicName>
, with range in the Western Ghats as doubtful, “? southern
<collectingCountry id="91D9FAD1DA0C78222023862BAD04F8BD" box="[256,314,1871,1897]" name="India" pageId="18" pageNumber="361">India</collectingCountry>
”.
</paragraph>
<paragraph id="E971BA41DA0C782321E48617A8AFFF01" blockId="18.[151,1437,1582,2042]" lastBlockId="19.[151,1436,151,213]" lastPageId="19" lastPageNumber="362" pageId="18" pageNumber="361">
The NHMUK has
<specimenCount id="FFC871C8DA0C782220B88617ADF8F859" box="[411,454,1907,1933]" count="8" pageId="18" pageNumber="361" type="male">8 ♂</specimenCount>
and
<specimenCount id="FFC871C8DA0C782220DC8617AE15F859" box="[511,555,1907,1933]" count="8" pageId="18" pageNumber="361" type="female">8 ♀</specimenCount>
specimens (four of each sex photographed and catalogued, see Materials and Methods) from Karwar, Karwar District,
<collectingRegion id="2B0A74A3DA0C7822234D86F3AEDAF865" box="[622,740,1943,1969]" country="India" name="Karnataka" pageId="18" pageNumber="361">Karnataka</collectingRegion>
, raised from early stages by T. R. Bell in 1920s, except:
<specimenCount id="FFC871C8DA0C7822245786F3ACF3F801" count="1" pageId="18" pageNumber="361" type="male">one male</specimenCount>
from “Malabar, Wynaad” from “Moore Coll. 99-234”,
<specimenCount id="FFC871C8DA0C7822226486DFAF90F801" box="[839,942,1979,2005]" count="1" pageId="18" pageNumber="361" type="male">one male</specimenCount>
from “Nilgiris 89-53
<quantity id="2E3617A4DA0C7822258986DFA939F801" box="[1194,1287,1979,2006]" metricMagnitude="3" metricUnit="m" metricValue="1.524" pageId="18" pageNumber="361" unit="ft" value="5000.0">5,000 ft</quantity>
”,
<specimenCount id="FFC871C8DA0C7822243C86DFA9A2F801" box="[1311,1436,1979,2005]" count="1" pageId="18" pageNumber="361" type="female">one female</specimenCount>
from “Nilgiris 89-53
<quantity id="2E3617A4DA0C782220B386BBADD0F82D" box="[400,494,2015,2041]" metricMagnitude="2" metricUnit="m" metricValue="9.144" pageId="18" pageNumber="361" unit="ft" value="3000.0">3,000 ft</quantity>
”, and
<specimenCount id="FFC871C8DA0C7822231986BBAE9DF82D" box="[570,675,2015,2041]" count="1" pageId="18" pageNumber="361" type="male">one male</specimenCount>
from “Watecolle [
<collectingRegion id="2B0A74A3DA0C7822225886BBAFF5F82D" box="[891,971,2015,2041]" country="India" name="Kerala" pageId="18" pageNumber="361">Kerala</collectingRegion>
] 16/11/60 [1960]” and
<specimenCount id="FFC871C8DA0C782225FE86BBA965F82D" box="[1245,1371,2015,2041]" count="1" pageId="18" pageNumber="361" type="female">one female</specimenCount>
from “Sollekolle 14/10/60” from A. J. Sharman Collections, which did not have voucher codes. MCZ and McGuire Centre for
<taxonomicName id="2ECEC1C2DA0D7823203381DFADA9FF01" box="[272,407,187,213]" class="Insecta" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
and Biodiversity do not have any specimens of the new subspecies.
</paragraph>
<caption id="BDB1EAC9DA0D782321B48454AF99FABF" ID-DOI="http://doi.org/10.5281/zenodo.14386332" ID-Zenodo-Dep="14386332" httpUri="https://zenodo.org/record/14386332/files/figure.png" pageId="19" pageNumber="362" startId="19.[151,250,1328,1352]" targetBox="[151,1436,237,1304]" targetPageId="19" targetType="figure">
<paragraph id="E971BA41DA0D782321B48454AF99FABF" blockId="19.[151,1437,1327,1388]" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321B48454AD21FA9C" bold="true" box="[151,287,1328,1352]" pageId="19" pageNumber="362">FIGURE 12.</emphasis>
Distributional range map of relevant taxa from the
<taxonomicName id="2ECEC1C2DA0D782322078454AF96FA9C" baseAuthorityName="Cramer" baseAuthorityYear="1780" box="[804,936,1328,1352]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0D782322078454AF96FA9C" box="[804,936,1328,1352]" italics="true" pageId="19" pageNumber="362">Athyma nefte</emphasis>
</taxonomicName>
species group.
<emphasis id="DBBA6653DA0D782325678454ACFDFABF" bold="true" pageId="19" pageNumber="362">
Pink:
<taxonomicName id="2ECEC1C2DA0D782325A68454A950FA93" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1157,1390,1327,1352]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="subSpecies" species="inara" subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0D782325A68454A950FA93" bold="true" box="[1157,1390,1327,1352]" italics="true" pageId="19" pageNumber="362">A. inara sahyadriensis</emphasis>
</taxonomicName>
ssp. nov.
</emphasis>
,
<emphasis id="DBBA6653DA0D782321F38430AD39FABF" bold="true" box="[208,263,1364,1387]" pageId="19" pageNumber="362">blue:</emphasis>
<emphasis id="DBBA6653DA0D7823202E8430AD09FAB8" box="[269,311,1364,1388]" italics="true" pageId="19" pageNumber="362">A. i.</emphasis>
<taxonomicName id="2ECEC1C2DA0D7823201E8431ADF0FABF" authority=", Purple" authorityName="Purple" box="[317,462,1364,1388]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0D7823201E8431AD4AFAB8" box="[317,372,1365,1388]" italics="true" pageId="19" pageNumber="362">inara</emphasis>
,
<emphasis id="DBBA6653DA0D782320A28430ADF0FABF" bold="true" box="[385,462,1364,1387]" pageId="19" pageNumber="362">Purple</emphasis>
</taxonomicName>
<emphasis id="DBBA6653DA0D782320ED8430ADEDFABF" bold="true" box="[462,467,1364,1387]" pageId="19" pageNumber="362">:</emphasis>
<taxonomicName id="2ECEC1C2DA0D782320FA8430AE41FABF" authority=", Yellow" authorityName="Yellow" box="[473,639,1364,1388]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="asita">
<emphasis id="DBBA6653DA0D782320FA8430AE18FAB8" box="[473,550,1364,1388]" italics="true" pageId="19" pageNumber="362">A. asita</emphasis>
,
<emphasis id="DBBA6653DA0D782323108430AE41FABF" bold="true" box="[563,639,1364,1387]" pageId="19" pageNumber="362">Yellow</emphasis>
</taxonomicName>
<emphasis id="DBBA6653DA0D7823235C8430AEBDFABF" bold="true" box="[639,643,1364,1387]" pageId="19" pageNumber="362">:</emphasis>
<taxonomicName id="2ECEC1C2DA0D782323AA8430AEE9FAB8" baseAuthorityName="Cramer" baseAuthorityYear="1780" box="[649,727,1364,1388]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0D782323AA8430AEE9FAB8" box="[649,727,1364,1388]" italics="true" pageId="19" pageNumber="362">A. nefte</emphasis>
</taxonomicName>
. Map: Viraj Nawge.
</paragraph>
</caption>
<paragraph id="E971BA41DA0D782321B484DAAE18FA0C" blockId="19.[151,550,1470,1496]" box="[151,550,1470,1496]" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321B484DAAE18FA0C" bold="true" box="[151,550,1470,1496]" pageId="19" pageNumber="362">Species Biology and other Notes</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA0D782321B48762AF80F900" pageId="19" pageNumber="362" type="distribution">
<paragraph id="E971BA41DA0D782321B48762AF80F900" blockId="19.[151,1437,1542,2037]" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321B48762AEC5F9F4" bold="true" box="[151,763,1542,1569]" pageId="19" pageNumber="362">
Distribution:
<taxonomicName id="2ECEC1C2DA0D7823201E8762AEB6F9F5" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[317,648,1542,1569]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0D7823201E8762AEB6F9F5" bold="true" box="[317,648,1542,1569]" italics="true" pageId="19" pageNumber="362">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0D782323B08762AEC5F9F4" box="[659,763,1542,1568]" pageId="19" pageNumber="362" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
is distributed almost throughout the Western Ghats, from Shendurney WLS in southern
<collectingRegion id="2B0A74A3DA0D782320CD874EAE07F990" box="[494,569,1578,1604]" country="India" name="Kerala" pageId="19" pageNumber="362">Kerala</collectingRegion>
to Phansad WLS, Raigad District (
<bibRefCitation id="8D5FC7B0DA0D782322EB874EA8CAF990" author="Bhopale, N. &amp; Athavale, S." box="[968,1268,1578,1604]" pageId="19" pageNumber="362" refId="ref12668" refString="Bhopale, N. &amp; Athavale, S. (2009) First record of the Colour Sergeant Athyma nefte in Phansad Wildlife Sanctuary in Raigad District, Maharashtra, India. Journal of the Bombay Natural History Society 106 (1), 117." type="journal volume" year="2009">Bhopale &amp; Athavale 2009</bibRefCitation>
), and Tamhini, Pune District, in northern
<collectingRegion id="2B0A74A3DA0D78232098872AAE79F9BC" box="[443,583,1614,1640]" country="India" name="Maharashtra" pageId="19" pageNumber="362">Maharashtra</collectingRegion>
(
<figureCitation id="71F5A6C4DA0D78232375872AAE94F9BC" box="[598,682,1614,1640]" captionStart="FIGURE 12" captionStartId="19.[151,250,1328,1352]" captionTargetBox="[151,1436,237,1304]" captionTargetId="figure-49@19.[151,1436,237,1304]" captionTargetPageId="19" captionText="FIGURE 12. Distributional range map of relevant taxa from the Athyma nefte species group. Pink: A. inara sahyadriensis ssp. nov., blue: A. i. inara, Purple: A. asita, Yellow: A. nefte. Map: Viraj Nawge." figureDoi="http://doi.org/10.5281/zenodo.14386332" httpUri="https://zenodo.org/record/14386332/files/figure.png" pageId="19" pageNumber="362">Fig. 12</figureCitation>
). In comparison,
<emphasis id="DBBA6653DA0D7823224E872AAFE1F9BC" box="[877,991,1614,1640]" italics="true" pageId="19" pageNumber="362">
A. i.
<taxonomicName id="2ECEC1C2DA0D78232281872BAFE1F9BC" box="[930,991,1615,1640]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="inara">inara</taxonomicName>
</emphasis>
occurs in NE
<collectingCountry id="91D9FAD1DA0D7823255C872AA887F9BC" box="[1151,1209,1614,1640]" name="India" pageId="19" pageNumber="362">India</collectingCountry>
and N
<collectingCountry id="91D9FAD1DA0D78232429872BA947F9BD" box="[1290,1401,1615,1641]" name="Myanmar" pageId="19" pageNumber="362">Myanmar</collectingCountry>
,
<taxonomicName id="2ECEC1C2DA0D782324A7872AACF3F958" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="asita">
<emphasis id="DBBA6653DA0D782324A7872AACF3F958" italics="true" pageId="19" pageNumber="362">A. asita</emphasis>
</taxonomicName>
(often treated as a subspecies of
<taxonomicName id="2ECEC1C2DA0D782323658716AEA2F958" baseAuthorityName="Cramer" baseAuthorityYear="1780" box="[582,668,1650,1676]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0D782323658716AEA2F958" box="[582,668,1650,1676]" italics="true" pageId="19" pageNumber="362">A. nefte</emphasis>
</taxonomicName>
, see Discussion) occurs throughout Indo-China (except S
<collectingCountry id="91D9FAD1DA0D7823240C8716A9AAF958" box="[1327,1428,1650,1676]" name="Thailand" pageId="19" pageNumber="362">Thailand</collectingCountry>
) and S
<collectingCountry id="91D9FAD1DA0D782321FF87F3AD75F965" box="[220,331,1687,1713]" name="Myanmar" pageId="19" pageNumber="362">Myanmar</collectingCountry>
, extending northward into S
<collectingCountry id="91D9FAD1DA0D782323A887F2AEEFF964" box="[651,721,1686,1712]" name="China" pageId="19" pageNumber="362">China</collectingCountry>
, and
<taxonomicName id="2ECEC1C2DA0D7823222987F2AF60F964" baseAuthorityName="Cramer" baseAuthorityYear="1780" box="[778,862,1686,1712]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="species" species="nefte">
<emphasis id="DBBA6653DA0D7823222987F2AF60F964" box="[778,862,1686,1712]" italics="true" pageId="19" pageNumber="362">A. nefte</emphasis>
</taxonomicName>
occurs from S
<collectingCountry id="91D9FAD1DA0D7823252487F2A855F964" box="[1031,1131,1686,1712]" name="Thailand" pageId="19" pageNumber="362">Thailand</collectingCountry>
to
<collectingCountry id="91D9FAD1DA0D782325AE87F2A8C8F964" box="[1165,1270,1686,1712]" name="Malaysia" pageId="19" pageNumber="362">Malaysia</collectingCountry>
, and
<collectingCountry id="91D9FAD1DA0D7823240C87F2A9A2F964" box="[1327,1436,1686,1712]" name="Indonesia" pageId="19" pageNumber="362">Indonesia</collectingCountry>
from
<collectingRegion id="2B0A74A3DA0D782321F687DFAD39F901" box="[213,263,1723,1749]" country="Indonesia" name="Jawa Timur" pageId="19" pageNumber="362">Java</collectingRegion>
to Borneo (
<figureCitation id="71F5A6C4DA0D782320AF87DEADE0F900" box="[396,478,1722,1748]" captionStart="FIGURE 12" captionStartId="19.[151,250,1328,1352]" captionTargetBox="[151,1436,237,1304]" captionTargetId="figure-49@19.[151,1436,237,1304]" captionTargetPageId="19" captionText="FIGURE 12. Distributional range map of relevant taxa from the Athyma nefte species group. Pink: A. inara sahyadriensis ssp. nov., blue: A. i. inara, Purple: A. asita, Yellow: A. nefte. Map: Viraj Nawge." figureDoi="http://doi.org/10.5281/zenodo.14386332" httpUri="https://zenodo.org/record/14386332/files/figure.png" pageId="19" pageNumber="362">Fig. 12</figureCitation>
), in subtly different subspecific variations.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA0D782421E487BAAF9FFE21" lastPageId="20" lastPageNumber="363" pageId="19" pageNumber="362" type="biology_ecology">
<paragraph id="E971BA41DA0D782321E487BAAF6BF85D" blockId="19.[151,1437,1542,2037]" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321E487BAAF8FF92C" bold="true" box="[199,945,1758,1785]" pageId="19" pageNumber="362">
Status, habitat, and habits:
<taxonomicName id="2ECEC1C2DA0D7823232A87BAAF77F92D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[521,841,1758,1785]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0D7823232A87BAAF77F92D" bold="true" box="[521,841,1758,1785]" italics="true" pageId="19" pageNumber="362">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0D7823226D87BAAF8FF92C" box="[846,945,1758,1784]" pageId="19" pageNumber="362" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
occurs in low-lying evergreen forest patches (usually below
<quantity id="2E3617A4DA0D782320608666ADFCF8C9" box="[323,450,1794,1821]" metricMagnitude="2" metricUnit="m" metricValue="7.5" metricValueMax="8.0" metricValueMin="7.0" pageId="19" pageNumber="362" unit="m" value="750.0" valueMax="800.0" valueMin="700.0">700–800 m</quantity>
asl), where it may be seasonally and locally not uncommon along shaded forest streams and forest edges. Males are encountered mud-puddling, or feeding on carnivore scats and rotting crabs, usually on the ground. Females tend to stay on vegetation, often feeding on bird droppings. They have not been observed feeding on flower nectar. Other habits are similar of the genus.
</paragraph>
<paragraph id="E971BA41DA0D782321E486F6AF76F804" blockId="19.[151,1437,1542,2037]" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321E486F6AF7DF878" bold="true" box="[199,835,1938,1965]" pageId="19" pageNumber="362">
Flight period:
<taxonomicName id="2ECEC1C2DA0D7823205D86F6AEF3F879" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[382,717,1938,1965]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0D7823205D86F6AEF3F879" bold="true" box="[382,717,1938,1965]" italics="true" pageId="19" pageNumber="362">Athyma inara sahyadriensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA0D782323FA86F6AF7DF878" box="[729,835,1938,1964]" pageId="19" pageNumber="362" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
is multivoltine, flying throughout the year, with a population peak from October to January (
<bibRefCitation id="8D5FC7B0DA0D7823235086D2AF02F804" author="Kunte, K. &amp; Sondhi, S. &amp; Roy, P. &amp; Chief Editors" box="[627,828,1974,2000]" pageId="19" pageNumber="362" refId="ref14101" refString="Kunte, K., Sondhi, S. &amp; Roy, P. (Chief Editors) (2024) Butterflies of India, v. 4.21. Indian Foundation for Butterflies, Bengaluru. Available from: https: // www. ifoundbutterflies. org (accessed 15 April 2024)." type="url" year="2024">
Kunte
<emphasis id="DBBA6653DA0D782323E386D3AEC6F804" box="[704,760,1974,2000]" italics="true" pageId="19" pageNumber="362">et al.</emphasis>
2024
</bibRefCitation>
).
</paragraph>
<paragraph id="E971BA41DA0D782421E486BEAE2FFF2D" blockId="19.[151,1437,1542,2037]" lastBlockId="20.[151,1437,151,501]" lastPageId="20" lastPageNumber="363" pageId="19" pageNumber="362">
<emphasis id="DBBA6653DA0D782321E486BEAE48F820" bold="true" box="[199,630,2010,2036]" pageId="19" pageNumber="362">Larval host plants and early stages:</emphasis>
The following larval host plants are known for
<emphasis id="DBBA6653DA0D782425B486BEAD3EFF65" bold="true" lastPageId="20" lastPageNumber="363" pageId="19" pageNumber="362">
<taxonomicName id="2ECEC1C2DA0D782325B486BEA9A2F821" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1175,1436,2010,2037]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="362" phylum="Arthropoda" rank="subSpecies" species="inara" subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0D782325B486BEA9A2F821" bold="true" box="[1175,1436,2010,2037]" italics="true" pageId="19" pageNumber="362">A. inara sahyadriensis</emphasis>
</taxonomicName>
ssp. nov.
</emphasis>
:
<taxonomicName id="2ECEC1C2DA0A7824203081F3AE0FFF65" box="[275,561,151,177]" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="species" species="lanceolarium">
<emphasis id="DBBA6653DA0A7824203081F3AE0FFF65" box="[275,561,151,177]" italics="true" pageId="20" pageNumber="363">Glochidion lanceolarium</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA0A7824236781F3AED5FF65" box="[580,747,151,177]" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="species" species="heyneanum">
<emphasis id="DBBA6653DA0A7824236781F3AED5FF65" box="[580,747,151,177]" italics="true" pageId="20" pageNumber="363">G. heyneanum</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA0A782423DE81F3AF9FFF65" box="[765,929,151,177]" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="species" species="zeylanicum">
<emphasis id="DBBA6653DA0A782423DE81F3AF9FFF65" box="[765,929,151,177]" italics="true" pageId="20" pageNumber="363">G. zeylanicum</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA0A7824229781F3A85AFF65" authorityName="Martinov" authorityYear="1820" box="[948,1124,151,177]" class="Magnoliopsida" family="Phyllanthaceae" kingdom="Plantae" order="Malpighiales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="family">Phyllanthaceae</taxonomicName>
), and
<taxonomicName id="2ECEC1C2DA0A7824258C81F3A9A3FF65" box="[1199,1437,151,177]" class="Magnoliopsida" family="Rubiaceae" genus="Mussaenda" kingdom="Plantae" order="Gentianales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="species" species="frondosa">
<emphasis id="DBBA6653DA0A7824258C81F3A9A3FF65" box="[1199,1437,151,177]" italics="true" pageId="20" pageNumber="363">Mussaenda frondosa</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA0A782421BC81DFAD29FF01" box="[159,279,187,213]" class="Magnoliopsida" family="Rubiaceae" kingdom="Plantae" order="Gentianales" pageId="20" pageNumber="363" phylum="Tracheophyta" rank="family">Rubiaceae</taxonomicName>
) (
<bibRefCitation id="8D5FC7B0DA0A7824200D81DFADCCFF01" author="Nitin, R. &amp; Balakrishnan, V. C. &amp; Churi, P. V &amp; Kalesh, S. &amp; Prakash, S. &amp; Kunte, K." box="[302,498,187,213]" pageId="20" pageNumber="363" pagination="11495 - 11550" refId="ref14504" refString="Nitin, R., Balakrishnan, V. C., Churi, P. V, Kalesh, S., Prakash, S. &amp; Kunte, K. (2018) Larval host plants of the butterflies of the Western Ghats, India. Journal of Threatened Taxa, 10 (4), 11495 - 11550. https: // doi. org / 10.11609 / jott. 3104.10.4.11495 - 11550" type="journal article" year="2018">
Nitin
<emphasis id="DBBA6653DA0A7824205181DFAD92FF01" box="[370,428,187,213]" italics="true" pageId="20" pageNumber="363">et al.</emphasis>
2018
</bibRefCitation>
). The early stages are well-documented (
<bibRefCitation id="8D5FC7B0DA0A782422E781DFA8EEFF01" author="Bhakare, M. &amp; Ogale, H." box="[964,1232,187,213]" pageId="20" pageNumber="363" refId="ref12582" refString="Bhakare, M. &amp; Ogale, H. (2018) A Guide to Butterflies of Western Ghats (India). Published privately by Milind Bhakare and Hemant Ogale, Satara, 496 pp." type="book" year="2018">Bhakare &amp; Ogale 2018</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA0A782425FD81DFA9AEFF01" author="Saji, K. &amp; Ogale, H. &amp; Debnath, S. &amp; Mandal, M." box="[1246,1424,187,213]" pageId="20" pageNumber="363" refId="ref14768" refString="Saji, K., Ogale, H., Debnath, S. &amp; Mandal, M. (2024) Athyma inara Westwood, 1850 - Colour Sergeant. In: Kunte, K., Sondhi, S. &amp; Roy, P. (Chief Editors). Butterflies of India, v. 4.21. Indian Foundation for Butterflies, Bengaluru. Available from: https: // www. ifoundbutterflies. org / athyma-inara (accessed 15 April 2024)." type="url" year="2024">
Saji
<emphasis id="DBBA6653DA0A7824243181DFA972FF01" box="[1298,1356,187,213]" italics="true" pageId="20" pageNumber="363">et al.</emphasis>
2024
</bibRefCitation>
), and also illustrated here (
<figureCitation id="71F5A6C4DA0A7824209281BBAE3DFF2D" box="[433,515,223,249]" captionStart="FIGURE 11" captionStartId="18.[151,250,1332,1356]" captionTargetBox="[159,1428,181,1309]" captionTargetId="figure-20@18.[159,1428,181,1309]" captionTargetPageId="18" captionText="FIGURE 11. Early stages of Athyma inara sahyadriensis ssp. nov. a: egg (2010/11/05); b: 1st instar (2010/09/20); c: 2nd instar (2010/09/26); d–e: 3rd instar (2010/10/02); f–i: 4th instar (2010/10/05–07); j–l: 5th instar (2010/10/08); m–n: 5th instar with colours changed just before pupation (2010/10/10). o–q: newly-formed pupa (o: lateral view; p: dorsal view; q: ventral view); r–s: pupa before eclosion (r: lateral view; s: dorsal view); t: eclosed female resting on pupal case, drying its wings. Images are of multiple individuals. Amboli, Sindhudurg District, Maharashtra, India. Photographs: Hemant Ogale, used with permission." figureDoi="http://doi.org/10.5281/zenodo.14386330" httpUri="https://zenodo.org/record/14386330/files/figure.png" pageId="20" pageNumber="363">Fig. 11</figureCitation>
).
</paragraph>
<paragraph id="E971BA41DA0A782421E48066ACC4FE79" blockId="20.[151,1437,151,501]" pageId="20" pageNumber="363">
<emphasis id="DBBA6653DA0A782421E48066AD8FFEC9" bold="true" box="[199,433,258,285]" pageId="20" pageNumber="363">
Sympatric
<taxonomicName id="2ECEC1C2DA0A7824206D8066AD96FEC9" box="[334,424,258,285]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA0A7824206D8066AD96FEC9" bold="true" box="[334,424,258,285]" italics="true" pageId="20" pageNumber="363">Athyma</emphasis>
</taxonomicName>
:
</emphasis>
The following
<taxonomicName id="2ECEC1C2DA0A7824234B8067AE80FEC9" box="[616,702,259,285]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA0A7824234B8067AE80FEC9" box="[616,702,259,285]" italics="true" pageId="20" pageNumber="363">Athyma</emphasis>
</taxonomicName>
occur in the Western Ghats along with
<emphasis id="DBBA6653DA0A782425B08067ACC2FE95" bold="true" pageId="20" pageNumber="363">
<taxonomicName id="2ECEC1C2DA0A782425B08067A9A2FEC9" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1171,1436,258,285]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="subSpecies" species="inara" subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0A782425B08067A9A2FEC9" bold="true" box="[1171,1436,258,285]" italics="true" pageId="20" pageNumber="363">A. inara sahyadriensis</emphasis>
</taxonomicName>
ssp. nov.
</emphasis>
:
<emphasis id="DBBA6653DA0A7824202F8043AD41FE94" box="[268,383,295,321]" italics="true" pageId="20" pageNumber="363">
<taxonomicName id="2ECEC1C2DA0A7824202F8043AD5CFE95" box="[268,354,295,321]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="genus">Athyma</taxonomicName>
p.
</emphasis>
<taxonomicName id="2ECEC1C2DA0A782420A58043AEA8FE95" authority="(Linnaeus, 1758)" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[390,662,294,321]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="perius">
<emphasis id="DBBA6653DA0A782420A58043ADF2FE94" box="[390,460,295,320]" italics="true" pageId="20" pageNumber="363">perius</emphasis>
(Linnaeus, 1758)
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA0A782423808043A846FE95" authority="Fruhstorfer, 1906" authorityName="Fruhstorfer" authorityYear="1906" box="[675,1144,295,321]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="subSpecies" species="ranga" subSpecies="karwara">
<emphasis id="DBBA6653DA0A782423808043AF94FE95" box="[675,938,295,321]" italics="true" pageId="20" pageNumber="363">Athyma ranga karwara</emphasis>
Fruhstorfer, 1906
</taxonomicName>
, and
<taxonomicName id="2ECEC1C2DA0A782425978043ADB3FEB1" authority="(Evans, 1924)" baseAuthorityName="Evans" baseAuthorityYear="1924" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="subSpecies" species="selenophora" subSpecies="kanara">
<emphasis id="DBBA6653DA0A782425978043ACD8FEB1" italics="true" pageId="20" pageNumber="363">Athyma selenophora kanara</emphasis>
(Evans, 1924)
</taxonomicName>
.
<taxonomicName id="2ECEC1C2DA0A782420B4802FAE07FEB0" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[407,569,331,357]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="perius">
<emphasis id="DBBA6653DA0A782420B4802FAE07FEB0" box="[407,569,331,357]" italics="true" pageId="20" pageNumber="363">Athyma perius</emphasis>
</taxonomicName>
normally occurs on open ridge-tops and plateaus, not in the shaded evergreen forest habitats as
<emphasis id="DBBA6653DA0A7824207D800BAEEFFE5D" bold="true" box="[350,721,366,393]" pageId="20" pageNumber="363">
<taxonomicName id="2ECEC1C2DA0A7824207D800BAE5DFE5D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[350,611,366,393]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="subSpecies" species="inara" subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA0A7824207D800BAE5DFE5D" bold="true" box="[350,611,366,393]" italics="true" pageId="20" pageNumber="363">A. inara sahyadriensis</emphasis>
</taxonomicName>
ssp. nov.
</emphasis>
, but
<taxonomicName id="2ECEC1C2DA0A7824222F800BAF51FE5C" authorityName="Moore" authorityYear="1858" box="[780,879,367,393]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="ranga">
<emphasis id="DBBA6653DA0A7824222F800BAF51FE5C" box="[780,879,367,393]" italics="true" pageId="20" pageNumber="363">A. ranga</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA0A7824228B800BA86CFE5D" baseAuthorityName="Kollar" baseAuthorityYear="1844" box="[936,1106,367,393]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="selenophora">
<emphasis id="DBBA6653DA0A7824228B800BA86CFE5D" box="[936,1106,367,393]" italics="true" pageId="20" pageNumber="363">A. selenophora</emphasis>
</taxonomicName>
may occur side-by-side with
<taxonomicName id="2ECEC1C2DA0A782421B480F7ACCDFE78" box="[151,243,403,429]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA0A782421B480F7ACCDFE78" box="[151,243,403,429]" italics="true" pageId="20" pageNumber="363">A. inara</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="E971BA41DA0A782421E480D3AF9FFE21" blockId="20.[151,1437,151,501]" pageId="20" pageNumber="363">
Further information will be made available on the species page of the Butterflies of
<collectingCountry id="91D9FAD1DA0A7824254B80D3A89CFE05" box="[1128,1186,439,465]" name="India" pageId="20" pageNumber="363">India</collectingCountry>
website (https://www. ifoundbutterflies.org/
<taxonomicName id="2ECEC1C2DA0A782420A580BFADDFFE21" box="[390,481,475,501]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="363" phylum="Arthropoda" rank="genus">Athyma</taxonomicName>
) as additional observations accumulate.
</paragraph>
</subSubSection>
</treatment>
</document>

142
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@ -1,142 +0,0 @@
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<emphasis id="DBBA6653DA13783D21B481F3AE8EFF65" bold="true" box="[151,688,151,178]" pageId="13" pageNumber="356">
<taxonomicName id="2ECEC1C2DA13783D21B481F3AE7EFF65" authority="Kunte" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[151,576,151,178]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="13" pageNumber="356" phylum="Arthropoda" rank="subSpecies" species="inara" status="subsp. nov." subSpecies="sahyadriensis">
<emphasis id="DBBA6653DA13783D21B481F3ADD5FF66" bold="true" box="[151,491,151,178]" italics="true" pageId="13" pageNumber="356">Athyma inara sahyadriensis</emphasis>
Kunte
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA13783D236481F3AE8EFF65" box="[583,688,151,177]" pageId="13" pageNumber="356" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
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</subSubSection>
<subSubSection id="A1D4E9CADA13783D21B481BBAE85FEB1" pageId="13" pageNumber="356" type="materials_examined">
<paragraph id="E971BA41DA13783D21B481BBAE85FEB1" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21B481BBAD50FF2D" bold="true" box="[151,366,223,249]" pageId="13" pageNumber="356">
<typeStatus id="367504E3DA13783D21B481BBAD3AFF2D" box="[151,260,223,249]" pageId="13" pageNumber="356" type="holotype">Holotype</typeStatus>
(
<figureCitation id="71F5A6C4DA13783D203681BBAD61FF2D" box="[277,351,223,249]" captionStart="FIGURE 7" captionStartId="14.[151,250,1820,1844]" captionTargetBox="[282,1304,194,1783]" captionTargetId="figure-20@14.[274,1314,181,1796]" captionTargetPageId="14" captionText="FIGURE 7. Type specimens of Athyma inara sahyadriensis ssp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386318" httpUri="https://zenodo.org/record/14386318/files/figure.png" pageId="13" pageNumber="356">Fig. 7</figureCitation>
):
</emphasis>
Voucher code IBC-PS982.
<specimenCount id="FFC871C8DA13783D238D8184AEF8FF2E" box="[686,710,224,250]" count="1" pageId="13" pageNumber="356" type="male">♂</specimenCount>
. Near
<collectingCountry id="91D9FAD1DA13783D22308184AF57FF2E" box="[787,873,224,250]" name="India" pageId="13" pageNumber="356">Kannur</collectingCountry>
,
<collectingCountry id="91D9FAD1DA13783D22558184AFF4FF2E" box="[886,970,224,250]" name="India" pageId="13" pageNumber="356">Kannur</collectingCountry>
District,
<collectingRegion id="2B0A74A3DA13783D251B81BBA8BBFF2D" box="[1080,1157,223,249]" country="India" name="Kerala" pageId="13" pageNumber="356">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA13783D25B081BBA8F3FF2D" box="[1171,1229,223,249]" name="India" pageId="13" pageNumber="356">India</collectingCountry>
(lat-long: approx.
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,
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). Approx.
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asl. Evergreen forest. Raised from a caterpillar collected from
<taxonomicName id="2ECEC1C2DA13783D243D8067AC89FE95" class="Magnoliopsida" family="Phyllanthaceae" genus="Glochidion" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="species" species="undetermined">
<emphasis id="DBBA6653DA13783D243D8067AC89FE95" italics="true" pageId="13" pageNumber="356">Glochidion sp.</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA13783D21E48043AD4AFE95" authorityName="Martinov" authorityYear="1820" box="[199,372,295,321]" class="Magnoliopsida" family="Phyllanthaceae" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="356" phylum="Tracheophyta" rank="family">Phyllanthaceae</taxonomicName>
) by Saji K., eclosed and pinned dry on
<date id="9D709C81DA13783D221F8043AF85FE95" box="[828,955,295,321]" pageId="13" pageNumber="356" value="2013-08-03">2013/03/08</date>
by Krushnamegh Kunte. Deposited in the Biodiversity Lab Research Collections at NCBS.
</paragraph>
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<emphasis id="DBBA6653DA13783D21E4800BAD89FE5D" bold="true" box="[199,439,367,393]" pageId="13" pageNumber="356">
Description (
<figureCitation id="71F5A6C4DA13783D2042800BAD96FE5D" box="[353,424,367,393]" captionStart="FIGURE 7" captionStartId="14.[151,250,1820,1844]" captionTargetBox="[282,1304,194,1783]" captionTargetId="figure-20@14.[274,1314,181,1796]" captionTargetPageId="14" captionText="FIGURE 7. Type specimens of Athyma inara sahyadriensis ssp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu, Ujwala Pawar, and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386318" httpUri="https://zenodo.org/record/14386318/files/figure.png" pageId="13" pageNumber="356">Fig. 7</figureCitation>
):
</emphasis>
</paragraph>
<paragraph id="E971BA41DA13783D21E480F0AF08FDCD" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E480F0ADBCFE79" box="[199,386,403,429]" italics="true" pageId="13" pageNumber="356">Forewing length</emphasis>
:
<quantity id="2E3617A4DA13783D20B080F7ADD9FE7A" box="[403,487,403,430]" metricMagnitude="-2" metricUnit="m" metricValue="2.9" pageId="13" pageNumber="356" unit="mm" value="29.0">29 mm</quantity>
.
<emphasis id="DBBA6653DA13783D20D080F7AEFDFE79" box="[499,707,403,429]" italics="true" pageId="13" pageNumber="356">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA13783D23F780F7AF16FE7A" box="[724,808,403,430]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="356" unit="mm" value="17.0">17 mm</quantity>
. Head black. Eyes pale grey, spotted dark brown (eyes turn dark brown or black upon preservation). Antennae black. Proboscis grey-brown. Thorax black above with two very faint, diffused white or pale grey spots; white below. Abdomen black above with white bands on the first two abdominal segments; white below. Legs pale greyish-white.
</paragraph>
<paragraph id="E971BA41DA13783D21E48347ADCDFC8A" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E48347ADF0FDE9" box="[199,462,547,573]" italics="true" pageId="13" pageNumber="356">Dorsal side: Forewings</emphasis>
black.A very narrow, faint, diffused white line from the base to the centre of discal cell, and a large white spot at the end of discal cell. Large white spots stacked in line in interspaces 1a, 1b and 2, and then again in interspaces 4–6. A very small, diffused pale greyish-white spot in interspace 3. Dark orange subapical spots in interspaces 4 and 5, with very small, inconspicuous spots in 3 and 5. An inconspicuous, diffused, pale grey submarginal line running parallel to termen, separated from a similar line that continues from the orange subapical spots to dorsum.
<emphasis id="DBBA6653DA13783D207683B3ADF0FD25" box="[341,462,727,753]" italics="true" pageId="13" pageNumber="356">Hindwings</emphasis>
black. A white band from the base through the cell to v6, the spots in 7 separated by the grey and black v7. A narrow, orange post-discal band interrupted by black veins from interspaces 1c to 7, its edges smooth, straight, not prominently convex. An inconspicuous, diffused, pale grey submarginal line running parallel to termen, from tornus to apex.
</paragraph>
<paragraph id="E971BA41DA13783D21E48203AF26FBC5" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<emphasis id="DBBA6653DA13783D21E48203ADF8FC55" box="[199,454,871,897]" italics="true" pageId="13" pageNumber="356">Ventral side: Forewing</emphasis>
tawny with grey and white markings that are similar to those on the dorsal side, and with additional black smudges surrounding the white spots. Basal line through the discal cell is broader and grey, and conspicuous (rather than narrow, white and inconspicuous, as on the dorsal side).
<emphasis id="DBBA6653DA13783D251682CBA89DFC1D" box="[1077,1187,943,969]" italics="true" pageId="13" pageNumber="356">Hindwing</emphasis>
grey post-discal band (corresponding to the narrow, orange post-discal band on dorsal side) is continued from interspace 1b to 4, but not beyond. HW submarginal grey specks between the veins.
</paragraph>
<paragraph id="E971BA41DA13783D21E4857FAF75FBA9" blockId="13.[151,1437,223,1150]" pageId="13" pageNumber="356">
<materialsCitation id="59A6B01CDA13783D21E4857FAF76FBA9" collectionCode="UNH" pageId="13" pageNumber="356" specimenCount="1" typeStatus="paratype">
This is a dry season form, in which
<collectionCode id="8FDF2284DA13783D23468578AE9CFBE2" box="[613,674,1052,1078]" country="0" httpUri="http://grbio.org/cool/24ny-b7ax" name="University of New Hampshire" pageId="13" pageNumber="356">UNH</collectionCode>
areas between the discal white band and post-discal grey band are plain, with dark reddish, highly diffused spots barely visible between the two bands (these spots are black and very prominent in the wet season form; see
<typeStatus id="367504E3DA13783D236B8500AE96FBAA" box="[584,680,1124,1150]" pageId="13" pageNumber="356" type="paratype">paratype</typeStatus>
IBC-AK310)
</materialsCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

134
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<mods:title id="BDDC9C8D75ED4DFC9AF44F3C5D661BD2">Two new taxa of brush-footed butterflies (Lepidoptera: Nymphalidae) from the Western Ghats biodiversity hotspot, southern India</mods:title>
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<mods:namePart id="624AFFDCC09956C9154D6273CEAE8F3E">Kunte, Krushnamegh</mods:namePart>
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<mods:affiliation id="23290914209E77FBCD851E8C38B547C5">National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK Campus, Bellary Road, Bengaluru 560 065, India &amp; Ashoka University, Rajiv Gandhi Education City, Sonipat, Haryana 131029, India</mods:affiliation>
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<subSubSection id="A1D4E9CADA1C783221B48067AE26FEC9" box="[151,536,258,285]" pageId="2" pageNumber="345" type="nomenclature">
<paragraph id="E971BA41DA1C783221B48067AE26FEC9" blockId="2.[151,1437,150,2013]" box="[151,536,258,285]" pageId="2" pageNumber="345">
4.
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<taxonomicName id="2ECEC1C2DA1C783221E48066AE31FEC9" authority="Fruhstorfer, 1904" authorityName="Fruhstorfer" authorityYear="1904" box="[199,527,258,285]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="friderici" subSpecies="friderici">
<emphasis id="DBBA6653DA1C783221E48066AE31FEC9" bold="true" box="[199,527,258,285]" italics="true" pageId="2" pageNumber="345">Amathusia friderici friderici</emphasis>
</taxonomicName>
:
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1C783223038067AD6BFE79" pageId="2" pageNumber="345" type="materials_examined">
<paragraph id="E971BA41DA1C783223038067AD6BFE79" blockId="2.[151,1437,150,2013]" pageId="2" pageNumber="345">
<materialsCitation id="59A6B01CDA1C783223038067AE9AFE95" collectionCode="BMNH" country="Myanmar" location="Tenasserim" pageId="2" pageNumber="345" specimenCount="1" specimenCount-male="1" typeStatus="holotype">
<specimenCount id="FFC871C8DA1C783223038067AE06FEC9" box="[544,568,259,285]" count="1" pageId="2" pageNumber="345" type="male">♂</specimenCount>
:
<collectionCode id="8FDF2284DA1C783223678067AEA5FEC9" box="[580,667,259,285]" pageId="2" pageNumber="345">BMNH</collectionCode>
(E) #141394, double-numbered with BMNH(E) #806351 (
<typeStatus id="367504E3DA1C783224088067A9A6FEC9" box="[1323,1432,259,285]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
.
<location id="EC11EC9ADA1C783221E48043AD77FE95" LSID="urn:lsid:plazi:treatment:61670B57DA1C783221B48067AD6BFE79:EC11EC9ADA1C783221E48043AD77FE95" box="[199,329,295,321]" country="Myanmar" name="Tenasserim" pageId="2" pageNumber="345">Tenasserim</location>
,
<collectingCountry id="91D9FAD1DA1C783220758043ADFBFE95" box="[342,453,295,321]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783220F28043AE2AFE95" box="[465,532,295,321]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
),
<collectionCode id="8FDF2284DA1C7832230F8043AEBDFE95" box="[556,643,295,321]" pageId="2" pageNumber="345">BMNH</collectionCode>
(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1C783223988042ACD8FEB1" collectionCode="NHMUK" collectorName="Karen Hills &amp; Tenasserim" country="Myanmar" location="northern Tenasserim Hills" municipality="Kawkareik" pageId="2" pageNumber="345" specimenCode="NHMUK010244425" specimenCount="1" typeStatus="holotype">
805388 (
<location id="EC11EC9ADA1C7832223D8043A87BFE95" LSID="urn:lsid:plazi:treatment:61670B57DA1C783221B48067AD6BFE79:EC11EC9ADA1C7832223D8043A87BFE95" box="[798,1093,295,321]" country="Myanmar" municipality="Kawkareik" name="northern Tenasserim Hills" pageId="2" pageNumber="345">northern Tenasserim Hills</location>
at
<collectingMunicipality id="0915203BDA1C783225488043A8D7FE95" box="[1131,1257,295,321]" pageId="2" pageNumber="345">Kawkareik</collectingMunicipality>
,
<collectingCountry id="91D9FAD1DA1C783225D68043A95AFE95" box="[1269,1380,295,321]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783224538043ACE9FEB1" captionStart="FIGURE 4" captionStartId="9.[151,250,1824,1848]" captionTargetBox="[180,1395,186,1785]" captionTargetId="figure-19@9.[170,1416,181,1801]" captionTargetPageId="9" captionText="FIGURE 4. Diagnostic characters of Amathusia travancorica sp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386309" httpUri="https://zenodo.org/record/14386309/files/figure.png" pageId="2" pageNumber="345">Fig. 4</figureCitation>
),
</materialsCitation>
<materialsCitation id="59A6B01CDA1C783221D0802EAFD9FEB1" box="[243,999,330,357]" collectionCode="NHMUK" collectorName="Karen Hills &amp; Tenasserim" country="Myanmar" location="northern Tenasserim Hills" municipality="Kawkareik" pageId="2" pageNumber="345" specimenCode="NHMUK010244425" specimenCount="1" typeStatus="holotype">
<specimenCode id="B968123ADA1C783221D0802EADE5FEB1" box="[243,475,330,357]" collectionCode="NHMUK" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="2" pageNumber="345" type="Museum">NHMUK010244425</specimenCode>
(
<collectorName id="443BDF97DA1C783220D3802FAE43FEB1" box="[496,637,331,357]" pageId="2" pageNumber="345">Karen Hills</collectorName>
,
<collectorName id="443BDF97DA1C783223AC802FAF2FFEB1" box="[655,785,331,357]" pageId="2" pageNumber="345">Tenasserim</collectorName>
,
<collectingCountry id="91D9FAD1DA1C78322200802FAFACFEB1" box="[803,914,331,357]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
. 1920)
</materialsCitation>
,
<materialsCitation id="59A6B01CDA1C783222DA802FAE59FE5D" collectionCode="BMNH" country="Malaysia" location="Selangor" pageId="2" pageNumber="345" specimenCount="1" specimenCount-female="1" stateProvince="Selangor" typeStatus="holotype">
BMNH(E) #984600 and BMNH(E) #984608 (
<collectingRegion id="2B0A74A3DA1C7832201B800BADA1FE5D" box="[312,415,367,393]" country="Malaysia" name="Selangor" pageId="2" pageNumber="345">Selangor</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C78322089800BAE2AFE5D" box="[426,532,367,393]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1961).
</materialsCitation>
<materialsCitation id="59A6B01CDA1C7832234E800BAD6CFE79" collectionCode="BMNH" collectorName="All" country="Malaysia" location="Selangor" pageId="2" pageNumber="345" specimenCount="1" specimenCount-female="1" stateProvince="Selangor" typeStatus="holotype">
<specimenCount id="FFC871C8DA1C7832234E800BAEBAFE5D" box="[621,644,367,393]" count="1" pageId="2" pageNumber="345" type="female">♀</specimenCount>
:
<collectionCode id="8FDF2284DA1C783223B2800BAED6FE5D" box="[657,744,367,393]" pageId="2" pageNumber="345">BMNH</collectionCode>
(E) #805379 (
<location id="EC11EC9ADA1C783222A3800BA838FE5D" LSID="urn:lsid:plazi:treatment:61670B57DA1C783221B48067AD6BFE79:EC11EC9ADA1C783222A3800BA838FE5D" box="[896,1030,367,393]" country="Myanmar" name="Karen Hills" pageId="2" pageNumber="345">Karen Hills</location>
,
<collectingCountry id="91D9FAD1DA1C78322532800BA8BEFE5D" box="[1041,1152,367,393]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783225A9800BA8D2FE5D" box="[1162,1260,367,393]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStartId-0="7.[151,250,1957,1981]" captionStartId-1="9.[151,250,1824,1848]" captionTargetBox-0="[155,1436,188,1922]" captionTargetBox-1="[180,1395,186,1785]" captionTargetId-0="figure-19@7.[151,1436,181,1933]" captionTargetId-1="figure-19@9.[170,1416,181,1801]" captionTargetPageId-0="7" captionTargetPageId-1="9" captionText-0="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." captionText-1="FIGURE 4. Diagnostic characters of Amathusia travancorica sp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar." figureDoi-0="http://doi.org/10.5281/zenodo.14386305" figureDoi-1="http://doi.org/10.5281/zenodo.14386309" httpUri-0="https://zenodo.org/record/14386305/files/figure.png" httpUri-1="https://zenodo.org/record/14386309/files/figure.png" pageId="2" pageNumber="345">Fig. 3–4</figureCitation>
).
<collectorName id="443BDF97DA1C783225DE800BA91FFE5D" box="[1277,1313,367,393]" pageId="2" pageNumber="345">All</collectorName>
specimens in
<collectionCode id="8FDF2284DA1C783221C780F7AD6CFE79" box="[228,338,403,429]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="2" pageNumber="345" type="Museum">NHMUK</collectionCode>
</materialsCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

112
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751155 (
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3.
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(
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. 1891.
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,
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(
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(
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:
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(
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?)
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<specimenCode id="B968123ADA1C7832207181BAAE04FF2D" box="[338,570,222,249]" collectionCode="NHMUK" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="2" pageNumber="345" type="Museum">NHMUK010244255</specimenCode>
(
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,
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. 1893). All specimens in NHMUK
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.
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<treatment id="61670B57DA1C783221B48203ADEFFA31" LSID="urn:lsid:plazi:treatment:61670B57DA1C783221B48203ADEFFA31" httpUri="http://treatment.plazi.org/id/61670B57DA1C783221B48203ADEFFA31" lastPageNumber="345" pageId="2" pageNumber="345">
<subSubSection id="A1D4E9CADA1C783221B48203AE01FC55" box="[151,575,870,897]" pageId="2" pageNumber="345" type="nomenclature">
<paragraph id="E971BA41DA1C783221B48203AE01FC55" blockId="2.[151,1437,150,2013]" box="[151,575,870,897]" pageId="2" pageNumber="345">
9.
<emphasis id="DBBA6653DA1C783221E48202AE01FC55" bold="true" box="[199,575,870,897]" pageId="2" pageNumber="345">
<taxonomicName id="2ECEC1C2DA1C783221E48202AE08FC55" ID-CoL="5TYM4" box="[199,566,870,897]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="phidippus">
<emphasis id="DBBA6653DA1C783221E48202AE08FC55" bold="true" box="[199,566,870,897]" italics="true" pageId="2" pageNumber="345">Amathusia phidippus phidippus</emphasis>
</taxonomicName>
:
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1C783223658203A8A3FC39" pageId="2" pageNumber="345" type="reference_group">
<paragraph id="E971BA41DA1C783223658203A8A3FC39" blockId="2.[151,1437,150,2013]" pageId="2" pageNumber="345">
This includes the following synonyms (
<bibRefCitation id="8D5FC7B0DA1C783225238203A89BFC55" author="Inayoshi, Y." box="[1024,1189,871,897]" pageId="2" pageNumber="345" refId="ref13540" refString="Inayoshi, Y. (2023) A Check List of Butterflies in Indo-China, Chiefly from Thailand, Laos &amp; Vietnam. Ching Mai. Available from: http: // yutaka. it-n. jp (accessed 15 April 2024)" type="url" year="2023">Inayoshi 2023</bibRefCitation>
), whose collections in NHMUK were also inspected: (a)
<taxonomicName id="2ECEC1C2DA1C7832236882EFA844FC71" authority="Fruhstorfer, 1904" authorityName="Fruhstorfer" authorityYear="1904" box="[587,1146,907,933]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="adustatus">
<emphasis id="DBBA6653DA1C7832236882EFAF92FC71" box="[587,940,907,933]" italics="true" pageId="2" pageNumber="345">Amathusia phidippus adustatus</emphasis>
Fruhstorfer, 1904
</taxonomicName>
, (b)
<taxonomicName id="2ECEC1C2DA1C7832258D82EFADC4FC1D" authority="Fruhstorfer, 1911" authorityName="Fruhstorfer" authorityYear="1911" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="chersias">
<emphasis id="DBBA6653DA1C7832258D82EFAD1AFC1D" italics="true" pageId="2" pageNumber="345">Amathusia phidippus chersias</emphasis>
Fruhstorfer, 1911
</taxonomicName>
, (c)
<taxonomicName id="2ECEC1C2DA1C7832231B82CBA848FC1D" authority="Fruhstorfer, 1911" authorityName="Fruhstorfer" authorityYear="1911" box="[568,1142,943,969]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="eutropius">
<emphasis id="DBBA6653DA1C7832231B82CBAF9EFC1C" box="[568,928,943,969]" italics="true" pageId="2" pageNumber="345">Amathusia phidippus eutropius</emphasis>
Fruhstorfer, 1911
</taxonomicName>
, (d)
<taxonomicName id="2ECEC1C2DA1C7832259782CBAE37FC39" authority="Corbet &amp; Pendlebury, 1936" authorityName="Corbet &amp; Pendlebury" authorityYear="1936" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="gunneryi">
<emphasis id="DBBA6653DA1C7832259782CBA9A3FC1C" box="[1204,1437,943,969]" italics="true" pageId="2" pageNumber="345">Amathusia gunneryi</emphasis>
Corbet &amp; Pendlebury, 1936
</taxonomicName>
, and (e)
<taxonomicName id="2ECEC1C2DA1C7832235682B7A8A7FC39" authority="Fruhstorfer, 1916" authorityName="Fruhstorfer" authorityYear="1916" box="[629,1177,979,1005]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="majada">
<emphasis id="DBBA6653DA1C7832235682B7AFFBFC39" box="[629,965,979,1005]" italics="true" pageId="2" pageNumber="345">Amathusia phidippus majada</emphasis>
Fruhstorfer, 1916
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1C7832258B82B0ADEFFA31" pageId="2" pageNumber="345" type="materials_examined">
<paragraph id="E971BA41DA1C7832258B82B0ADEFFA31" blockId="2.[151,1437,150,2013]" pageId="2" pageNumber="345">
<materialsCitation id="59A6B01CDA1C7832258B82B0ADEFFA31" pageId="2" pageNumber="345">
<emphasis id="DBBA6653DA1C7832258B82B0A8F8FC24" bold="true" box="[1192,1222,980,1008]" pageId="2" pageNumber="345">
<specimenCount id="FFC871C8DA1C7832258B82B0A881FC24" box="[1192,1215,980,1008]" count="1" pageId="2" pageNumber="345" type="male">♂</specimenCount>
:
</emphasis>
BMNH # 141395 (
<typeStatus id="367504E3DA1C783221EC8293AD07FBC5" box="[207,313,1015,1041]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
<emphasis id="DBBA6653DA1C783220458293ADEFFBC5" box="[358,465,1015,1041]" italics="true" pageId="2" pageNumber="345">adustatus</emphasis>
. Nawngchik, Bansai Kau,
<collectingCountry id="91D9FAD1DA1C783222338293AF44FBC5" box="[784,890,1015,1041]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1901.
<figureCitation id="71F5A6C4DA1C783222F08293A827FBC5" box="[979,1049,1015,1041]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), NHMUK010244505 (
<collectingRegion id="2B0A74A3DA1C7832240D8293A9A6FBC5" box="[1326,1432,1015,1041]" country="Thailand" name="Krung Thep Mahanakhon" pageId="2" pageNumber="345">Bangkok</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783221E4857FAD10FBE1" box="[199,302,1051,1077]" name="Thailand" pageId="2" pageNumber="345">Thailand</collectingCountry>
. 1901. This has a type label attached for
<emphasis id="DBBA6653DA1C78322200857FAFB0FBE1" box="[803,910,1051,1077]" italics="true" pageId="2" pageNumber="345">adustatus</emphasis>
, in error), BMNH(E) #806352 (
<typeStatus id="367504E3DA1C7832242C857FA947FBE1" box="[1295,1401,1051,1077]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
<emphasis id="DBBA6653DA1C783221E4855BAD1AFB8D" box="[199,292,1087,1113]" italics="true" pageId="2" pageNumber="345">chersias</emphasis>
. Location not specified but assumed to be
<collectingCountry id="91D9FAD1DA1C78322225855BAF56FB8D" box="[774,872,1087,1113]" name="Malaysia" pageId="2" pageNumber="345">Malacca</collectingCountry>
,
<collectingCountry id="91D9FAD1DA1C78322250855BAFE3FB8D" box="[883,989,1087,1113]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783222CB855BA814FB8D" box="[1000,1066,1087,1113]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), BMNH # 141445 (
<typeStatus id="367504E3DA1C78322437855BA940FB8D" box="[1300,1406,1087,1113]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
<taxonomicName id="2ECEC1C2DA1C783221E48507AD15FBA8" authorityName="Corbet &amp; Pendlebury" authorityYear="1936" box="[199,299,1123,1148]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="gunneryi">
<emphasis id="DBBA6653DA1C783221E48507AD15FBA8" box="[199,299,1123,1148]" italics="true" pageId="2" pageNumber="345">gunneryi</emphasis>
</taxonomicName>
.
<collectingRegion id="2B0A74A3DA1C783220198507ADDFFBA9" box="[314,481,1123,1149]" country="Malaysia" name="Kuala Lumpur" pageId="2" pageNumber="345">Kuala Lumpur</collectingRegion>
,
<collectingRegion id="2B0A74A3DA1C783220CE8507AE6CFBA9" box="[493,594,1123,1149]" country="Malaysia" name="Selangor" pageId="2" pageNumber="345">Selangor</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C7832237C8507AEF7FBA9" box="[607,713,1123,1149]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1930.
<figureCitation id="71F5A6C4DA1C7832223F8507AF5EFBA9" box="[796,864,1123,1149]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), NHMUK010244885 (
<typeStatus id="367504E3DA1C7832254C8507A8E7FBA9" box="[1135,1241,1123,1149]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
<emphasis id="DBBA6653DA1C783224238507A96AFBA9" box="[1280,1364,1123,1149]" italics="true" pageId="2" pageNumber="345">majada</emphasis>
.
<collectingRegion id="2B0A74A3DA1C783224408507A9A6FBA9" box="[1379,1432,1123,1149]" country="Indonesia" name="Jawa Timur" pageId="2" pageNumber="345">Java</collectingRegion>
.
<figureCitation id="71F5A6C4DA1C783221E485E3AD16FB75" box="[199,296,1159,1185]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStartId-0="7.[151,250,1957,1981]" captionStartId-1="9.[151,250,1824,1848]" captionTargetBox-0="[155,1436,188,1922]" captionTargetBox-1="[180,1395,186,1785]" captionTargetId-0="figure-19@7.[151,1436,181,1933]" captionTargetId-1="figure-19@9.[170,1416,181,1801]" captionTargetPageId-0="7" captionTargetPageId-1="9" captionText-0="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." captionText-1="FIGURE 4. Diagnostic characters of Amathusia travancorica sp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar." figureDoi-0="http://doi.org/10.5281/zenodo.14386305" figureDoi-1="http://doi.org/10.5281/zenodo.14386309" httpUri-0="https://zenodo.org/record/14386305/files/figure.png" httpUri-1="https://zenodo.org/record/14386309/files/figure.png" pageId="2" pageNumber="345">Fig. 3–4</figureCitation>
), BMNH(E) #984660 (
<collectingRegion id="2B0A74A3DA1C7832230E85E3AE5DFB75" box="[557,611,1159,1185]" country="Indonesia" name="Jawa Timur" pageId="2" pageNumber="345">Java</collectingRegion>
. 1921), BMNH(E) #984642 (
<collectingRegion id="2B0A74A3DA1C7832228C85E3AFDBFB75" box="[943,997,1159,1185]" country="Indonesia" name="Jawa Timur" pageId="2" pageNumber="345">Java</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783222D385E3A85FFB75" box="[1008,1121,1159,1185]" name="Indonesia" pageId="2" pageNumber="345">Indonesia</collectingCountry>
. 1898), BMNH(E) #984594 (
<collectingRegion id="2B0A74A3DA1C783221EC85CFAD0FFB11" box="[207,305,1195,1221]" country="Indonesia" name="Sumatera Utara" pageId="2" pageNumber="345">Sumatra</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C7832206285CFAD8CFB11" box="[321,434,1195,1221]" name="Indonesia" pageId="2" pageNumber="345">Indonesia</collectingCountry>
), BMNH(E) #984573 (
<collectingCountry id="91D9FAD1DA1C783223E585CFAF01FB11" box="[710,831,1195,1221]" name="Singapore" pageId="2" pageNumber="345">Singapore</collectingCountry>
?), BMNH(E) #984574 (
<collectingRegion id="2B0A74A3DA1C7832257E85CFA8ADFB11" box="[1117,1171,1195,1221]" country="Indonesia" name="Jawa Timur" pageId="2" pageNumber="345">Java</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C7832258785CFA92BFB11" box="[1188,1301,1195,1221]" name="Indonesia" pageId="2" pageNumber="345">Indonesia</collectingCountry>
. 1898.
<figureCitation id="71F5A6C4DA1C7832245385CFACE8FB3D" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), NHMUK010244361 (
<collectingCountry id="91D9FAD1DA1C783220CA85ABAE70FB3D" box="[489,590,1231,1257]" name="Vietnam" pageId="2" pageNumber="345">Vietnam</collectingCountry>
), NHMUK010244151 (“Lagi
<collectingCountry id="91D9FAD1DA1C7832228585ABAFC0FB3D" box="[934,1022,1231,1257]" name="Vietnam" pageId="2" pageNumber="345">Annam</collectingCountry>
”. 1921), NHMUK010244540 (Java. 1895).
<emphasis id="DBBA6653DA1C783220358590AD0AFAC4" bold="true" box="[278,308,1268,1296]" pageId="2" pageNumber="345">
<specimenCount id="FFC871C8DA1C783220358590AD13FAC4" box="[278,301,1268,1296]" count="1" pageId="2" pageNumber="345" type="female">♀</specimenCount>
:
</emphasis>
BMNH # 141393 (
<collectingRegion id="2B0A74A3DA1C7832232F8597AE48FAD9" box="[524,630,1267,1293]" country="Thailand" name="Krung Thep Mahanakhon" pageId="2" pageNumber="345">Bangkok</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783223A38597AED9FAD9" box="[640,743,1267,1293]" name="Thailand" pageId="2" pageNumber="345">Thailand</collectingCountry>
. 1901. This is a presumed, but unconfirmed, type of
<emphasis id="DBBA6653DA1C783224128597A9A2FAD9" box="[1329,1436,1267,1293]" italics="true" pageId="2" pageNumber="345">adustatus</emphasis>
based on a red label attached), BMNH(E) #806353 (
<typeStatus id="367504E3DA1C783222288473AF51FAE5" box="[779,879,1303,1329]" pageId="2" pageNumber="345" type="allotype">Allotype</typeStatus>
of
<emphasis id="DBBA6653DA1C783222B18473AFD1FAE5" box="[914,1007,1303,1329]" italics="true" pageId="2" pageNumber="345">chersias</emphasis>
.
<collectingCountry id="91D9FAD1DA1C783222DE8473A861FAE5" box="[1021,1119,1303,1329]" name="Malaysia" pageId="2" pageNumber="345">Malacca</collectingCountry>
,
<collectingCountry id="91D9FAD1DA1C7832254A8473A8EDFAE5" box="[1129,1235,1303,1329]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783225FE8473A920FAE5" box="[1245,1310,1303,1329]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), BMNH # 141442 (
<typeStatus id="367504E3DA1C7832200B845FADB2FA81" box="[296,396,1339,1365]" pageId="2" pageNumber="345" type="allotype">Allotype</typeStatus>
of
<taxonomicName id="2ECEC1C2DA1C7832208E845FAE2FFA80" authorityName="Corbet &amp; Pendlebury" authorityYear="1936" box="[429,529,1339,1364]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="gunneryi">
<emphasis id="DBBA6653DA1C7832208E845FAE2FFA80" box="[429,529,1339,1364]" italics="true" pageId="2" pageNumber="345">gunneryi</emphasis>
</taxonomicName>
.
<collectingRegion id="2B0A74A3DA1C7832233E845FAEFFFA81" box="[541,705,1339,1365]" country="Malaysia" name="Kuala Lumpur" pageId="2" pageNumber="345">Kuala Lumpur</collectingRegion>
,
<collectingRegion id="2B0A74A3DA1C783223E9845FAF11FA81" box="[714,815,1339,1365]" country="Malaysia" name="Selangor" pageId="2" pageNumber="345">Selangor</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C7832221B845FAF9CFA81" box="[824,930,1339,1365]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1930.
<figureCitation id="71F5A6C4DA1C783222CC845FA80EFA81" box="[1007,1072,1339,1365]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), NHMUK010244706 (
<typeStatus id="367504E3DA1C7832241B845FA9A2FA81" box="[1336,1436,1339,1365]" pageId="2" pageNumber="345" type="allotype">Allotype</typeStatus>
of
<emphasis id="DBBA6653DA1C783221C4843BAD05FAAD" box="[231,315,1375,1401]" italics="true" pageId="2" pageNumber="345">majada</emphasis>
. Java.
<figureCitation id="71F5A6C4DA1C783220AF843BADEEFAAD" box="[396,464,1375,1401]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
), NHMUK010244332 (Java. 1898), NHMUK010244426 (
<collectingRegion id="2B0A74A3DA1C78322548843BA8EBFAAD" box="[1131,1237,1375,1401]" country="Thailand" name="Krung Thep Mahanakhon" pageId="2" pageNumber="345">Bangkok</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783225C2843BA976FAAD" box="[1249,1352,1375,1401]" name="Thailand" pageId="2" pageNumber="345">Thailand</collectingCountry>
. 1914), NHMUK010244420 (Java. 1898), BMNH(E) #984479 (Mergui,
<collectingCountry id="91D9FAD1DA1C7832229884E7A814FA49" box="[955,1066,1411,1437]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
.
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), BMNH(E) #984619 (
<collectingRegion id="2B0A74A3DA1C783223B284C3AEF9FA15" box="[657,711,1447,1473]" country="Indonesia" name="Jawa Timur" pageId="2" pageNumber="345">Java</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783223F784C3AF7BFA15" box="[724,837,1447,1473]" name="Indonesia" pageId="2" pageNumber="345">Indonesia</collectingCountry>
. 1912), BMNH(E) #984629 (Marang,
<collectingCountry id="91D9FAD1DA1C783225D884C3A959FA15" box="[1275,1383,1447,1473]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
). All specimens in NHMUK.
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7.
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),
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(
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specimens in
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8.
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<emphasis id="DBBA6653DA1C783221E4839EAE0AFCC1" bold="true" box="[199,564,762,789]" italics="true" pageId="2" pageNumber="345">Amathusia perakana perakana</emphasis>
Honrath, [1888]
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:
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<specimenCount id="FFC871C8DA1C78322236839FAF13FCC1" box="[789,813,763,789]" count="1" pageId="2" pageNumber="345" type="male">♂</specimenCount>
:
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, double-numbered with BMNH(E) #806354 (
<typeStatus id="367504E3DA1C7832201D827BAD95FCED" box="[318,427,799,825]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
.
<location id="EC11EC9ADA1C78322098827BADC5FCED" LSID="urn:lsid:plazi:treatment:61670B57DA1C783221B4839FA83AFC89:EC11EC9ADA1C78322098827BADC5FCED" box="[443,507,799,825]" country="Malaysia" name="Larut" pageId="2" pageNumber="345" stateProvince="Perak">Larut</location>
,
<collectingRegion id="2B0A74A3DA1C78322328827BAE70FCED" box="[523,590,799,825]" country="Malaysia" name="Perak" pageId="2" pageNumber="345">Perak</collectingRegion>
,
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. 1885.
<figureCitation id="71F5A6C4DA1C78322206827BAF53FCED" box="[805,877,799,825]" captionStart="FIGURE 3" captionStartId="7.[151,250,1957,1981]" captionTargetBox="[155,1436,188,1922]" captionTargetId="figure-19@7.[151,1436,181,1933]" captionTargetPageId="7" captionText="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="2" pageNumber="345">Fig. 3</figureCitation>
),
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<specimenCode id="B968123ADA1C783222AB827BA84EFCED" box="[904,1136,799,825]" collectionCode="NHMUK" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="2" pageNumber="345" type="Museum">NHMUK010244339</specimenCode>
(
<collectingRegion id="2B0A74A3DA1C783225A6827BA8F7FCED" box="[1157,1225,799,825]" country="Malaysia" name="Perak" pageId="2" pageNumber="345">Perak</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C783225F9827BA97AFCED" box="[1242,1348,799,825]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1890)
</materialsCitation>
,
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<specimenCode id="B968123ADA1C783221E48227AD91FC89" box="[199,431,835,861]" collectionCode="NHMUK" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="2" pageNumber="345" type="Museum">NHMUK010244268</specimenCode>
(
<collectingRegion id="2B0A74A3DA1C7832209D8227AE3CFC89" box="[446,514,835,861]" country="Malaysia" name="Perak" pageId="2" pageNumber="345">Perak</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1C7832232E8227AE49FC89" box="[525,631,835,861]" name="Malaysia" pageId="2" pageNumber="345">Malaysia</collectingCountry>
. 1897).
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specimens in
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10.
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<emphasis id="DBBA6653DA1C783221E4848AAE50F9DD" bold="true" box="[199,622,1518,1545]" italics="true" pageId="2" pageNumber="345">Amathusia schoenbergi schoenbergi</emphasis>
Honrath, [1888]
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<collectingCountry id="91D9FAD1DA1C783221E48777AD17F9F9" box="[199,297,1555,1581]" name="Malaysia" pageId="2" pageNumber="345">Malacca</collectingCountry>
,
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. 1886), NHMUK010244523 (Borneo), NHMUK010244525 (Borneo). All specimens in NHMUK. This includes
<taxonomicName id="2ECEC1C2DA1C783220FF8753A812F985" authority="Fruhstorfer, 1899" authorityName="Fruhstorfer" authorityYear="1899" box="[476,1068,1591,1617]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="schoenbergi" subSpecies="borneensis">
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12.
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<taxonomicName id="2ECEC1C2DA1C783221E48656AD86F899" box="[199,440,1842,1869]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="subSpecies" species="inara" subSpecies="inara">
<emphasis id="DBBA6653DA1C783221E48656AD86F899" bold="true" box="[199,440,1842,1869]" italics="true" pageId="2" pageNumber="345">Athyma inara inara</emphasis>
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:
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This includes the following synonyms: (a)
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<emphasis id="DBBA6653DA1C783222F08657A8B6F898" box="[979,1160,1843,1869]" italics="true" pageId="2" pageNumber="345">Athyma inarina</emphasis>
(
<bibRefCitation id="8D5FC7B0DA1C783225BE8657A913F899" author="Evans, W. H." box="[1181,1325,1843,1869]" pageId="2" pageNumber="345" refId="ref13172" refString="Evans, W. H. (1932) The Identification of Indian Butterflies (2 nd edn.). Bombay Natural History Society, Mumbai, 454 pp." type="book" year="1932">Evans 1932</bibRefCitation>
)
</taxonomicName>
, and (b)
<taxonomicName id="2ECEC1C2DA1C783221E48633AE03F8A5" authority="Swinhoe, 1917" authorityName="Swinhoe" authorityYear="1917" box="[199,573,1879,1905]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA1C783221E48633ADB4F8A4" box="[199,394,1879,1904]" italics="true" pageId="2" pageNumber="345">Pantoporia mera</emphasis>
<bibRefCitation id="8D5FC7B0DA1C783220B08633AE03F8A5" author="Swinhoe, C." box="[403,573,1879,1905]" pageId="2" pageNumber="345" pagination="407 - 412" refId="ref15041" refString="Swinhoe, C. (1917) New Indo-Malayan Lepidoptera. Annals and Magazine of Natural History, 20, 407 - 412. https: // doi. org / 10.1080 / 00222931709487028" type="journal article" year="1917">Swinhoe, 1917</bibRefCitation>
</taxonomicName>
(see below for a note on synonymy of this taxon).
</paragraph>
</subSubSection>
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<paragraph id="E971BA41DA1C783325B2863CAD6BFEB1" blockId="2.[151,1437,150,2013]" lastBlockId="3.[151,1437,150,753]" lastPageId="3" lastPageNumber="346" pageId="2" pageNumber="345">
<materialsCitation id="59A6B01CDA1C783325B2863CAD6CFEB1" lastPageId="3" lastPageNumber="346" pageId="2" pageNumber="345">
<emphasis id="DBBA6653DA1C783225B2863CA88EF8A0" bold="true" box="[1169,1200,1880,1908]" pageId="2" pageNumber="345">
<specimenCount id="FFC871C8DA1C783225B2863CA896F8A0" box="[1169,1192,1880,1908]" count="1" pageId="2" pageNumber="345" type="male">♂</specimenCount>
:
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BMNH(E) #933444 (
<typeStatus id="367504E3DA1C783221EC861FAD07F841" box="[207,313,1915,1941]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
<taxonomicName id="2ECEC1C2DA1C78322046861FAD9CF840" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[357,418,1915,1940]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="inara">
<emphasis id="DBBA6653DA1C78322046861FAD9CF840" box="[357,418,1915,1940]" italics="true" pageId="2" pageNumber="345">inara</emphasis>
</taxonomicName>
. Silhet,
<collectingCountry id="91D9FAD1DA1C78322325861FAEB2F841" box="[518,652,1915,1941]" name="Bangladesh" pageId="2" pageNumber="345">Bangladesh</collectingCountry>
.
<figureCitation id="71F5A6C4DA1C783223B8861FAF33F841" box="[667,781,1915,1941]" captionStart-0="FIGURE 9" captionStart-1="FIGURE 10" captionStartId-0="16.[151,250,1981,2005]" captionStartId-1="17.[151,250,1191,1215]" captionTargetBox-0="[168,1415,786,1947]" captionTargetBox-1="[156,1434,187,1159]" captionTargetId-0="figure-368@16.[164,1423,777,1958]" captionTargetId-1="figure-19@17.[151,1436,181,1167]" captionTargetPageId-0="16" captionTargetPageId-1="17" captionText-0="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." captionText-1="FIGURE 10. Diagnostic characters of Athyma inara sahyadriensis ssp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the subspecies. Male of A. i. sahyadriensis ssp. nov. is compared with males of A. i. inara and two subspecies of A. nefte. Numbers point to distinctive characters: 1: FW subapical markings, 2: FW submarginal markings, 3: FW spot in interspace 3, 4: FW markings in the discal cell, 5: HW post-discal band, 6: UNF markings in the discal cell, 7: UN base colour of the wings, 8: UNF submarginal markings, 9: large white spot in interspace 2, 10: UNH post-discal band (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte." figureDoi-0="http://doi.org/10.5281/zenodo.14386322" figureDoi-1="http://doi.org/10.5281/zenodo.14386326" httpUri-0="https://zenodo.org/record/14386322/files/figure.png" httpUri-1="https://zenodo.org/record/14386326/files/figure.png" pageId="2" pageNumber="345">Fig. 9–10</figureCitation>
), BMNH(E) #933429 (
<typeStatus id="367504E3DA1C78322538861FA8BBF841" box="[1051,1157,1915,1941]" pageId="2" pageNumber="345" type="holotype">Holotype</typeStatus>
of
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<emphasis id="DBBA6653DA1C78322592861FA93DF840" box="[1201,1283,1915,1940]" italics="true" pageId="2" pageNumber="345">inarina</emphasis>
</taxonomicName>
. “
<collectingRegion id="2B0A74A3DA1C7832243C861FA94EF841" box="[1311,1392,1915,1941]" country="India" name="Assam" pageId="2" pageNumber="345">Assam</collectingRegion>
” as per a specimen label; border of
<collectingRegion id="2B0A74A3DA1C7832230E86FBAE47F86D" box="[557,633,1951,1977]" country="India" name="Assam" pageId="2" pageNumber="345">Assam</collectingRegion>
and
<collectingRegion id="2B0A74A3DA1C7832239186FBAF2AF86D" box="[690,788,1951,1977]" country="India" name="Manipur" pageId="2" pageNumber="345">Manipur</collectingRegion>
, NE
<collectingCountry id="91D9FAD1DA1C7832226E86FBAFB7F86D" box="[845,905,1951,1977]" name="India" pageId="2" pageNumber="345">India</collectingCountry>
, as per original description.
<figureCitation id="71F5A6C4DA1C783225E986FBA930F86D" box="[1226,1294,1951,1977]" captionStart="FIGURE 9" captionStartId="16.[151,250,1981,2005]" captionTargetBox="[168,1415,786,1947]" captionTargetId="figure-368@16.[164,1423,777,1958]" captionTargetPageId="16" captionText="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386322" httpUri="https://zenodo.org/record/14386322/files/figure.png" pageId="2" pageNumber="345">Fig. 9</figureCitation>
), BMNH(E) #933424 (
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of
<taxonomicName id="2ECEC1C2DA1C783220E686A7ADC1F808" authorityName="Swinhoe" authorityYear="1917" box="[453,511,1987,2012]" class="Insecta" family="Nymphalidae" genus="Pantoporia" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="345" phylum="Arthropoda" rank="species" species="mera">
<emphasis id="DBBA6653DA1C783220E686A7ADC1F808" box="[453,511,1987,2012]" italics="true" pageId="2" pageNumber="345">mera</emphasis>
</taxonomicName>
. “Borneo”, in error, collected almost certainly from either NE
<collectingCountry id="91D9FAD1DA1C7832259486A7A8CFF809" box="[1207,1265,1987,2013]" name="India" pageId="2" pageNumber="345">India</collectingCountry>
or N
<collectingCountry id="91D9FAD1DA1C7832240C86A7A9A2F809" box="[1327,1436,1987,2013]" name="Myanmar" pageId="2" pageNumber="345">Myanmar</collectingCountry>
based on the phenotype.
<figureCitation id="71F5A6C4DA1D783320D081F3AE02FF65" box="[499,572,151,177]" captionStart="FIGURE 9" captionStartId="16.[151,250,1981,2005]" captionTargetBox="[168,1415,786,1947]" captionTargetId="figure-368@16.[164,1423,777,1958]" captionTargetPageId="16" captionText="FIGURE 9. Comparison of Athyma inara sahyadriensis ssp. nov. with other similar Athyma (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386322" httpUri="https://zenodo.org/record/14386322/files/figure.png" pageId="3" pageNumber="346">Fig. 9</figureCitation>
), BMNH(E) #933502 (Darjeeling,
<collectingRegion id="2B0A74A3DA1D783322FF81F3A84CFF65" box="[988,1138,151,177]" country="India" name="West Bengal" pageId="3" pageNumber="346">West Bengal</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1D783325A081F3A8FEFF65" box="[1155,1216,151,177]" name="India" pageId="3" pageNumber="346">India</collectingCountry>
. 1894), BMNH(E) #933423 (
<collectingRegion id="2B0A74A3DA1D7833201581DFADB2FF01" box="[310,396,187,213]" country="India" name="Sikkim" pageId="3" pageNumber="346">Sikkim</collectingRegion>
. 1893), BMNH(E) #933496 (
<collectingRegion id="2B0A74A3DA1D783323ED81DFAF1AFF01" box="[718,804,187,213]" country="India" name="Sikkim" pageId="3" pageNumber="346">Sikkim</collectingRegion>
. 1893), BMNH(E) #933472 (Kurseong, near Darjeeling), BMNH(E) #933481 (Darjeeling. 1894).
<emphasis id="DBBA6653DA1D7833238B8184AEF8FF28" bold="true" box="[680,710,224,252]" pageId="3" pageNumber="346">
<specimenCount id="FFC871C8DA1D7833238B8184AE81FF28" box="[680,703,224,252]" count="1" pageId="3" pageNumber="346" type="female">♀</specimenCount>
:
</emphasis>
BMNH(E) #933428 (Darjeeling. 1894), BMNH(E) #933461 (Darjeeling. 1909), BMNH(E) #933418 (‘
<collectingRegion id="2B0A74A3DA1D7833238E8067AF39FEC9" box="[685,775,259,285]" country="India" name="Sikkim" pageId="3" pageNumber="346">Sikkim</collectingRegion>
, Singlah’, likely from Darjeeling/ Singalila area. 1894), BMNH(E) #933425 (‘
<collectingRegion id="2B0A74A3DA1D7833209D8043AE26FE95" box="[446,536,295,321]" country="India" name="Sikkim" pageId="3" pageNumber="346">Sikkim</collectingRegion>
, Teesta’. 1927), BMNH(E) #933400 (Silhet,
<collectingCountry id="91D9FAD1DA1D783325398043A89EFE95" box="[1050,1184,295,321]" name="Bangladesh" pageId="3" pageNumber="346">Bangladesh</collectingCountry>
. 1909). All specimens in NHMUK
</materialsCitation>
.
</paragraph>
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144
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13.
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<emphasis id="DBBA6653DA1D783321E4800AAD5FFE5D" bold="true" box="[199,353,366,393]" italics="true" pageId="3" pageNumber="346">Athyma asita</emphasis>
Moore, 1858
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:
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:
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(E) #933438 (
<typeStatus id="367504E3DA1D78332217800BAF9FFE5D" box="[820,929,367,393]" pageId="3" pageNumber="346" type="holotype">Holotype</typeStatus>
. Collected likely from S
<collectingCountry id="91D9FAD1DA1D783325EB800BA90BFE5D" box="[1224,1333,367,393]" name="Myanmar" pageId="3" pageNumber="346">Myanmar</collectingCountry>
or
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),
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(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1D7833232680F7A865FE79" box="[517,1115,403,429]" collectionCode="BMNH" country="Myanmar" location="Karen Hills" pageId="3" pageNumber="346" specimenCount="1" typeStatus="holotype">
933494 (
<location id="EC11EC9ADA1D7833234A80F7AECFFE79" LSID="urn:lsid:plazi:treatment:61670B57DA1D783321B4800BA895FE21:EC11EC9ADA1D7833234A80F7AECFFE79" box="[617,753,403,429]" country="Myanmar" name="Karen Hills" pageId="3" pageNumber="346">Karen Hills</location>
, S
<collectingCountry id="91D9FAD1DA1D7833223580F7AFBBFE79" box="[790,901,403,429]" name="Myanmar" pageId="3" pageNumber="346">Myanmar</collectingCountry>
. 1923),
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(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1D7833255280F7AD01FE05" collectionCode="BMNH" country="Myanmar" location="Gokteik'" pageId="3" pageNumber="346" specimenCount="1" typeStatus="holotype">
933493 (‘
<location id="EC11EC9ADA1D783325F880F7A976FE79" LSID="urn:lsid:plazi:treatment:61670B57DA1D783321B4800BA895FE21:EC11EC9ADA1D783325F880F7A976FE79" box="[1243,1352,403,429]" country="Myanmar" name="Gokteik'" pageId="3" pageNumber="346">Gokteik’</location>
. 1924),
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(E)
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#
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933495 (
<location id="EC11EC9ADA1D7833209B80D3AE72FE05" LSID="urn:lsid:plazi:treatment:61670B57DA1D783321B4800BA895FE21:EC11EC9ADA1D7833209B80D3AE72FE05" box="[440,588,439,465]" country="Myanmar" name="Tavoy valley" pageId="3" pageNumber="346">Tavoy valley</location>
,
<collectingCountry id="91D9FAD1DA1D7833237B80D3AEF9FE05" box="[600,711,439,465]" name="Myanmar" pageId="3" pageNumber="346">Myanmar</collectingCountry>
. 1893).
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<specimenCount id="FFC871C8DA1D7833220180DCAF07FE00" box="[802,825,440,468]" count="1" pageId="3" pageNumber="346" type="female">♀</specimenCount>
:
</emphasis>
<collectionCode id="8FDF2284DA1D7833226B80D3AFA1FE05" box="[840,927,439,465]" pageId="3" pageNumber="346">BMNH</collectionCode>
(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1D783322F580D3ADA6FE21" collectionCode="BMNH" country="Myanmar" location="Karen Hills" pageId="3" pageNumber="346" specimenCount="1" typeStatus="holotype">
933466 (
<location id="EC11EC9ADA1D7833251A80D3A8FEFE05" LSID="urn:lsid:plazi:treatment:61670B57DA1D783321B4800BA895FE21:EC11EC9ADA1D7833251A80D3A8FEFE05" box="[1081,1216,439,465]" country="Myanmar" name="Karen Hills" pageId="3" pageNumber="346">Karen Hills</location>
, S
<collectingCountry id="91D9FAD1DA1D783325C080D3A96CFE05" box="[1251,1362,439,465]" name="Myanmar" pageId="3" pageNumber="346">Myanmar</collectingCountry>
. 1923.
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),
<collectionCode id="8FDF2284DA1D7833200380BFAD49FE21" box="[288,375,475,501]" pageId="3" pageNumber="346">BMNH</collectionCode>
(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1D7833208E80BEA896FE21" box="[429,1192,474,501]" collectionCode="NHMUK" collectorName="All" country="Myanmar" location="Location" pageId="3" pageNumber="346" specimenCount="1" typeStatus="holotype">
933452 (
<location id="EC11EC9ADA1D7833232C80BFAE4AFE21" LSID="urn:lsid:plazi:treatment:61670B57DA1D783321B4800BA895FE21:EC11EC9ADA1D7833232C80BFAE4AFE21" box="[527,628,475,501]" country="Myanmar" name="Location" pageId="3" pageNumber="346">Location</location>
name illegible. 1922).
<collectorName id="443BDF97DA1D7833225480BFAFA5FE21" box="[887,923,475,501]" pageId="3" pageNumber="346">All</collectorName>
specimens in
<collectionCode id="8FDF2284DA1D7833251980BFA896FE21" box="[1082,1192,475,501]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" lsid="urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="3" pageNumber="346" type="Museum">NHMUK</collectionCode>
</materialsCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>

122
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14.
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:
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<collectionCode id="8FDF2284DA1D783320EE809BAE1AFDCD" box="[461,548,511,537]" pageId="3" pageNumber="346">BMNH</collectionCode>
(E)
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#
<materialsCitation id="59A6B01CDA1D78332374809BA83BFDCD" box="[599,1029,511,537]" collectionCode="BMNH" location="Java" pageId="3" pageNumber="346" specimenCount="1" stateProvince="Jawa Timur">
933413 (
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),
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(E)
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#
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933401 (
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. 1912).
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<specimenCount id="FFC871C8DA1D783324218364A927FDC8" box="[1282,1305,512,540]" count="1" pageId="3" pageNumber="346" type="female">♀</specimenCount>
:
</emphasis>
<collectionCode id="8FDF2284DA1D78332407809BA945FDCD" box="[1316,1403,511,537]" pageId="3" pageNumber="346">BMNH</collectionCode>
(E)
</materialsCitation>
#
<materialsCitation id="59A6B01CDA1D783321F68346AEAEFDE9" box="[213,656,546,573]" collectionCode="BMNH" location="Java" pageId="3" pageNumber="346" specimenCount="1" stateProvince="Jawa Timur">
933454 (
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. 1912.
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),
<collectionCode id="8FDF2284DA1D7833233B8347AE51FDE9" box="[536,623,547,573]" pageId="3" pageNumber="346">BMNH</collectionCode>
(E)
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#
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933440 (
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specimens in
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15.
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<taxonomicName id="2ECEC1C2DA1D783321E48322AE4AFDB4" authority="Moore, 1858" authorityName="Moore" authorityYear="1858" box="[199,628,582,609]" class="Insecta" family="Nymphalidae" genus="Athyma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="3" pageNumber="346" phylum="Arthropoda" rank="subSpecies" species="nefte" subSpecies="subrata">
<emphasis id="DBBA6653DA1D783321E48322ADF4FDB5" bold="true" box="[199,458,582,609]" italics="true" pageId="3" pageNumber="346">Athyma nefte subrata</emphasis>
Moore, 1858
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:
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This includes the following synonyms (
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): (a)
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<emphasis id="DBBA6653DA1D783324648323AD20FD51" italics="true" pageId="3" pageNumber="346">Athyma nivifera</emphasis>
Butler, 1879
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, and (b)
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<emphasis id="DBBA6653DA1D7833233E830FAF71FD51" box="[541,847,619,645]" italics="true" pageId="3" pageNumber="346">Pantoporia nefte subratina</emphasis>
Fruhstorfer, 1912
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.
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:
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BMNH(E) #933480 (
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<specimenCount id="FFC871C8DA1D7833246B8308A963FD5C" box="[1352,1373,620,648]" count="1" pageId="3" pageNumber="346" type="male">♂</specimenCount>
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,
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.
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), BMNH(E) #933448 (
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,
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).
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:
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BMNH(E) #933457 (
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<emphasis id="DBBA6653DA1D7833209183D7AE37FD19" box="[434,521,691,717]" italics="true" pageId="3" pageNumber="346">nivifera</emphasis>
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.
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,
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.
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), BMNH(E) #933431 (
<collectingRegion id="2B0A74A3DA1D7833256183D7A89AFD19" box="[1090,1188,691,717]" country="Indonesia" name="Sumatera Utara" pageId="3" pageNumber="346">Sumatra</collectingRegion>
,
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. 1903.
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), BMNH(E) #933501 (
<collectingRegion id="2B0A74A3DA1D783320FF83B3AE00FD25" box="[476,574,727,753]" country="Indonesia" name="Sumatera Utara" pageId="3" pageNumber="346">Sumatra</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1D7833236A83B3AE84FD25" box="[585,698,727,753]" name="Indonesia" pageId="3" pageNumber="346">Indonesia</collectingCountry>
. 1903). All specimens in NHMUK.
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<heading id="B2390D2DDA1D783321B484E6AE48FA49" bold="true" box="[151,630,1410,1437]" fontSize="11" level="1" pageId="3" pageNumber="346" reason="1">
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<emphasis id="DBBA6653DA1D783321B484E6AD82FA49" bold="true" box="[151,444,1410,1437]" italics="true" pageId="3" pageNumber="346">Amathusia travancorica</emphasis>
Kunte
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(
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):
</emphasis>
Voucher code IBC-PW981.
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. Thenmala FRH, near Shendurney WLS, Kollam District,
<collectingRegion id="2B0A74A3DA1D7833246884A0A9A6FA0A" box="[1355,1432,1476,1502]" country="India" name="Kerala" pageId="3" pageNumber="346">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1D783321B4848CACEFF9D6" box="[151,209,1512,1538]" name="India" pageId="3" pageNumber="346">India</collectingCountry>
(lat-long:
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,
<geoCoordinate id="8CFADC86DA1D783320E6848CAE01F9D6" box="[453,575,1512,1538]" degrees="77.0626" direction="east" orientation="longitude" pageId="3" pageNumber="346" precision="5" value="77.0626">77.0626 E</geoCoordinate>
). Approx.
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asl. Raised from caterpillar on a
<taxonomicName id="2ECEC1C2DA1D78332556848CA8F5F9D6" box="[1141,1227,1512,1538]" class="Liliopsida" family="Arecaceae" genus="Licuala" kingdom="Plantae" order="Arecales" pageId="3" pageNumber="346" phylum="Tracheophyta" rank="genus">
<emphasis id="DBBA6653DA1D78332556848CA8F5F9D6" box="[1141,1227,1512,1538]" italics="true" pageId="3" pageNumber="346">Licuala</emphasis>
</taxonomicName>
palm (
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) on
<date id="9D709C81DA1D7833219E8768AD01F9F2" box="[189,319,1548,1574]" pageId="3" pageNumber="346" value="2012-12-19">2012/12/19</date>
, eclosed and preserved on
<date id="9D709C81DA1D7833235B8768AEC1F9F2" box="[632,767,1548,1574]" pageId="3" pageNumber="346" value="2013-01-14">2013/01/14.</date>
Collected by Krushnamegh Kunte. Preserved dry, pinned, with thoracic muscle tissue and two legs (mid- and hind-leg on the right side) preserved in 100% molecular-grade ethanol. Deposited in the Biodiversity Lab Research Collections at NCBS.
</paragraph>
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<emphasis id="DBBA6653DA1D783321E4871CAD86F946" bold="true" box="[199,440,1656,1682]" pageId="3" pageNumber="346">
Description (
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<emphasis id="DBBA6653DA1D783321E487F9ADBCF962" box="[199,386,1692,1718]" italics="true" pageId="3" pageNumber="346">Forewing length</emphasis>
:
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.
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:
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. Head brown above; palps white; eyes pale brown with vertical dark bands (eyes turn dark brown upon preservation). Antennae pale reddish-brown. Proboscis brown. Thorax and abdomen brown above, white on thorax and pale yellow on abdomen below. Abdomen with tufts of long, dense scales on the side, which are longest (
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) on segments 9–12, almost black above, yellow below. Legs pale brown.
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Wings broad.
<emphasis id="DBBA6653DA1D783320428635ADF3F8BE" box="[353,461,1873,1898]" italics="true" pageId="3" pageNumber="346">Forewing</emphasis>
termen straight, not concave; dorsum straight.
<emphasis id="DBBA6653DA1D783322F08634A87FF8BE" box="[979,1089,1872,1898]" italics="true" pageId="3" pageNumber="346">Hindwing</emphasis>
very broad and rounded except the tornus, which is produced into a lobe. Three tufts of long, black scent scales: (a) along v1b, (b) in an adjoining fold in interspace 1c, and (c) between v7 and v8.
</paragraph>
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<emphasis id="DBBA6653DA1A783421B48624AD2AF88C" bold="true" box="[151,276,1856,1880]" pageId="4" pageNumber="347">FIGURE 1.</emphasis>
Type specimens of
<emphasis id="DBBA6653DA1A783420C18624AF07F88C" bold="true" box="[482,825,1856,1880]" pageId="4" pageNumber="347">
<taxonomicName id="2ECEC1C2DA1A783420C18624AEDEF88C" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[482,736,1856,1880]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="347" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1A783420C18624AEDEF88C" bold="true" box="[482,736,1856,1880]" italics="true" pageId="4" pageNumber="347">Amathusia travancorica</emphasis>
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<taxonomicNameLabel id="C089DB28DA1A783423C48624AF07F88C" box="[743,825,1856,1880]" pageId="4" pageNumber="347" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu and Krushnamegh Kunte.
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<paragraph id="E971BA41DA1B783521E481F3AE52FEC9" blockId="5.[151,1437,151,537]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E481F3ADE8FF64" box="[199,470,151,177]" italics="true" pageId="5" pageNumber="348">Dorsal side: Forewings</emphasis>
pale rusty-brown, without markings, but markings from the ventral side are visible faintly through the wings; i.e., markings from the ventral side may show through on the dorsal side.
<emphasis id="DBBA6653DA1B7835240081DFA9A2FF01" box="[1315,1436,187,213]" italics="true" pageId="5" pageNumber="348">Hindwings</emphasis>
pale rusty-brown, same as the forewing; dorsum area that covers the abdomen is white. Tornal lobe with two small, white-bordered black spots on either side.
</paragraph>
<paragraph id="E971BA41DA1B783521E48043AEADFDCD" blockId="5.[151,1437,151,537]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E48043AD68FE95" box="[199,342,295,321]" italics="true" pageId="5" pageNumber="348">Ventral side:</emphasis>
pale yellowish-brown with dark brown and white bands spanning across forewing and hindwing, similar to other members of the genus: three dark bands at the wing bases and in the discal cell, alternating with white bands; a pale broad band just beyond the discal cells; a white-bordered dark brown, broad band from FW costa to the large post-discal eyespot on HW (this band is the darkest), followed by a very broad pale brown band and a paler yellowish-brown submarginal band. The terminal margin is dark brown with a pale line on it. There are two large eyespots, one in interspace 2, and the other in interspace 6. Tornal lobe has two small, white-bordered black spots on either side, similar to the upper side.
</paragraph>
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</treatment>
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810
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<heading id="B2390D2DDA1D783321B484E6AE48FA49" bold="true" box="[151,630,1410,1437]" fontSize="11" level="1" pageId="3" pageNumber="346" reason="1">
<emphasis id="DBBA6653DA1D783321B484E6AE48FA49" bold="true" box="[151,630,1410,1437]" pageId="3" pageNumber="346">
<taxonomicName id="2ECEC1C2DA1D783321B484E6AE2FFA49" authority="Kunte" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[151,529,1410,1437]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="3" pageNumber="346" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1D783321B484E6AD82FA49" bold="true" box="[151,444,1410,1437]" italics="true" pageId="3" pageNumber="346">Amathusia travancorica</emphasis>
Kunte
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1D7833233A84E7AE48FA49" box="[537,630,1411,1437]" pageId="3" pageNumber="346" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1D783321B484A0AFE4F9BA" pageId="3" pageNumber="346" type="materials_examined">
<paragraph id="E971BA41DA1D783321B484A0AFE4F9BA" blockId="3.[151,1437,1476,1971]" pageId="3" pageNumber="346">
<materialsCitation id="59A6B01CDA1D783321B484A0AFE4F9BA" collectingDate="2012-12-19" collectorName="Krushnamegh Kunte" country="India" elevation="90" latitude="8.9594" location="Thenmala FRH" longLatPrecision="7" longitude="77.0626" municipality="Kollam District" pageId="3" pageNumber="346" specimenCode="IBC-PW981" specimenCount="1" specimenCount-male="1" stateProvince="Kerala" typeStatus="holotype">
<emphasis id="DBBA6653DA1D783321B484A0AD57FA0A" bold="true" box="[151,361,1476,1502]" pageId="3" pageNumber="346">
<typeStatus id="367504E3DA1D783321B484A0AD3AFA0A" box="[151,260,1476,1502]" pageId="3" pageNumber="346" type="holotype">Holotype</typeStatus>
(
<figureCitation id="71F5A6C4DA1D7833203184A0AD69FA0A" box="[274,343,1476,1502]" captionStart="FIGURE 1" captionStartId="4.[151,250,1856,1880]" captionTargetBox="[385,1197,186,1823]" captionTargetId="figure-20@4.[381,1207,181,1832]" captionTargetPageId="4" captionText="FIGURE 1. Type specimens of Amathusia travancorica sp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386301" httpUri="https://zenodo.org/record/14386301/files/figure.png" pageId="3" pageNumber="346">Fig. 1</figureCitation>
):
</emphasis>
Voucher code
<specimenCode id="B968123ADA1D7833232D84A0AEA0FA0A" box="[526,670,1476,1502]" pageId="3" pageNumber="346">IBC-PW981</specimenCode>
.
<specimenCount id="FFC871C8DA1D7833238484A1AE81FA0B" box="[679,703,1477,1503]" count="1" pageId="3" pageNumber="346" type="male">♂</specimenCount>
.
<location id="EC11EC9ADA1D783323EA84A0AF47FA0B" LSID="urn:lsid:plazi:treatment:61670B57DA1D783C21B484E6AFFEFC55:EC11EC9ADA1D783323EA84A0AF47FA0B" box="[713,889,1476,1503]" country="India" latitude="8.9594" longLatPrecision="7" longitude="77.0626" municipality="Kollam District" name="Thenmala FRH" pageId="3" pageNumber="346" stateProvince="Kerala">Thenmala FRH</location>
, near
<location id="EC11EC9ADA1D7833229984A0A8BAFA0A" LSID="urn:lsid:plazi:treatment:61670B57DA1D783C21B484E6AFFEFC55:EC11EC9ADA1D7833229984A0A8BAFA0A" box="[954,1156,1476,1502]" country="India" latitude="8.9594" longLatPrecision="7" longitude="77.0626" municipality="Kollam District" name="Shendurney WLS" pageId="3" pageNumber="346" stateProvince="Kerala">Shendurney WLS</location>
,
<collectingMunicipality id="0915203BDA1D783325AC84A0A97EFA0A" box="[1167,1344,1476,1502]" pageId="3" pageNumber="346">Kollam District</collectingMunicipality>
,
<collectingRegion id="2B0A74A3DA1D7833246884A0A9A6FA0A" box="[1355,1432,1476,1502]" country="India" name="Kerala" pageId="3" pageNumber="346">Kerala</collectingRegion>
,
<collectingCountry id="91D9FAD1DA1D783321B4848CACEFF9D6" box="[151,209,1512,1538]" name="India" pageId="3" pageNumber="346">India</collectingCountry>
(lat-long:
<geoCoordinate id="8CFADC86DA1D78332068848CAD86F9D7" box="[331,440,1512,1539]" degrees="8.9594" direction="north" orientation="latitude" pageId="3" pageNumber="346" precision="5" value="8.9594">8.9594 N</geoCoordinate>
,
<geoCoordinate id="8CFADC86DA1D783320E6848CAE01F9D6" box="[453,575,1512,1538]" degrees="77.0626" direction="east" orientation="longitude" pageId="3" pageNumber="346" precision="5" value="77.0626">77.0626 E</geoCoordinate>
). Approx.
<quantity id="2E3617A4DA1D7833239B848CAECDF9D7" box="[696,755,1512,1539]" metricMagnitude="1" metricUnit="m" metricValue="9.0" pageId="3" pageNumber="346" unit="m" value="90.0">
<elevation id="62E35D72DA1D7833239B848CAECDF9D7" box="[696,755,1512,1539]" metricMagnitude="1" metricUnit="m" metricValue="9.0" pageId="3" pageNumber="346" unit="m" value="90.0">90 m</elevation>
</quantity>
asl. Raised from caterpillar on a
<taxonomicName id="2ECEC1C2DA1D78332556848CA8F5F9D6" box="[1141,1227,1512,1538]" class="Liliopsida" family="Arecaceae" genus="Licuala" kingdom="Plantae" order="Arecales" pageId="3" pageNumber="346" phylum="Tracheophyta" rank="genus">
<emphasis id="DBBA6653DA1D78332556848CA8F5F9D6" box="[1141,1227,1512,1538]" italics="true" pageId="3" pageNumber="346">Licuala</emphasis>
</taxonomicName>
palm (
<taxonomicName id="2ECEC1C2DA1D7833243E848CA9ABF9D6" box="[1309,1429,1512,1538]" class="Liliopsida" family="Arecaceae" kingdom="Plantae" order="Arecales" pageId="3" pageNumber="346" phylum="Tracheophyta" rank="family">Arecaceae</taxonomicName>
) on
<date id="9D709C81DA1D7833219E8768AD01F9F2" box="[189,319,1548,1574]" pageId="3" pageNumber="346" value="2012-12-19">
<collectingDate id="8D346569DA1D7833219E8768AD01F9F2" box="[189,319,1548,1574]" pageId="3" pageNumber="346" value="2012-12-19">2012/12/19</collectingDate>
</date>
, eclosed and preserved on
<date id="9D709C81DA1D7833235B8768AEC1F9F2" box="[632,767,1548,1574]" pageId="3" pageNumber="346" value="2013-01-14">2013/01/14.</date>
Collected by
<collectorName id="443BDF97DA1D783322818768A8AEF9F3" box="[930,1168,1548,1575]" pageId="3" pageNumber="346">Krushnamegh Kunte</collectorName>
. Preserved dry, pinned, with thoracic muscle tissue and two legs (mid- and hind-leg on the right side) preserved in 100% molecular-grade ethanol. Deposited in the Biodiversity Lab Research Collections at NCBS.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1D783521E4871CAEADFDCD" lastPageId="5" lastPageNumber="348" pageId="3" pageNumber="346" type="description">
<paragraph id="E971BA41DA1D783321E4871CAD86F946" blockId="3.[151,1437,1476,1971]" box="[199,440,1656,1682]" pageId="3" pageNumber="346">
<emphasis id="DBBA6653DA1D783321E4871CAD86F946" bold="true" box="[199,440,1656,1682]" pageId="3" pageNumber="346">
Description (
<figureCitation id="71F5A6C4DA1D78332042871CAD98F946" box="[353,422,1656,1682]" captionStart="FIGURE 1" captionStartId="4.[151,250,1856,1880]" captionTargetBox="[385,1197,186,1823]" captionTargetId="figure-20@4.[381,1207,181,1832]" captionTargetPageId="4" captionText="FIGURE 1. Type specimens of Amathusia travancorica sp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu and Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386301" httpUri="https://zenodo.org/record/14386301/files/figure.png" pageId="3" pageNumber="346">Fig. 1</figureCitation>
):
</emphasis>
</paragraph>
<paragraph id="E971BA41DA1D783321E487F9AD67F892" blockId="3.[151,1437,1476,1971]" pageId="3" pageNumber="346">
<emphasis id="DBBA6653DA1D783321E487F9ADBCF962" box="[199,386,1692,1718]" italics="true" pageId="3" pageNumber="346">Forewing length</emphasis>
:
<quantity id="2E3617A4DA1D783320B187F8ADDBF963" box="[402,485,1692,1719]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="3" pageNumber="346" unit="mm" value="50.0">50 mm</quantity>
.
<emphasis id="DBBA6653DA1D783320D287F8AEFEF962" box="[497,704,1692,1718]" italics="true" pageId="3" pageNumber="346">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA1D783323F387F8AF1AF963" box="[720,804,1692,1719]" metricMagnitude="-2" metricUnit="m" metricValue="2.2" pageId="3" pageNumber="346" unit="mm" value="22.0">22 mm</quantity>
. Head brown above; palps white; eyes pale brown with vertical dark bands (eyes turn dark brown upon preservation). Antennae pale reddish-brown. Proboscis brown. Thorax and abdomen brown above, white on thorax and pale yellow on abdomen below. Abdomen with tufts of long, dense scales on the side, which are longest (
<quantity id="2E3617A4DA1D783323EC866CAF17F8F6" box="[719,809,1800,1826]" metricMagnitude="-3" metricUnit="m" metricValue="7.5" metricValueMax="8.0" metricValueMin="7.0" pageId="3" pageNumber="346" unit="mm" value="7.5" valueMax="8.0" valueMin="7.0">7-8 mm</quantity>
) on segments 9–12, almost black above, yellow below. Legs pale brown.
</paragraph>
<paragraph id="E971BA41DA1D783321E48634AE86F866" blockId="3.[151,1437,1476,1971]" pageId="3" pageNumber="346">
Wings broad.
<emphasis id="DBBA6653DA1D783320428635ADF3F8BE" box="[353,461,1873,1898]" italics="true" pageId="3" pageNumber="346">Forewing</emphasis>
termen straight, not concave; dorsum straight.
<emphasis id="DBBA6653DA1D783322F08634A87FF8BE" box="[979,1089,1872,1898]" italics="true" pageId="3" pageNumber="346">Hindwing</emphasis>
very broad and rounded except the tornus, which is produced into a lobe. Three tufts of long, black scent scales: (a) along v1b, (b) in an adjoining fold in interspace 1c, and (c) between v7 and v8.
</paragraph>
<caption id="BDB1EAC9DA1A783421B48624AE32F8A8" ID-DOI="http://doi.org/10.5281/zenodo.14386301" ID-Zenodo-Dep="14386301" httpUri="https://zenodo.org/record/14386301/files/figure.png" pageId="4" pageNumber="347" startId="4.[151,250,1856,1880]" targetBox="[385,1197,186,1823]" targetPageId="4" targetType="figure">
<paragraph id="E971BA41DA1A783421B48624AE32F8A8" blockId="4.[151,1436,1856,1916]" pageId="4" pageNumber="347">
<emphasis id="DBBA6653DA1A783421B48624AD2AF88C" bold="true" box="[151,276,1856,1880]" pageId="4" pageNumber="347">FIGURE 1.</emphasis>
Type specimens of
<emphasis id="DBBA6653DA1A783420C18624AF07F88C" bold="true" box="[482,825,1856,1880]" pageId="4" pageNumber="347">
<taxonomicName id="2ECEC1C2DA1A783420C18624AEDEF88C" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[482,736,1856,1880]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="4" pageNumber="347" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1A783420C18624AEDEF88C" bold="true" box="[482,736,1856,1880]" italics="true" pageId="4" pageNumber="347">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1A783423C48624AF07F88C" box="[743,825,1856,1880]" pageId="4" pageNumber="347" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu and Krushnamegh Kunte.
</paragraph>
</caption>
<paragraph id="E971BA41DA1B783521E481F3AE52FEC9" blockId="5.[151,1437,151,537]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E481F3ADE8FF64" box="[199,470,151,177]" italics="true" pageId="5" pageNumber="348">Dorsal side: Forewings</emphasis>
pale rusty-brown, without markings, but markings from the ventral side are visible faintly through the wings; i.e., markings from the ventral side may show through on the dorsal side.
<emphasis id="DBBA6653DA1B7835240081DFA9A2FF01" box="[1315,1436,187,213]" italics="true" pageId="5" pageNumber="348">Hindwings</emphasis>
pale rusty-brown, same as the forewing; dorsum area that covers the abdomen is white. Tornal lobe with two small, white-bordered black spots on either side.
</paragraph>
<paragraph id="E971BA41DA1B783521E48043AEADFDCD" blockId="5.[151,1437,151,537]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E48043AD68FE95" box="[199,342,295,321]" italics="true" pageId="5" pageNumber="348">Ventral side:</emphasis>
pale yellowish-brown with dark brown and white bands spanning across forewing and hindwing, similar to other members of the genus: three dark bands at the wing bases and in the discal cell, alternating with white bands; a pale broad band just beyond the discal cells; a white-bordered dark brown, broad band from FW costa to the large post-discal eyespot on HW (this band is the darkest), followed by a very broad pale brown band and a paler yellowish-brown submarginal band. The terminal margin is dark brown with a pale line on it. There are two large eyespots, one in interspace 2, and the other in interspace 6. Tornal lobe has two small, white-bordered black spots on either side, similar to the upper side.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1B783521B4830FAFD3FCC9" pageId="5" pageNumber="348" type="materials_examined">
<materialsCitation id="59A6B01CDA1B783521B4830FAFCDFD15" collectionCode="IBC" pageId="5" pageNumber="348" specimenCode="IBC- PW982" specimenCount="1" specimenCount-male="1" typeStatus="paratype">
<paragraph id="E971BA41DA1B783521B4830FAED3FD50" blockId="5.[151,749,618,645]" box="[151,749,618,645]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521B4830FAED3FD50" bold="true" box="[151,749,618,645]" pageId="5" pageNumber="348">
<typeStatus id="367504E3DA1B783521B4830FAD28FD51" box="[151,278,619,645]" pageId="5" pageNumber="348" type="paratype">Paratypes</typeStatus>
, and intraspecific variation (
<figureCitation id="71F5A6C4DA1B7835235D830EAEDDFD50" box="[638,739,618,644]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStartId-0="4.[151,250,1856,1880]" captionStartId-1="6.[151,250,1817,1841]" captionStartId-2="7.[151,250,1957,1981]" captionTargetBox-0="[385,1197,186,1823]" captionTargetBox-1="[151,1436,417,1793]" captionTargetBox-2="[155,1436,188,1922]" captionTargetId-0="figure-20@4.[381,1207,181,1832]" captionTargetId-1="figure-151@6.[151,1436,417,1793]" captionTargetId-2="figure-19@7.[151,1436,181,1933]" captionTargetPageId-0="4" captionTargetPageId-1="6" captionTargetPageId-2="7" captionText-0="FIGURE 1. Type specimens of Amathusia travancorica sp. nov. A millimetre scale is at the bottom. Photographs: Dipendra Nath Basu and Krushnamegh Kunte." captionText-1="FIGURE 2. Male genitalia of Amathusia travancorica sp. nov. (paratype, IBC-PW982). a: lateral view of genital capsule or armature showing the inside of the left valve (right valve removed), b: dorsal view of genital capsule, c: outside of the right valve, d: Phallus (aedeagus), e: dorsal view of the tip of uncus, along with its shape tracing, and f: shape tracings of dorsal view of other Indo-Malayan Amathusia. Photographs: Dipendra Nath Basu and Ujwala Pawar. Genitalia tracings are redrawn from Uémura (1986), except that of A. travancorica sp. nov. (this study)." captionText-2="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." figureDoi-0="http://doi.org/10.5281/zenodo.14386301" figureDoi-1="http://doi.org/10.5281/zenodo.14386303" figureDoi-2="http://doi.org/10.5281/zenodo.14386305" httpUri-0="https://zenodo.org/record/14386301/files/figure.png" httpUri-1="https://zenodo.org/record/14386303/files/figure.png" httpUri-2="https://zenodo.org/record/14386305/files/figure.png" pageId="5" pageNumber="348">Fig. 1–3</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="E971BA41DA1B783521B483CEAFCDFD15" blockId="5.[151,1436,682,798]" box="[151,1011,682,706]" pageId="5" pageNumber="348">
<specimenCount id="FFC871C8DA1B783521B483CEAC93FD15" box="[151,173,682,705]" count="1" pageId="5" pageNumber="348" type="male">♂</specimenCount>
: voucher code
<specimenCode id="B968123ADA1B7835206583CEADF0FD16" box="[326,462,682,706]" pageId="5" pageNumber="348">IBC- PW982</specimenCode>
. Forewing length:
<quantity id="2E3617A4DA1B783523A883CEAEE8FD15" box="[651,726,682,706]" metricMagnitude="-2" metricUnit="m" metricValue="4.9" pageId="5" pageNumber="348" unit="mm" value="49.0">49 mm</quantity>
.
<emphasis id="DBBA6653DA1B783523FC83CEAFA6FD16" box="[735,920,682,706]" italics="true" pageId="5" pageNumber="348">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA1B7835228683CEAFCEFD15" box="[933,1008,682,706]" metricMagnitude="-2" metricUnit="m" metricValue="2.1" pageId="5" pageNumber="348" unit="mm" value="21.0">21 mm</quantity>
.
</paragraph>
</materialsCitation>
<paragraph id="E971BA41DA1B783521B483ADA8EEFD2B" blockId="5.[151,1436,682,798]" pageId="5" pageNumber="348">
<materialsCitation id="59A6B01CDA1B783521B483ADA8EEFD2B" pageId="5" pageNumber="348" specimenCode="IBC-PS668" specimenCount="1" specimenCount-female="1">
<specimenCount id="FFC871C8DA1B783521B483ADAC92FD34" box="[151,172,713,736]" count="1" pageId="5" pageNumber="348" type="female">♀</specimenCount>
: voucher code
<specimenCode id="B968123ADA1B7835206583ACAD80FD34" box="[326,446,712,736]" pageId="5" pageNumber="348">IBC-PS668</specimenCode>
. Forewing length:
<quantity id="2E3617A4DA1B7835235F83ACAEF9FD34" box="[636,711,712,736]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="5" pageNumber="348" unit="mm" value="50.0">50 mm</quantity>
.
<emphasis id="DBBA6653DA1B783523F283ACAFB4FD34" box="[721,906,712,737]" italics="true" pageId="5" pageNumber="348">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA1B783522B483ADAFDCFD34" box="[919,994,713,737]" metricMagnitude="-2" metricUnit="m" metricValue="2.2" pageId="5" pageNumber="348" unit="mm" value="22.0">22 mm</quantity>
. Although the designation of allotype is not regulated but it is allowed by ICZN Rules, this paratype may be considered an allotype of the species.
</materialsCitation>
</paragraph>
<paragraph id="E971BA41DA1B783521B48262AFD3FCC9" blockId="5.[151,1436,682,798]" box="[151,1005,774,798]" pageId="5" pageNumber="348">
<materialsCitation id="59A6B01CDA1B783521B48262AFD3FCC9" box="[151,1005,774,798]" pageId="5" pageNumber="348" specimenCode="IBC-PW976" specimenCount="1" specimenCount-female="1">
<specimenCount id="FFC871C8DA1B783521B48262AC92FCC9" box="[151,172,774,797]" count="1" pageId="5" pageNumber="348" type="female">♀</specimenCount>
: voucher code
<specimenCode id="B968123ADA1B783520658262ADF9FCCA" box="[326,455,774,798]" pageId="5" pageNumber="348">IBC-PW976</specimenCode>
. Forewing length:
<quantity id="2E3617A4DA1B783523A68262AEEEFCC9" box="[645,720,774,798]" metricMagnitude="-2" metricUnit="m" metricValue="4.9" pageId="5" pageNumber="348" unit="mm" value="49.0">49 mm</quantity>
.
<emphasis id="DBBA6653DA1B783523FA8262AFACFCCA" box="[729,914,774,798]" italics="true" pageId="5" pageNumber="348">Length of antenna</emphasis>
:
<quantity id="2E3617A4DA1B783522BC8262AFD7FCC9" box="[927,1001,774,798]" metricMagnitude="-2" metricUnit="m" metricValue="2.2" pageId="5" pageNumber="348" unit="mm" value="22.0">22 mm</quantity>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1B783521B48228AFB9FA56" pageId="5" pageNumber="348" type="description">
<paragraph id="E971BA41DA1B783521B48228AF53FC06" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
Collection and preservation protocols, collection locality and other details same as the
<typeStatus id="367504E3DA1B7835255D8228A8DDFCB2" box="[1150,1251,844,870]" pageId="5" pageNumber="348" type="holotype">holotype</typeStatus>
, except that: (a) IBC- PW982 caterpillar collected on
<date id="9D709C81DA1B783523638214AEFDFC5E" box="[576,707,880,906]" pageId="5" pageNumber="348" value="2012-12-18">2012/12/18</date>
, and mid- and hind-leg on the left side of the adult preserved in ethanol, (b) IBC-PS668 eclosed on
<date id="9D709C81DA1B7835230882F0AE90FC7A" box="[555,686,916,942]" pageId="5" pageNumber="348" value="2013-01-17">2013/01/17</date>
, and both pairs of mid- and hindlegs preserved in ethanol, and (c) IBC-PW976 both mid-legs and left hindleg preserved in ethanol.
</paragraph>
<paragraph id="E971BA41DA1B783521E482B8AD93FB7F" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
The male
<typeStatus id="367504E3DA1B7835201682B9ADABFC23" box="[309,405,989,1015]" pageId="5" pageNumber="348" type="paratype">paratype</typeStatus>
(IBC- PW982) is very similar to the
<typeStatus id="367504E3DA1B7835221382B8AFABFC22" box="[816,917,988,1014]" pageId="5" pageNumber="348" type="holotype">holotype</typeStatus>
, with no significant differences in morphology or markings. Both female
<typeStatus id="367504E3DA1B7835209B8565AE1DFBCF" box="[440,547,1025,1051]" pageId="5" pageNumber="348" type="paratype">paratypes</typeStatus>
have ventral sides similar to the
<typeStatus id="367504E3DA1B783522AD8564AFCEFBCE" box="[910,1008,1024,1050]" pageId="5" pageNumber="348" type="holotype">holotype</typeStatus>
but their discal bands are much darker chocolate-brown. Their dorsal sides are similar to each other, but differ from that of males as follows: (a) Bands from ventral side appear on the dorsal side more prominently, (b) there is a broad, diffused yellow patch from costa till interspace 4, which is separated by a dark brown band from another narrow, yellow submarginal band that spans from interspaces 1b to 5.
</paragraph>
<paragraph id="E971BA41DA1B783521E485D0A9A5FB26" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
Apart from these sexually dimorphic differences, all the
<typeStatus id="367504E3DA1B7835226C85D1AFB4FB1B" box="[847,906,1205,1231]" pageId="5" pageNumber="348">types</typeStatus>
differ to a very small degree from each other, presenting a uniform phenotype in the Western Ghats. Seasonal variation, if any occurs, has not yet been recorded.
</paragraph>
<paragraph id="E971BA41DA1B783521E48598AFB9FA56" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E48598AD01FAC2" bold="true" box="[199,319,1276,1302]" pageId="5" pageNumber="348">Genitalia:</emphasis>
<emphasis id="DBBA6653DA1B783520658598ADBEFAC2" box="[326,384,1276,1302]" italics="true" pageId="5" pageNumber="348">Male</emphasis>
(IBC-PW982,
<figureCitation id="71F5A6C4DA1B7835230A8598AEA8FAC2" box="[553,662,1276,1302]" captionStart="FIGURE 2" captionStartId="6.[151,250,1817,1841]" captionTargetBox="[151,1436,417,1793]" captionTargetId="figure-151@6.[151,1436,417,1793]" captionTargetPageId="6" captionText="FIGURE 2. Male genitalia of Amathusia travancorica sp. nov. (paratype, IBC-PW982). a: lateral view of genital capsule or armature showing the inside of the left valve (right valve removed), b: dorsal view of genital capsule, c: outside of the right valve, d: Phallus (aedeagus), e: dorsal view of the tip of uncus, along with its shape tracing, and f: shape tracings of dorsal view of other Indo-Malayan Amathusia. Photographs: Dipendra Nath Basu and Ujwala Pawar. Genitalia tracings are redrawn from Uémura (1986), except that of A. travancorica sp. nov. (this study)." figureDoi="http://doi.org/10.5281/zenodo.14386303" httpUri="https://zenodo.org/record/14386303/files/figure.png" pageId="5" pageNumber="348">Fig. 2a–e</figureCitation>
): Tips of the uncus relatively slender, long, diverging at a small angle but their two sides are deeply cleft up to the base, and the tip is strongly arched near the joint with tegumen. In lateral view, valve broad, gently sloping, ending in a narrow point, with long hair at the lower margin. Phallus (aedeagus) long, narrow, prominently curving upward for the distal 1/3 length.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA1B783A21E484E8AF7DFE21" lastPageId="10" lastPageNumber="353" pageId="5" pageNumber="348" type="diagnosis">
<paragraph id="E971BA41DA1B783521E484E8AF7EF8DA" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
<emphasis id="DBBA6653DA1B783521E484E8AF7EFA72" bold="true" box="[199,832,1419,1446]" pageId="5" pageNumber="348">
Diagnosis (
<figureCitation id="71F5A6C4DA1B7835206984E8AD8EFA72" box="[330,432,1420,1446]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStartId-0="7.[151,250,1957,1981]" captionStartId-1="9.[151,250,1824,1848]" captionTargetBox-0="[155,1436,188,1922]" captionTargetBox-1="[180,1395,186,1785]" captionTargetId-0="figure-19@7.[151,1436,181,1933]" captionTargetId-1="figure-19@9.[170,1416,181,1801]" captionTargetPageId-0="7" captionTargetPageId-1="9" captionText-0="FIGURE 3. Comparison of Amathusia travancorica sp. nov. with other similar Amathusia (see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar." captionText-1="FIGURE 4. Diagnostic characters of Amathusia travancorica sp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar." figureDoi-0="http://doi.org/10.5281/zenodo.14386305" figureDoi-1="http://doi.org/10.5281/zenodo.14386309" httpUri-0="https://zenodo.org/record/14386305/files/figure.png" httpUri-1="https://zenodo.org/record/14386309/files/figure.png" pageId="5" pageNumber="348">Fig. 3–4</figureCitation>
):
<taxonomicName id="2ECEC1C2DA1B783520E584EFAEE0FA72" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[454,734,1419,1446]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1B783520E584EFAEE0FA72" bold="true" box="[454,734,1419,1446]" italics="true" pageId="5" pageNumber="348">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1B783523C584E8AF7EFA72" box="[742,832,1420,1446]" pageId="5" pageNumber="348" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
has a distinctive phenotype compared to all other SE Asian and Andamanese
<taxonomicName id="2ECEC1C2DA1B7835208084D4AE25FA1E" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[419,539,1456,1482]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA1B7835208084D4AE25FA1E" box="[419,539,1456,1482]" italics="true" pageId="5" pageNumber="348">Amathusia</emphasis>
</taxonomicName>
. A combination of four main characters readily and consistently differentiates
<emphasis id="DBBA6653DA1B783524A084D4ADB4FA3A" bold="true" pageId="5" pageNumber="348">
<taxonomicName id="2ECEC1C2DA1B783524A084D4AD17FA3A" authorityName="Kunte &amp; Basu" authorityYear="2024" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1B783524A084D4AD17FA3A" bold="true" italics="true" pageId="5" pageNumber="348">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1B7835201384B0ADB4FA3A" box="[304,394,1492,1518]" pageId="5" pageNumber="348" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
from related taxa (
<figureCitation id="71F5A6C4DA1B7835237C84B0AE9DFA3A" box="[607,675,1492,1518]" captionStart="FIGURE 4" captionStartId="9.[151,250,1824,1848]" captionTargetBox="[180,1395,186,1785]" captionTargetId="figure-19@9.[170,1416,181,1801]" captionTargetPageId="9" captionText="FIGURE 4. Diagnostic characters of Amathusia travancorica sp. nov., marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar." figureDoi="http://doi.org/10.5281/zenodo.14386309" httpUri="https://zenodo.org/record/14386309/files/figure.png" pageId="5" pageNumber="348">Fig. 4</figureCitation>
, in which numbered pointers correspond to the following numbered characters): (1) FW termen straight in both sexes, not concave (prominently concave in
<taxonomicName id="2ECEC1C2DA1B783525A1849CA8C0F9C6" authorityName="Fruhstorfer" authorityYear="1904" box="[1154,1278,1528,1554]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA1B783525A1849CA8C0F9C6" box="[1154,1278,1528,1554]" italics="true" pageId="5" pageNumber="348">A. friderici</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA1B7835242E849CA9A2F9C6" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[1293,1436,1528,1554]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA1B7835242E849CA9A2F9C6" box="[1293,1436,1528,1554]" italics="true" pageId="5" pageNumber="348">A. phidippus</emphasis>
</taxonomicName>
and all other SE Asian species, especially in
<taxonomicName id="2ECEC1C2DA1B783523898779AF24F9E2" box="[682,794,1565,1590]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="masina">
<emphasis id="DBBA6653DA1B783523898779AF24F9E2" box="[682,794,1565,1590]" italics="true" pageId="5" pageNumber="348">P. masina</emphasis>
</taxonomicName>
); (2) HW termen conspicuously more convex, and not prominently produced leading up to the tornal lobe (HW termen straighter and tornus prominently elongated and produced in
<taxonomicName id="2ECEC1C2DA1B783520078700AD9EF9AA" authorityName="Fruhstorfer" authorityYear="1904" box="[292,416,1636,1662]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA1B783520078700AD9EF9AA" box="[292,416,1636,1662]" italics="true" pageId="5" pageNumber="348">A. friderici</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA1B7835208D8700AE05F9AA" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[430,571,1636,1662]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA1B7835208D8700AE05F9AA" box="[430,571,1636,1662]" italics="true" pageId="5" pageNumber="348">A. phidippus</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA1B783523698700AE85F9AA" box="[586,699,1636,1662]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="masina">
<emphasis id="DBBA6653DA1B783523698700AE85F9AA" box="[586,699,1636,1662]" italics="true" pageId="5" pageNumber="348">A. masina</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA1B783523D18700AF42F9AA" authorityName="Honrath" authorityYear="1888" box="[754,892,1636,1662]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="perakana">
<emphasis id="DBBA6653DA1B783523D18700AF42F9AA" box="[754,892,1636,1662]" italics="true" pageId="5" pageNumber="348">A. perakana</emphasis>
</taxonomicName>
); (3) In male, UPF is uniform pale rusty-brown, without any yellow markings (very prominent yellow subapical band almost joining the submarginal yellow band in
<taxonomicName id="2ECEC1C2DA1B7835219587C8AD0DF912" authorityName="Fruhstorfer" authorityYear="1904" box="[182,307,1708,1734]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA1B7835219587C8AD0DF912" box="[182,307,1708,1734]" italics="true" pageId="5" pageNumber="348">A. friderici</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA1B7835206187C8ADF3F912" authorityName="Fruhstorfer" authorityYear="1904" box="[322,461,1708,1734]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="binghami">
<emphasis id="DBBA6653DA1B7835206187C8ADF3F912" box="[322,461,1708,1734]" italics="true" pageId="5" pageNumber="348">A. binghami</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA1B7835232487C8AE8EF912" authorityName="Honrath" authorityYear="1888" box="[519,688,1708,1734]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="schoenbergi">
<emphasis id="DBBA6653DA1B7835232487C8AE8EF912" box="[519,688,1708,1734]" italics="true" pageId="5" pageNumber="348">A. schoenbergi</emphasis>
</taxonomicName>
; pale submarginal band showing through more prominently in
<taxonomicName id="2ECEC1C2DA1B783524A787C8AD3BF93E" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA1B783524A787C8AD3BF93E" italics="true" pageId="5" pageNumber="348">A. phidippus</emphasis>
</taxonomicName>
), and (4) In female, UPF yellow markings are more faint and diffused (yellow markings very prominent and more clearly delineated in
<taxonomicName id="2ECEC1C2DA1B783520D28790AE52F8DA" authorityName="Fruhstorfer" authorityYear="1904" box="[497,620,1780,1806]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA1B783520D28790AE52F8DA" box="[497,620,1780,1806]" italics="true" pageId="5" pageNumber="348">A. friderici</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA1B783523818790AF0EF8DA" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[674,816,1780,1806]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA1B783523818790AF0EF8DA" box="[674,816,1780,1806]" italics="true" pageId="5" pageNumber="348">A. phidippus</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="E971BA41DA1B783521E4867CA87BF8AE" blockId="5.[151,1437,844,2023]" pageId="5" pageNumber="348">
Additional diagnostic characters of
<emphasis id="DBBA6653DA1B7835237D867CAF48F8E6" bold="true" box="[606,886,1816,1842]" pageId="5" pageNumber="348">
<taxonomicName id="2ECEC1C2DA1B7835237D867CAF2CF8E6" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[606,786,1816,1842]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1B7835237D867CAF2CF8E6" bold="true" box="[606,786,1816,1842]" italics="true" pageId="5" pageNumber="348">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1B78352238867CAF48F8E6" box="[795,886,1816,1842]" pageId="5" pageNumber="348" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
are: (a) UNH has two large eyespots (two large spots and a smaller spot in between the two large spots in
<taxonomicName id="2ECEC1C2DA1B783522058658AFD8F882" authorityName="Fruhstorfer" authorityYear="1899" box="[806,998,1852,1878]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="andamanensis">
<emphasis id="DBBA6653DA1B783522058658AFD8F882" box="[806,998,1852,1878]" italics="true" pageId="5" pageNumber="348">A. andamanensis</emphasis>
</taxonomicName>
), (b) UNF outer edge of the large dark discal band smooth (conspicuously jagged in
<taxonomicName id="2ECEC1C2DA1B783523B68604AF69F8AE" authorityName="Honrath" authorityYear="1888" box="[661,855,1888,1914]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="ochraceofusca">
<emphasis id="DBBA6653DA1B783523B68604AF69F8AE" box="[661,855,1888,1914]" italics="true" pageId="5" pageNumber="348">A. ochraceofusca</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA1B783522AD8604A80BF8AE" authorityName="Honrath" authorityYear="1888" box="[910,1077,1888,1914]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="schoenbergi">
<emphasis id="DBBA6653DA1B783522AD8604A80BF8AE" box="[910,1077,1888,1914]" italics="true" pageId="5" pageNumber="348">A. schoenbergi</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="E971BA41DA1B783621E486E0AFA8FE5D" blockId="5.[151,1437,844,2023]" lastBlockId="6.[151,1437,151,393]" lastPageId="6" lastPageNumber="349" pageId="5" pageNumber="348">
The male genitalia of
<emphasis id="DBBA6653DA1B783520E186E0AEE7F84A" bold="true" box="[450,729,1924,1950]" pageId="5" pageNumber="348">
<taxonomicName id="2ECEC1C2DA1B783520E186E0AE4BF84A" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[450,629,1924,1950]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA1B783520E186E0AE4BF84A" bold="true" box="[450,629,1924,1950]" italics="true" pageId="5" pageNumber="348">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA1B7835235D86E0AEE7F84A" box="[638,729,1924,1950]" pageId="5" pageNumber="348" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
may be distinguished from all other Indo-Malayan
<taxonomicName id="2ECEC1C2DA1B7835240786E0A9A2F84A" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1316,1436,1924,1950]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA1B7835240786E0A9A2F84A" box="[1316,1436,1924,1950]" italics="true" pageId="5" pageNumber="348">Amathusia</emphasis>
</taxonomicName>
by their slender, long tips of the uncus, which diverge from each other at a small angle but they are deeply cleft up to the base (
<figureCitation id="71F5A6C4DA1B7835203E86A8AD50F832" box="[285,366,1996,2022]" captionStart="FIGURE 2" captionStartId="6.[151,250,1817,1841]" captionTargetBox="[151,1436,417,1793]" captionTargetId="figure-151@6.[151,1436,417,1793]" captionTargetPageId="6" captionText="FIGURE 2. Male genitalia of Amathusia travancorica sp. nov. (paratype, IBC-PW982). a: lateral view of genital capsule or armature showing the inside of the left valve (right valve removed), b: dorsal view of genital capsule, c: outside of the right valve, d: Phallus (aedeagus), e: dorsal view of the tip of uncus, along with its shape tracing, and f: shape tracings of dorsal view of other Indo-Malayan Amathusia. Photographs: Dipendra Nath Basu and Ujwala Pawar. Genitalia tracings are redrawn from Uémura (1986), except that of A. travancorica sp. nov. (this study)." figureDoi="http://doi.org/10.5281/zenodo.14386303" httpUri="https://zenodo.org/record/14386303/files/figure.png" pageId="5" pageNumber="348">Fig. 2e</figureCitation>
). In contrast, uncus tips of other
<taxonomicName id="2ECEC1C2DA1B783523F886A8AF6DF832" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[731,851,1996,2022]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="5" pageNumber="348" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA1B783523F886A8AF6DF832" box="[731,851,1996,2022]" italics="true" pageId="5" pageNumber="348">Amathusia</emphasis>
</taxonomicName>
diverge from each other at a much greater angle and therefore their tips are widely separated; those in
<taxonomicName id="2ECEC1C2DA187836239E81F3AF63FF65" authorityName="Fruhstorfer" authorityYear="1899" box="[701,861,151,177]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="andamanensis">
<emphasis id="DBBA6653DA187836239E81F3AF63FF65" box="[701,861,151,177]" italics="true" pageId="6" pageNumber="349">andamanensis</emphasis>
</taxonomicName>
are relatively broader and shorter, those in
<taxonomicName id="2ECEC1C2DA187836246381F3A9A2FF65" authorityName="Fruhstorfer" authorityYear="1904" box="[1344,1436,151,177]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA187836246381F3A9A2FF65" box="[1344,1436,151,177]" italics="true" pageId="6" pageNumber="349">friderici</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA18783621E581DFAD29FF00" box="[198,279,187,212]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="masina">
<emphasis id="DBBA6653DA18783621E581DFAD29FF00" box="[198,279,187,212]" italics="true" pageId="6" pageNumber="349">masina</emphasis>
</taxonomicName>
are bent outward at the tip and more widely separated at the base, and those in
<taxonomicName id="2ECEC1C2DA18783625AA81DFA8C9FF01" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[1161,1271,187,213]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA18783625AA81DFA8C9FF01" box="[1161,1271,187,213]" italics="true" pageId="6" pageNumber="349">phidippus</emphasis>
</taxonomicName>
are not deeply cleft up to the base (
<figureCitation id="71F5A6C4DA187836205581BBADFCFF2D" box="[374,450,223,249]" captionStart="FIGURE 2" captionStartId="6.[151,250,1817,1841]" captionTargetBox="[151,1436,417,1793]" captionTargetId="figure-151@6.[151,1436,417,1793]" captionTargetPageId="6" captionText="FIGURE 2. Male genitalia of Amathusia travancorica sp. nov. (paratype, IBC-PW982). a: lateral view of genital capsule or armature showing the inside of the left valve (right valve removed), b: dorsal view of genital capsule, c: outside of the right valve, d: Phallus (aedeagus), e: dorsal view of the tip of uncus, along with its shape tracing, and f: shape tracings of dorsal view of other Indo-Malayan Amathusia. Photographs: Dipendra Nath Basu and Ujwala Pawar. Genitalia tracings are redrawn from Uémura (1986), except that of A. travancorica sp. nov. (this study)." figureDoi="http://doi.org/10.5281/zenodo.14386303" httpUri="https://zenodo.org/record/14386303/files/figure.png" pageId="6" pageNumber="349">Fig. 2f</figureCitation>
) (
<bibRefCitation id="8D5FC7B0DA18783620FB81BBAEAFFF2D" author="Aoki, T. &amp; Yamaguchi, S. &amp; Uemura, Y." box="[472,657,223,249]" pageId="6" pageNumber="349" refId="ref12426" refString="Aoki, T., Yamaguchi, S. &amp; Uemura, Y. (1982) Butterflies of the South East Asian Islands. Vol. 3. Satyridae, Amathusiidae and Libytheidae. Plapac Co. Ltd., Tokyo, 530 pp." type="book" year="1982">
Aoki
<emphasis id="DBBA6653DA187836233B81BBAE6EFF2D" box="[536,592,223,249]" italics="true" pageId="6" pageNumber="349">et al.</emphasis>
1982
</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA18783623BF81BBAF56FF2D" author="Corbet, A. S. &amp; Pendlebury, H. M. &amp; Eliot, J. N." box="[668,872,223,249]" pageId="6" pageNumber="349" refId="ref12929" refString="Corbet, A. S., Pendlebury, H. M. &amp; Eliot, J. N. (2020) The Butterflies of the Malay Peninsula (5 th edition revised by George and Nancy van der Poorten). Lepodon Books, Toronto, 630 pp." type="book" year="2020">
Corbet
<emphasis id="DBBA6653DA18783623CD81BBAF18FF2D" box="[750,806,223,249]" italics="true" pageId="6" pageNumber="349">et al.</emphasis>
2020
</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA187836225181BBAFD4FF2D" author="Eliot, J. N." box="[882,1002,223,249]" pageId="6" pageNumber="349" pagination="155 - 161" refId="ref13125" refString="Eliot, J. N. (1964) A note on the phidippus L. group of the genus Amathusia (Lepidoptera: Amathusiidae), with special reference to the Malayan foms. The Entomologist, 97 (1), 155 - 161." type="journal article" year="1964">Eliot 1964</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA18783622D681BBA8AFFF2D" author="Uemura, Y." box="[1013,1169,223,249]" pageId="6" pageNumber="349" pagination="2 - 8" refId="ref15108" refString="Uemura, Y. (1986) Identification guide to the species of the genus Amathusia (Lepidoptera: Satyridae; Amathusiinae). Yadoriga, 125, 2 - 8." type="journal article" year="1986">Uémura 1986</bibRefCitation>
). Additionally, the long, narrow phallus in
<emphasis id="DBBA6653DA18783620418067AE48FEC9" bold="true" box="[354,630,259,285]" pageId="6" pageNumber="349">
<taxonomicName id="2ECEC1C2DA18783620418067AE2BFEC9" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[354,533,259,285]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA18783620418067AE2BFEC9" bold="true" box="[354,533,259,285]" italics="true" pageId="6" pageNumber="349">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA187836233F8067AE48FEC9" box="[540,630,259,285]" pageId="6" pageNumber="349" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is prominently curved upward for the distal 1/3 length, whereas phallus in
<taxonomicName id="2ECEC1C2DA18783621958043AD69FE95" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[182,343,295,321]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA18783621958043AD69FE95" box="[182,343,295,321]" italics="true" pageId="6" pageNumber="349">A. phidippus</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA18783620B28043AE27FE95" authorityName="Honrath" authorityYear="1888" box="[401,537,295,321]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="schoenbergi">
<emphasis id="DBBA6653DA18783620B28043AE27FE95" box="[401,537,295,321]" italics="true" pageId="6" pageNumber="349">schoenbergi</emphasis>
</taxonomicName>
is nearly straight and curved upward gently only for the distal 1/4 length, that in
<taxonomicName id="2ECEC1C2DA1878362196802FAD21FEB1" authorityName="Fruhstorfer" authorityYear="1904" box="[181,287,331,357]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="binghami">
<emphasis id="DBBA6653DA1878362196802FAD21FEB1" box="[181,287,331,357]" italics="true" pageId="6" pageNumber="349">binghami</emphasis>
</taxonomicName>
,
<taxonomicName id="2ECEC1C2DA187836200D802FADEFFEB1" authorityName="Honrath" authorityYear="1888" box="[302,465,331,357]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="ochraceofusca">
<emphasis id="DBBA6653DA187836200D802FADEFFEB1" box="[302,465,331,357]" italics="true" pageId="6" pageNumber="349">ochraceofusca</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA1878362329802FAE4AFEB1" authorityName="Honrath" authorityYear="1888" box="[522,628,331,357]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="perakana">
<emphasis id="DBBA6653DA1878362329802FAE4AFEB1" box="[522,628,331,357]" italics="true" pageId="6" pageNumber="349">perakana</emphasis>
</taxonomicName>
prominently bent upward at half length (it is also much more robust in
<taxonomicName id="2ECEC1C2DA18783621B4800BAD3FFE5D" authorityName="Honrath" authorityYear="1888" box="[151,257,367,393]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="perakana">
<emphasis id="DBBA6653DA18783621B4800BAD3FFE5D" box="[151,257,367,393]" italics="true" pageId="6" pageNumber="349">perakana</emphasis>
</taxonomicName>
), and the tip is much broader in
<taxonomicName id="2ECEC1C2DA187836234D800BAFAFFE5D" authority="(Aoki et al. 1982)" baseAuthorityName="Aoki" baseAuthorityYear="1982" box="[622,913,367,393]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="masina">
<emphasis id="DBBA6653DA187836234D800BAE81FE5C" box="[622,703,367,392]" italics="true" pageId="6" pageNumber="349">masina</emphasis>
(
<bibRefCitation id="8D5FC7B0DA18783623EE800BAFB7FE5D" author="Aoki, T. &amp; Yamaguchi, S. &amp; Uemura, Y." box="[717,905,367,393]" pageId="6" pageNumber="349" refId="ref12426" refString="Aoki, T., Yamaguchi, S. &amp; Uemura, Y. (1982) Butterflies of the South East Asian Islands. Vol. 3. Satyridae, Amathusiidae and Libytheidae. Plapac Co. Ltd., Tokyo, 530 pp." type="book" year="1982">
Aoki
<emphasis id="DBBA6653DA187836222D800BAF78FE5D" box="[782,838,367,393]" italics="true" pageId="6" pageNumber="349">et al.</emphasis>
1982
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<caption id="BDB1EAC9DA18783621B4867DAF77F815" ID-DOI="http://doi.org/10.5281/zenodo.14386303" ID-Zenodo-Dep="14386303" httpUri="https://zenodo.org/record/14386303/files/figure.png" pageId="6" pageNumber="349" startId="6.[151,250,1817,1841]" targetBox="[151,1436,417,1793]" targetPageId="6" targetType="figure">
<paragraph id="E971BA41DA18783621B4867DAF77F815" blockId="6.[151,1436,1817,1985]" pageId="6" pageNumber="349">
<emphasis id="DBBA6653DA18783621B4867DAD2AF8E5" bold="true" box="[151,276,1817,1841]" pageId="6" pageNumber="349">FIGURE 2.</emphasis>
Male genitalia of
<emphasis id="DBBA6653DA18783620F2867DAF19F8E5" bold="true" box="[465,807,1817,1841]" pageId="6" pageNumber="349">
<taxonomicName id="2ECEC1C2DA18783620F2867DAEF0F8E5" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[465,718,1817,1841]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA18783620F2867DAEF0F8E5" bold="true" box="[465,718,1817,1841]" italics="true" pageId="6" pageNumber="349">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA18783623F5867DAF19F8E5" box="[726,807,1817,1841]" pageId="6" pageNumber="349" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(paratype, IBC-PW982).
<emphasis id="DBBA6653DA1878362513867DA878F8E5" bold="true" box="[1072,1094,1817,1841]" pageId="6" pageNumber="349">a:</emphasis>
lateral view of genital capsule or armature showing the inside of the left valve (right valve removed),
<emphasis id="DBBA6653DA18783622788659AF4CF881" bold="true" box="[859,882,1853,1877]" pageId="6" pageNumber="349">b:</emphasis>
dorsal view of genital capsule,
<emphasis id="DBBA6653DA187836259F8659A8EEF881" bold="true" box="[1212,1232,1853,1877]" pageId="6" pageNumber="349">c:</emphasis>
outside of the right valve,
<emphasis id="DBBA6653DA18783621F98605ACCFF8AD" bold="true" box="[218,241,1889,1913]" pageId="6" pageNumber="349">d:</emphasis>
Phallus (aedeagus),
<emphasis id="DBBA6653DA18783620E38605ADEAF8AD" bold="true" box="[448,468,1889,1913]" pageId="6" pageNumber="349">e:</emphasis>
dorsal view of the tip of uncus, along with its shape tracing, and
<emphasis id="DBBA6653DA18783625428605A84FF8AD" bold="true" box="[1121,1137,1889,1913]" pageId="6" pageNumber="349">f:</emphasis>
shape tracings of dorsal view of other Indo-Malayan
<taxonomicName id="2ECEC1C2DA18783620A586E1ADCCF849" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[390,498,1925,1949]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA18783620A586E1ADCCF849" box="[390,498,1925,1949]" italics="true" pageId="6" pageNumber="349">Amathusia</emphasis>
</taxonomicName>
. Photographs: Dipendra Nath Basu and Ujwala Pawar. Genitalia tracings are redrawn from
<bibRefCitation id="8D5FC7B0DA18783621B486CDAD0BF815" author="Uemura, Y." box="[151,309,1961,1985]" pageId="6" pageNumber="349" pagination="2 - 8" refId="ref15108" refString="Uemura, Y. (1986) Identification guide to the species of the genus Amathusia (Lepidoptera: Satyridae; Amathusiinae). Yadoriga, 125, 2 - 8." type="journal article" year="1986">Uémura (1986)</bibRefCitation>
, except that of
<emphasis id="DBBA6653DA18783620F386CDAEF6F815" bold="true" box="[464,712,1961,1985]" pageId="6" pageNumber="349">
<taxonomicName id="2ECEC1C2DA18783620F386CDAE4FF815" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[464,625,1961,1985]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="6" pageNumber="349" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA18783620F386CDAE4FF815" bold="true" box="[464,625,1961,1985]" italics="true" pageId="6" pageNumber="349">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA187836235B86CDAEF6F815" box="[632,712,1961,1985]" pageId="6" pageNumber="349" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(this study).
</paragraph>
</caption>
<caption id="BDB1EAC9DA19783721B486C1A93AF835" ID-DOI="http://doi.org/10.5281/zenodo.14386305" ID-Zenodo-Dep="14386305" httpUri="https://zenodo.org/record/14386305/files/figure.png" pageId="7" pageNumber="350" startId="7.[151,250,1957,1981]" targetBox="[155,1436,188,1922]" targetPageId="7" targetType="figure">
<paragraph id="E971BA41DA19783721B486C1A93AF835" blockId="7.[151,1437,1957,2017]" pageId="7" pageNumber="350">
<emphasis id="DBBA6653DA19783721B486C1AD2BF869" bold="true" box="[151,277,1957,1981]" pageId="7" pageNumber="350">FIGURE 3.</emphasis>
Comparison of
<emphasis id="DBBA6653DA197837209C86C1AF27F869" bold="true" box="[447,793,1957,1981]" pageId="7" pageNumber="350">
<taxonomicName id="2ECEC1C2DA197837209C86C1AE80F869" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[447,702,1957,1981]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="350" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA197837209C86C1AE80F869" bold="true" box="[447,702,1957,1981]" italics="true" pageId="7" pageNumber="350">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA19783723E586C1AF27F869" box="[710,793,1957,1981]" pageId="7" pageNumber="350" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
with other similar
<taxonomicName id="2ECEC1C2DA19783722C086C1A871F869" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[995,1103,1957,1981]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="7" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA19783722C086C1A871F869" box="[995,1103,1957,1981]" italics="true" pageId="7" pageNumber="350">Amathusia</emphasis>
</taxonomicName>
(see Materials and Methods for specimen details). A millimetre scale is at the bottom. Photographs: Krushnamegh Kunte and Gaurav Agavekar.
</paragraph>
</caption>
<caption id="BDB1EAC9DA16783821B486F6ADAAF87E" box="[151,404,1938,1962]" isContinuationCaption="true" pageId="8" pageNumber="351" startId="8.[151,250,1938,1962]" targetBox="[151,1430,191,1903]" targetPageId="8" targetType="figure">
<paragraph id="E971BA41DA16783821B486F6ADAAF87E" blockId="8.[151,404,1938,1962]" box="[151,404,1938,1962]" pageId="8" pageNumber="351">
<emphasis id="DBBA6653DA16783821B486F6AD2DF87E" bold="true" box="[151,275,1938,1962]" pageId="8" pageNumber="351">FIGURE 3.</emphasis>
(Continued)
</paragraph>
</caption>
<caption id="BDB1EAC9DA17783921B48644AD71F839" ID-DOI="http://doi.org/10.5281/zenodo.14386309" ID-Zenodo-Dep="14386309" httpUri="https://zenodo.org/record/14386309/files/figure.png" pageId="9" pageNumber="352" startId="9.[151,250,1824,1848]" targetBox="[180,1395,186,1785]" targetPageId="9" targetType="figure">
<paragraph id="E971BA41DA17783921B48644AD71F839" blockId="9.[151,1437,1824,2029]" pageId="9" pageNumber="352">
<emphasis id="DBBA6653DA17783921B48644AD28F8EC" bold="true" box="[151,278,1824,1848]" pageId="9" pageNumber="352">FIGURE 4.</emphasis>
Diagnostic characters of
<emphasis id="DBBA6653DA17783923058644AFBDF8ED" bold="true" box="[550,899,1824,1849]" pageId="9" pageNumber="352">
<taxonomicName id="2ECEC1C2DA17783923058644AF1BF8ED" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[550,805,1824,1849]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="9" pageNumber="352" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA17783923058644AF1BF8ED" bold="true" box="[550,805,1824,1849]" italics="true" pageId="9" pageNumber="352">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA177839220C8645AFBDF8ED" box="[815,899,1825,1849]" pageId="9" pageNumber="352" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, marked on digitally enhanced images from which wing-wear has been removed to illustrate the distinctive phenotype of the species. The new species is compared with two species whose names have been applied to the Western Ghats populations in the past. Numbers point to distinctive characters: 1: forewing termen, 2: hindwing termen and tornus area, 3: yellow markings on forewing of male, 4: yellow markings on forewing of female (further details in the text). Images are not to scale. Photographs: Krushnamegh Kunte, Dipendra Nath Basu and Gaurav Agavekar.
</paragraph>
</caption>
<paragraph id="E971BA41DA14783A21E481F3AF7DFE21" blockId="10.[151,1437,151,501]" pageId="10" pageNumber="353">
Several authors have earlier described the placement and colour of hair-pencils at v1b, interspace 1c, and between v7 and v8 among
<taxonomicName id="2ECEC1C2DA14783A20EA81DFAEA2FF01" box="[457,668,187,213]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A20EA81DFAE7FFF01" box="[457,577,187,213]" italics="true" pageId="10" pageNumber="353">Amathusia</emphasis>
species
</taxonomicName>
, in addition to the hair pencils on the side of abdomen (
<bibRefCitation id="8D5FC7B0DA14783A243781DFA9AEFF01" author="Eliot, J. N." box="[1300,1424,187,213]" pageId="10" pageNumber="353" pagination="155 - 161" refId="ref13125" refString="Eliot, J. N. (1964) A note on the phidippus L. group of the genus Amathusia (Lepidoptera: Amathusiidae), with special reference to the Malayan foms. The Entomologist, 97 (1), 155 - 161." type="journal article" year="1964">Eliot 1964</bibRefCitation>
). We have observed that the size and colour of these hair-pencils are variable among individuals within each species. Younger individuals have larger hair-pencils, and older individuals have much smaller hair-pencils, sometimes having lost all the long scales. The colour is usually variable, ranging from pale brown to black. All these variations have been observed among individuals of
<emphasis id="DBBA6653DA14783A2344802FAF48FEB1" bold="true" box="[615,886,331,357]" pageId="10" pageNumber="353">
<taxonomicName id="2ECEC1C2DA14783A2344802FAF26FEB1" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[615,792,331,357]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA14783A2344802FAF26FEB1" bold="true" box="[615,792,331,357]" italics="true" pageId="10" pageNumber="353">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA14783A223D802FAF48FEB1" box="[798,886,331,357]" pageId="10" pageNumber="353" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, including the
<typeStatus id="367504E3DA14783A2539802FA874FEB1" box="[1050,1098,331,357]" pageId="10" pageNumber="353">type</typeStatus>
specimens described above. If there are consistent species-level differences among
<taxonomicName id="2ECEC1C2DA14783A23FD800BAF8EFE5D" box="[734,944,367,393]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A23FD800BAF68FE5D" box="[734,854,367,393]" italics="true" pageId="10" pageNumber="353">Amathusia</emphasis>
species
</taxonomicName>
, then they need to be characterized carefully and reliably before they may be used as diagnostic, especially between
<emphasis id="DBBA6653DA14783A229080F7A8F8FE79" bold="true" box="[947,1222,403,429]" pageId="10" pageNumber="353">
<taxonomicName id="2ECEC1C2DA14783A229080F7A858FE79" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[947,1126,403,429]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA14783A229080F7A858FE79" bold="true" box="[947,1126,403,429]" italics="true" pageId="10" pageNumber="353">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA14783A254E80F7A8F8FE79" box="[1133,1222,403,429]" pageId="10" pageNumber="353" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
,
<taxonomicName id="2ECEC1C2DA14783A25F780F7A970FE79" authorityName="Fruhstorfer" authorityYear="1904" box="[1236,1358,403,429]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="friderici">
<emphasis id="DBBA6653DA14783A25F780F7A970FE79" box="[1236,1358,403,429]" italics="true" pageId="10" pageNumber="353">A. friderici</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA14783A24A780F7AD3BFE05" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA14783A24A780F7AD3BFE05" italics="true" pageId="10" pageNumber="353">A. phidippus</emphasis>
</taxonomicName>
. In museum specimens these hair-pencils are often damaged during pinning, or those inside wing folds are difficult to evert, as also observed earlier by
<bibRefCitation id="8D5FC7B0DA14783A239780BFAF01FE21" author="Eliot, J. N." box="[692,831,475,501]" pageId="10" pageNumber="353" pagination="155 - 161" refId="ref13125" refString="Eliot, J. N. (1964) A note on the phidippus L. group of the genus Amathusia (Lepidoptera: Amathusiidae), with special reference to the Malayan foms. The Entomologist, 97 (1), 155 - 161." type="journal article" year="1964">Eliot (1964)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA14783A21B48323AD20FDB5" box="[151,286,583,609]" pageId="10" pageNumber="353" type="etymology">
<paragraph id="E971BA41DA14783A21B48323AD20FDB5" blockId="10.[151,286,583,609]" box="[151,286,583,609]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21B48323AD20FDB5" bold="true" box="[151,286,583,609]" pageId="10" pageNumber="353">Etymology</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA14783A21B483EAAF20FCC1" pageId="10" pageNumber="353" type="vernacular_names">
<paragraph id="E971BA41DA14783A21B483EAADE2FD19" blockId="10.[151,1437,654,1293]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21B483EAAD66FD7D" bold="true" box="[151,344,654,681]" pageId="10" pageNumber="353">Scientific name:</emphasis>
We name
<emphasis id="DBBA6653DA14783A20EC83EAAF76FD7D" bold="true" box="[463,840,654,681]" pageId="10" pageNumber="353">
<taxonomicName id="2ECEC1C2DA14783A20EC83EAAED9FD7D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[463,743,654,681]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA14783A20EC83EAAED9FD7D" bold="true" box="[463,743,654,681]" italics="true" pageId="10" pageNumber="353">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA14783A23CD83EBAF76FD7D" box="[750,840,655,681]" pageId="10" pageNumber="353" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
after its endemic distributional range in the southern Travancore region of
<collectingRegion id="2B0A74A3DA14783A20A983D7ADE6FD19" box="[394,472,691,717]" country="India" name="Kerala" pageId="10" pageNumber="353">Kerala</collectingRegion>
.
</paragraph>
<paragraph id="E971BA41DA14783A21E483B2AF20FCC1" blockId="10.[151,1437,654,1293]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21E483B2AD48FD25" bold="true" box="[199,374,726,753]" pageId="10" pageNumber="353">English name:</emphasis>
We propose the English name ‘Travancore Palmking’ for
<emphasis id="DBBA6653DA14783A253F83B2A9A2FD25" bold="true" box="[1052,1436,726,753]" pageId="10" pageNumber="353">
<taxonomicName id="2ECEC1C2DA14783A253F83B2A908FD25" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[1052,1334,726,753]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA14783A253F83B2A908FD25" bold="true" box="[1052,1334,726,753]" italics="true" pageId="10" pageNumber="353">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA14783A246383B3A9A2FD25" box="[1344,1436,727,753]" pageId="10" pageNumber="353" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
based on its endemism to the southern Travancore region.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA14783A21E4827AAE18FAAD" pageId="10" pageNumber="353" type="discussion">
<paragraph id="E971BA41DA14783A21E4827AAF2BFAD9" blockId="10.[151,1437,654,1293]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21E4827AAD8DFCED" bold="true" box="[199,435,798,825]" pageId="10" pageNumber="353">Historical Records:</emphasis>
<bibRefCitation id="8D5FC7B0DA14783A209D827BAF7FFCED" author="Marshall, G. F. L. &amp; de Niceville, L." box="[446,833,799,825]" pageId="10" pageNumber="353" refId="ref14246" refString="Marshall, G. F. L. &amp; de Niceville, L. (1882) The Butterflies of India, Burmah and Ceylon. Volume I. Nymphalidae. Danainae, Satyrinae, Elymniinae, Morphinae, Acraeinae. The Calcutta Central Press Co., Kolkata, 327 pp." type="book" year="1882">Marshall and de Nicéville (1882)</bibRefCitation>
,
<bibRefCitation id="8D5FC7B0DA14783A2272827BAFCDFCED" author="Moore, F." box="[849,1011,799,825]" pageId="10" pageNumber="353" refId="ref14413" refString="Moore, F. (1896) Lepidoptera Indica. Vol. II. Rhopalocera. Family Nymphalidae. Sub-families Satyrinae (continued), Elymniinae, Amathusiinae, Nymphalinae (group Charaxina) [1893 - 1896]. Reeve &amp; Co, London, 274 pp." type="book" year="1896">Moore (1896)</bibRefCitation>
,
<bibRefCitation id="8D5FC7B0DA14783A2522827BA89EFCED" author="Talbot, G." box="[1025,1184,799,825]" pageId="10" pageNumber="353" refId="ref15075" refString="Talbot, G. (1947) The Fauna of British India, including Ceylon and Burma: Butterflies, Vol. 2. Taylor &amp; Francis, London, 506 pp." type="book" year="1947">Talbot (1947)</bibRefCitation>
, and
<bibRefCitation id="8D5FC7B0DA14783A25C1827BA9A2FCED" author="d'Abrera, B." box="[1250,1436,798,825]" pageId="10" pageNumber="353" refId="ref12980" refString="d'Abrera, B. (1985) Butterflies of the Oriental Region. Part II: Nymphalidae, Satyridae &amp; Amathusidae. Hill House, Melbourne, 534 pp." type="book" year="1985">d’Abrera (1985)</bibRefCitation>
did not mention the Western Ghats populations of
<taxonomicName id="2ECEC1C2DA14783A23C68227AF63FC89" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[741,861,835,861]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA14783A23C68227AF63FC89" box="[741,861,835,861]" italics="true" pageId="10" pageNumber="353">Amathusia</emphasis>
</taxonomicName>
.
<bibRefCitation id="8D5FC7B0DA14783A224D8227A839FC89" author="Evans, W. H." box="[878,1031,835,861]" pageId="10" pageNumber="353" refId="ref13172" refString="Evans, W. H. (1932) The Identification of Indian Butterflies (2 nd edn.). Bombay Natural History Society, Mumbai, 454 pp." type="book" year="1932">Evans (1932)</bibRefCitation>
listed
<taxonomicName id="2ECEC1C2DA14783A257B8227A971FC89" authorityName="Fruhstorfer" authorityYear="1904" box="[1112,1359,835,861]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="friderici">
<emphasis id="DBBA6653DA14783A257B8227A971FC89" box="[1112,1359,835,861]" italics="true" pageId="10" pageNumber="353">A. phidippus friderici</emphasis>
</taxonomicName>
in “S.
<collectingCountry id="91D9FAD1DA14783A21B48203ACEBFC55" box="[151,213,871,897]" name="India" pageId="10" pageNumber="353">India</collectingCountry>
”, and
<bibRefCitation id="8D5FC7B0DA14783A203F8203AE39FC55" author="Wynter-Blyth, M. A." box="[284,519,870,897]" pageId="10" pageNumber="353" refId="ref15140" refString="Wynter-Blyth, M. A. (1957) Butterflies of the Indian Region. Bombay Natural History Society, Mumbai, 523 pp." type="book" year="1957">Wynter-Blyth (1957)</bibRefCitation>
as
<taxonomicName id="2ECEC1C2DA14783A230D8203AE82FC55" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[558,700,871,897]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA14783A230D8203AE82FC55" box="[558,700,871,897]" italics="true" pageId="10" pageNumber="353">A. phidippus</emphasis>
</taxonomicName>
in “Travancore”, but neither mentioned specific locations. These two mentions appear to be based on
<specimenCount id="FFC871C8DA14783A231482EFAED2FC71" box="[567,748,907,933]" count="3" pageId="10" pageNumber="353" type="generic">three specimens</specimenCount>
collected from near Thiruvananthapuram, presumably in the 1920s, but we have been unable to locate these specimens (they may have been destroyed in an unknown museum in the Thiruvananthapuram area). NHMUK has a single female from Kottayam,
<collectingRegion id="2B0A74A3DA14783A250382B7A853FC39" box="[1056,1133,979,1005]" country="India" name="Kerala" pageId="10" pageNumber="353">Kerala</collectingRegion>
, collected in 1952, but this location is apparently not published in scientific literature (see further details in Materials and Methods). More recently, C. Susanth, B. V. Premkrishnan and R. Murukesh recorded two individuals, one each from Palode (a small village situated
<quantity id="2E3617A4DA14783A2073855AADA5FB8D" box="[336,411,1086,1113]" metricMagnitude="4" metricUnit="m" metricValue="3.5" pageId="10" pageNumber="353" unit="km" value="35.0">35 km</quantity>
from Thiruvananthapuram) and Kattilappara, Shendurney WLS (https://web.archive.org/ web/20080213063439/http://susanthsbirdsandbutterflies.blog.com/1770510/), in 2000/07 and 2002/05. Mathew and Pulikkal described the early stages and a small population of
<taxonomicName id="2ECEC1C2DA14783A225385E3AFC3FB75" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[880,1021,1159,1185]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA14783A225385E3AFC3FB75" box="[880,1021,1159,1185]" italics="true" pageId="10" pageNumber="353">A. phidippus</emphasis>
</taxonomicName>
from Thenmala (
<bibRefCitation id="8D5FC7B0DA14783A259D85E3ACEAFB11" author="Mathew, G. &amp; Pulikkal, U. K." pageId="10" pageNumber="353" pagination="118 - 120" refId="ref14364" refString="Mathew, G. &amp; Pulikkal, U. K. (2009) Biology of the Palm King Amathusia phidippus, an extremely rare and endangered butterfly of Peninsular India. Journal of the Bombay Natural History Society, 106 (1), 118 - 120." type="journal article" year="2009">Mathew &amp; Pulikkal 2009</bibRefCitation>
). Likewise, Bhakare and Ogale specify the occurrence of
<taxonomicName id="2ECEC1C2DA14783A227D85CFA871FB11" authorityName="Fruhstorfer" authorityYear="1904" box="[862,1103,1195,1221]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="subSpecies" species="phidippus" subSpecies="friderici">
<emphasis id="DBBA6653DA14783A227D85CFA871FB11" box="[862,1103,1195,1221]" italics="true" pageId="10" pageNumber="353">A. phidippus friderici</emphasis>
</taxonomicName>
in “Southern Western Ghats” (
<bibRefCitation id="8D5FC7B0DA14783A21BC85ABAD95FB3D" author="Bhakare, M. &amp; Ogale, H." box="[159,427,1231,1257]" pageId="10" pageNumber="353" refId="ref12582" refString="Bhakare, M. &amp; Ogale, H. (2018) A Guide to Butterflies of Western Ghats (India). Published privately by Milind Bhakare and Hemant Ogale, Satara, 496 pp." type="book" year="2018">Bhakare &amp; Ogale 2018</bibRefCitation>
). There are no specimens in the McGuire Centre for
<taxonomicName id="2ECEC1C2DA14783A252285ABA8B6FB3D" box="[1025,1160,1231,1257]" class="Insecta" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
and Biodiversity, MCZ, or the Smithsonian National Museum of Natural History.
</paragraph>
<paragraph id="E971BA41DA14783A21B4843AAE18FAAD" blockId="10.[151,550,1374,1401]" box="[151,550,1374,1401]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21B4843AAE18FAAD" bold="true" box="[151,550,1374,1401]" pageId="10" pageNumber="353">Species Biology and other Notes</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA14783A21B484C2A908F9A1" pageId="10" pageNumber="353" type="distribution">
<paragraph id="E971BA41DA14783A21B484C2A908F9A1" blockId="10.[151,1437,1446,1869]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21B484C2AD0DFA14" bold="true" box="[151,307,1446,1472]" pageId="10" pageNumber="353">Distribution:</emphasis>
Although the species has been reported broadly from “S.
<collectingCountry id="91D9FAD1DA14783A229884C3AFC9FA15" box="[955,1015,1447,1473]" name="India" pageId="10" pageNumber="353">India</collectingCountry>
” and “Travancore”, the species is not at all widespread. Only four confirmed locations are known (
<figureCitation id="71F5A6C4DA14783A227F84AFAF9AFA31" box="[860,932,1483,1509]" captionStart="FIGURE 6" captionStartId="12.[151,250,1936,1960]" captionTargetBox="[309,1273,928,1907]" captionTargetId="figure-346@12.[303,1283,921,1913]" captionTargetPageId="12" captionText="FIGURE 6. Distributional range map of Amathusia travancorica sp. nov. Map: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386314" httpUri="https://zenodo.org/record/14386314/files/figure.png" pageId="10" pageNumber="353">Fig. 6</figureCitation>
): Thenmala (
<typeStatus id="367504E3DA14783A251E84AFA851FA31" box="[1085,1135,1483,1509]" pageId="10" pageNumber="353">type</typeStatus>
locality) and Kattilappara (both in Shendurney WLS), Palode (Nedumangad Taluk, Thiruvananthapuram District) and Kottayam (Kottayam District, the Sharman location; see Materials and Methods). All four locations are in
<collectingRegion id="2B0A74A3DA14783A255E8777A8F5F9F9" box="[1149,1227,1555,1581]" country="India" name="Kerala" pageId="10" pageNumber="353">Kerala</collectingRegion>
. This narrows the distributional range to southern Travancore, although the species may likely be found in other low-lying evergreen forest patches of Travancore in southern
<collectingRegion id="2B0A74A3DA14783A2343873FAE90F9A1" box="[608,686,1627,1653]" country="India" name="Kerala" pageId="10" pageNumber="353">Kerala</collectingRegion>
. In any case, the species appears to be endemic to
<collectingRegion id="2B0A74A3DA14783A25C0873FA90FF9A1" box="[1251,1329,1627,1653]" country="India" name="Kerala" pageId="10" pageNumber="353">Kerala</collectingRegion>
.
</paragraph>
</subSubSection>
<subSubSection id="A1D4E9CADA14783C21E4871AAFFEFC55" lastPageId="12" lastPageNumber="355" pageId="10" pageNumber="353" type="biology_ecology">
<paragraph id="E971BA41DA14783A21E4871AA86DF899" blockId="10.[151,1437,1446,1869]" pageId="10" pageNumber="353">
<emphasis id="DBBA6653DA14783A21E4871AAFBCF94D" bold="true" box="[199,898,1662,1689]" pageId="10" pageNumber="353">
Status, habitat, and habits:
<taxonomicName id="2ECEC1C2DA14783A232E871AAF1DF94D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[525,803,1662,1689]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="10" pageNumber="353" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA14783A232E871AAF1DF94D" bold="true" box="[525,803,1662,1689]" italics="true" pageId="10" pageNumber="353">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA14783A220A871BAFBCF94D" box="[809,898,1663,1689]" pageId="10" pageNumber="353" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
has been described as ‘Very rare’ (
<bibRefCitation id="8D5FC7B0DA14783A2422871BACEEF969" author="Wynter-Blyth, M. A." pageId="10" pageNumber="353" refId="ref15140" refString="Wynter-Blyth, M. A. (1957) Butterflies of the Indian Region. Bombay Natural History Society, Mumbai, 523 pp." type="book" year="1957">Wynter-Blyth 1957</bibRefCitation>
) and ‘Rare’ (
<bibRefCitation id="8D5FC7B0DA14783A204187C7ADD4F969" author="Evans, W. H." box="[354,490,1699,1725]" pageId="10" pageNumber="353" refId="ref13172" refString="Evans, W. H. (1932) The Identification of Indian Butterflies (2 nd edn.). Bombay Natural History Society, Mumbai, 454 pp." type="book" year="1932">Evans 1932</bibRefCitation>
). All the above historical records appear to be based on less than a dozen sightings in nature and fewer specimens than that ever collected and deposited in research collections. Thus, the species appears to be truly very rare. Published sightings were from densely shaded parts of low-lying evergreen forests, sometimes at forest edges. All the known sightings are below
<quantity id="2E3617A4DA14783A23FA866AAF1FF8FD" box="[729,801,1806,1833]" metricMagnitude="2" metricUnit="m" metricValue="5.0" pageId="10" pageNumber="353" unit="m" value="500.0">500 m</quantity>
asl. Hence, the species appears to be a forest specialist, with a high conservation value due to its narrow distribution and rarity (
<bibRefCitation id="8D5FC7B0DA14783A229E8657A879F899" author="Kunte, K." box="[957,1095,1843,1869]" pageId="10" pageNumber="353" pagination="729 - 735" refId="ref13815" refString="Kunte, K. (2008) The Wildlife (Protection) Act and conservation prioritization of butterflies of the Western Ghats, southwestern India. Current Science, 94 (6), 729 - 735. https: // www. jstor. org / stable / 24100626" type="journal article" year="2008">Kunte 2008</bibRefCitation>
).
</paragraph>
<caption id="BDB1EAC9DA15783B21B4876AAF30F962" ID-DOI="http://doi.org/10.5281/zenodo.14386312" ID-Zenodo-Dep="14386312" httpUri="https://zenodo.org/record/14386312/files/figure.png" pageId="11" pageNumber="354" startId="11.[151,250,1550,1574]" targetBox="[159,1429,181,1526]" targetPageId="11" targetType="figure">
<paragraph id="E971BA41DA15783B21B4876AAF30F962" blockId="11.[151,1437,1550,1718]" pageId="11" pageNumber="354">
<emphasis id="DBBA6653DA15783B21B4876AAF6BF9F2" bold="true" box="[151,853,1550,1574]" pageId="11" pageNumber="354">
FIGURE 5. Early stages of
<taxonomicName id="2ECEC1C2DA15783B20E6876AAEFDF9F2" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[453,707,1550,1574]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="11" pageNumber="354" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA15783B20E6876AAEFDF9F2" bold="true" box="[453,707,1550,1574]" italics="true" pageId="11" pageNumber="354">Amathusia travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA15783B23E8876AAF23F9F2" box="[715,797,1550,1574]" pageId="11" pageNumber="354" rank="species">sp. nov.</taxonomicNameLabel>
a–h:
</emphasis>
5
<superScript id="1EBB1709DA15783B224A876AAF4AF9C8" attach="left" box="[873,884,1550,1564]" fontSize="6" pageId="11" pageNumber="354">th</superScript>
instar caterpillar. 2012/12/23 (a, b: lateral view; d, h: dorsal view; c (frontal) and e (lateral) close-up of the head; f: close-up of abdominal proleg; g: close-up of processes at the tip of the abdomen);
<emphasis id="DBBA6653DA15783B200C8732AD6CF9BA" bold="true" box="[303,338,1622,1646]" pageId="11" pageNumber="354">i–l:</emphasis>
pupa. 2013/01/15 (i: ventral view; j: dorsal view; k: close-up of the head and thorax in dorsal view; l: lateral view);
<emphasis id="DBBA6653DA15783B21FD871EAD2BF946" bold="true" box="[222,277,1658,1682]" pageId="11" pageNumber="354">m–o:</emphasis>
pupa just before eclosion (2013/01/17);
<emphasis id="DBBA6653DA15783B2393871EAEF9F946" bold="true" box="[688,711,1658,1682]" pageId="11" pageNumber="354">p:</emphasis>
eclosed female resting on the pupal case (2013/01/17). Thenmala FRH, Shendurney WLS, Kerala. Photographs: Krushnamegh Kunte.
</paragraph>
</caption>
<paragraph id="E971BA41DA15783C21E4866DA8A1FEB1" blockId="11.[151,1436,1800,2043]" lastBlockId="12.[151,1437,151,897]" lastPageId="12" lastPageNumber="355" pageId="11" pageNumber="354">
Known habits of
<taxonomicName id="2ECEC1C2DA15783B20AB866CAE3EF8F6" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[392,512,1800,1826]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="11" pageNumber="354" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA15783B20AB866CAE3EF8F6" box="[392,512,1800,1826]" italics="true" pageId="11" pageNumber="354">Amathusia</emphasis>
</taxonomicName>
are based largely on descriptions of
<taxonomicName id="2ECEC1C2DA15783B22BB866DA81BF8F6" baseAuthorityName="Linnaeus" baseAuthorityYear="1763" box="[920,1061,1800,1827]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="11" pageNumber="354" phylum="Arthropoda" rank="species" species="phidippus">
<emphasis id="DBBA6653DA15783B22BB866DA81BF8F6" box="[920,1061,1800,1827]" italics="true" pageId="11" pageNumber="354">A. phidippus</emphasis>
</taxonomicName>
in SE Asia, although in historical literature they have been loosely applied to the Western Ghats populations as well, and these agree closely with our own observations in the field. The species is known to be strictly crepuscular, spending the day resting in shaded areas. If disturbed during this time, the butterflies fly only a short distance and settle again, often in more shaded and inaccessible areas. Around sunset they start flying in relatively open areas under closed or partial forest canopy, often flying along the same routes for hours. They may venture in coconut plantations near forests in search of mates and larval host plants (palms; see below). They normally fly relatively slowly and in a bobbing manner to dodge obstructing dense vegetation among which they fly. However, they are capable of flying rapidly if disturbed and efficiently navigate dense vegetation owing to their broad wings, slight flicks of which can help butterflies change course. They settle down to feed on fermenting fruits, tree sap, and droppings of mammalian carnivores as well as herbivores. Similar to other
<taxonomicName id="2ECEC1C2DA12783C20EC8067AE62FEC9" box="[463,604,259,285]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="tribe" tribe="Amathusiini">Amathusiini</taxonomicName>
, they have not been observed feeding on flower nectar. They come to light rarely. One individual recorded by Susanth and team had come to a veranda light in a forest guest house after the rains, and the second—a female—was fluttering by a rattan straggler apparently to lay eggs.
</paragraph>
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<emphasis id="DBBA6653DA12783C21E4800AAD52FE5C" bold="true" box="[199,364,366,392]" pageId="12" pageNumber="355">Flight period:</emphasis>
Sightings of
<emphasis id="DBBA6653DA12783C20DC800BAF31FE5D" bold="true" box="[511,783,367,393]" pageId="12" pageNumber="355">
<taxonomicName id="2ECEC1C2DA12783C20DC800BAE8EFE5D" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[511,688,367,393]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA12783C20DC800BAE8EFE5D" bold="true" box="[511,688,367,393]" italics="true" pageId="12" pageNumber="355">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA12783C2395800BAF31FE5D" box="[694,783,367,393]" pageId="12" pageNumber="355" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
have been infrequent and sporadic. Hence, its flight period and voltinism are poorly understood, although it is expected to be multi-voltine.
</paragraph>
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<emphasis id="DBBA6653DA12783C21E480D3AEBCFE05" bold="true" box="[199,642,439,465]" pageId="12" pageNumber="355">Larval host plants and early stages:</emphasis>
<taxonomicName id="2ECEC1C2DA12783C23AE80D3AF04FE05" box="[653,826,439,465]" class="Liliopsida" family="Arecaceae" genus="Cocos" kingdom="Plantae" order="Arecales" pageId="12" pageNumber="355" phylum="Tracheophyta" rank="species" species="nucifera">
<emphasis id="DBBA6653DA12783C23AE80D3AF04FE05" box="[653,826,439,465]" italics="true" pageId="12" pageNumber="355">Cocos nucifera</emphasis>
</taxonomicName>
and
<taxonomicName id="2ECEC1C2DA12783C225B80D3AFF0FE05" box="[888,974,439,465]" class="Liliopsida" family="Arecaceae" genus="Licuala" kingdom="Plantae" order="Arecales" pageId="12" pageNumber="355" phylum="Tracheophyta" rank="genus">
<emphasis id="DBBA6653DA12783C225B80D3AFF0FE05" box="[888,974,439,465]" italics="true" pageId="12" pageNumber="355">Licuala</emphasis>
</taxonomicName>
(
<taxonomicName id="2ECEC1C2DA12783C22FC80D3A869FE05" box="[991,1111,439,465]" class="Liliopsida" family="Arecaceae" kingdom="Plantae" order="Arecales" pageId="12" pageNumber="355" phylum="Tracheophyta" rank="family">Arecaceae</taxonomicName>
) are commonly recorded as larval host plants of
<emphasis id="DBBA6653DA12783C205F80BFAEAEFE21" bold="true" box="[380,656,475,501]" pageId="12" pageNumber="355">
<taxonomicName id="2ECEC1C2DA12783C205F80BFAE11FE21" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[380,559,475,501]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="species" species="travancorica" status="sp. nov.">
<emphasis id="DBBA6653DA12783C205F80BFAE11FE21" bold="true" box="[380,559,475,501]" italics="true" pageId="12" pageNumber="355">A. travancorica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C089DB28DA12783C231580BFAEAEFE21" box="[566,656,475,501]" pageId="12" pageNumber="355" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, along with
<taxonomicName id="2ECEC1C2DA12783C223F80BFAFCDFE21" box="[796,1011,475,501]" class="Insecta" family="Acrididae" genus="Calamus" kingdom="Animalia" order="Orthoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="species" species="thwaitesii">
<emphasis id="DBBA6653DA12783C223F80BFAFCDFE21" box="[796,1011,475,501]" italics="true" pageId="12" pageNumber="355">Calamus thwaitesii</emphasis>
</taxonomicName>
that is recorded less frequently (
<bibRefCitation id="8D5FC7B0DA12783C244280BFAD27FDCD" author="Nitin, R. &amp; Balakrishnan, V. C. &amp; Churi, P. V &amp; Kalesh, S. &amp; Prakash, S. &amp; Kunte, K." pageId="12" pageNumber="355" pagination="11495 - 11550" refId="ref14504" refString="Nitin, R., Balakrishnan, V. C., Churi, P. V, Kalesh, S., Prakash, S. &amp; Kunte, K. (2018) Larval host plants of the butterflies of the Western Ghats, India. Journal of Threatened Taxa, 10 (4), 11495 - 11550. https: // doi. org / 10.11609 / jott. 3104.10.4.11495 - 11550" type="journal article" year="2018">
Nitin
<emphasis id="DBBA6653DA12783C21B4809BACECFDCD" box="[151,210,511,537]" italics="true" pageId="12" pageNumber="355">et al.</emphasis>
2018
</bibRefCitation>
). Susanth
<emphasis id="DBBA6653DA12783C20B2809BADF2FDCD" box="[401,460,511,537]" italics="true" pageId="12" pageNumber="355">et al.</emphasis>
(on the blog post cited above) suggest that the species may feed on rattan palms in dense forests. It is possible that the species feeds on almost any available palm in its habitat. Leaves of the exotic, ornamental palm
<taxonomicName id="2ECEC1C2DA12783C207D8323AE24FDB5" box="[350,538,583,609]" class="Liliopsida" family="Arecaceae" genus="Dypsis" kingdom="Plantae" order="Arecales" pageId="12" pageNumber="355" phylum="Tracheophyta" rank="species" species="lutescens">
<emphasis id="DBBA6653DA12783C207D8323AE24FDB5" box="[350,538,583,609]" italics="true" pageId="12" pageNumber="355">Dypsis lutescens</emphasis>
</taxonomicName>
were readily accepted by caterpillars, on which the lifecycle was completed in captivity (
<bibRefCitation id="8D5FC7B0DA12783C2024830FAE18FD51" author="Mathew, G. &amp; Pulikkal, U. K." box="[263,550,619,645]" pageId="12" pageNumber="355" pagination="118 - 120" refId="ref14364" refString="Mathew, G. &amp; Pulikkal, U. K. (2009) Biology of the Palm King Amathusia phidippus, an extremely rare and endangered butterfly of Peninsular India. Journal of the Bombay Natural History Society, 106 (1), 118 - 120." type="journal article" year="2009">Mathew &amp; Pulikkal 2009</bibRefCitation>
). The early stages are documented in detail (
<bibRefCitation id="8D5FC7B0DA12783C2533830FA910FD51" author="Mathew, G. &amp; Pulikkal, U. K." box="[1040,1326,619,645]" pageId="12" pageNumber="355" pagination="118 - 120" refId="ref14364" refString="Mathew, G. &amp; Pulikkal, U. K. (2009) Biology of the Palm King Amathusia phidippus, an extremely rare and endangered butterfly of Peninsular India. Journal of the Bombay Natural History Society, 106 (1), 118 - 120." type="journal article" year="2009">Mathew &amp; Pulikkal 2009</bibRefCitation>
), and also illustrated here with additional images showing close-ups of caterpillar head and different views of the pupa that have not been illustrated in previous literature (
<figureCitation id="71F5A6C4DA12783C238A83D7AED0FD19" box="[681,750,690,717]" captionStart="FIGURE 5" captionStartId="11.[151,250,1550,1574]" captionTargetBox="[159,1429,181,1526]" captionTargetId="figure-19@11.[159,1429,181,1526]" captionTargetPageId="11" captionText="FIGURE 5. Early stages of Amathusia travancorica sp. nov. a–h: 5th instar caterpillar. 2012/12/23 (a, b: lateral view; d, h: dorsal view; c (frontal) and e (lateral) close-up of the head; f: close-up of abdominal proleg; g: close-up of processes at the tip of the abdomen); i–l: pupa. 2013/01/15 (i: ventral view; j: dorsal view; k: close-up of the head and thorax in dorsal view; l: lateral view); m–o: pupa just before eclosion (2013/01/17); p: eclosed female resting on the pupal case (2013/01/17). Thenmala FRH, Shendurney WLS, Kerala. Photographs: Krushnamegh Kunte." figureDoi="http://doi.org/10.5281/zenodo.14386312" httpUri="https://zenodo.org/record/14386312/files/figure.png" pageId="12" pageNumber="355">Fig. 5</figureCitation>
).
</paragraph>
<paragraph id="E971BA41DA12783C21E483B2AD10FCED" blockId="12.[151,1437,151,897]" pageId="12" pageNumber="355">
<emphasis id="DBBA6653DA12783C21E483B2ADE8FD25" bold="true" box="[199,470,726,753]" pageId="12" pageNumber="355">
Sympatric
<taxonomicName id="2ECEC1C2DA12783C206C83B2ADF3FD25" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[335,461,726,753]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA12783C206C83B2ADF3FD25" bold="true" box="[335,461,726,753]" italics="true" pageId="12" pageNumber="355">Amathusia</emphasis>
</taxonomicName>
:
</emphasis>
No other
<taxonomicName id="2ECEC1C2DA12783C237583B3AEF0FD25" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[598,718,727,753]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="genus">
<emphasis id="DBBA6653DA12783C237583B3AEF0FD25" box="[598,718,727,753]" italics="true" pageId="12" pageNumber="355">Amathusia</emphasis>
</taxonomicName>
occurs in the Western Ghats. The only other Indian species,
<taxonomicName id="2ECEC1C2DA12783C21B4839FAEAEFCC1" authority="Fruhstorfer, 1899" authorityName="Fruhstorfer" authorityYear="1899" box="[151,656,763,789]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="species" species="andamanensis">
<emphasis id="DBBA6653DA12783C21B4839FAD84FCC1" box="[151,442,763,789]" italics="true" pageId="12" pageNumber="355">Amathusia andamanensis</emphasis>
Fruhstorfer, 1899
</taxonomicName>
, is endemic to the Andaman Islands (
<bibRefCitation id="8D5FC7B0DA12783C256C839FA8C3FCC1" author="d'Abrera, B." box="[1103,1277,762,789]" pageId="12" pageNumber="355" refId="ref12980" refString="d'Abrera, B. (1985) Butterflies of the Oriental Region. Part II: Nymphalidae, Satyridae &amp; Amathusidae. Hill House, Melbourne, 534 pp." type="book" year="1985">d’Abrera 1985</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA12783C242E839FA9A6FCC1" author="Evans, W. H." box="[1293,1432,763,789]" pageId="12" pageNumber="355" refId="ref13172" refString="Evans, W. H. (1932) The Identification of Indian Butterflies (2 nd edn.). Bombay Natural History Society, Mumbai, 454 pp." type="book" year="1932">Evans 1932</bibRefCitation>
;
<bibRefCitation id="8D5FC7B0DA12783C21B4827BAD1FFCED" author="Talbot, G." box="[151,289,799,825]" pageId="12" pageNumber="355" refId="ref15075" refString="Talbot, G. (1947) The Fauna of British India, including Ceylon and Burma: Butterflies, Vol. 2. Taylor &amp; Francis, London, 506 pp." type="book" year="1947">Talbot 1947</bibRefCitation>
).
</paragraph>
<paragraph id="E971BA41DA12783C21E48227AFFEFC55" blockId="12.[151,1437,151,897]" pageId="12" pageNumber="355">
Further information will be made available on the species page of the Butterflies of
<collectingCountry id="91D9FAD1DA12783C254B8227A89CFC89" box="[1128,1186,835,861]" name="India" pageId="12" pageNumber="355">India</collectingCountry>
website (https://www. ifoundbutterflies.org/
<taxonomicName id="2ECEC1C2DA12783C20A58203AE3FFC55" authorityName="Kunte &amp; Basu" authorityYear="2024" box="[390,513,871,897]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="12" pageNumber="355" phylum="Arthropoda" rank="genus">Amathusia</taxonomicName>
) as additional observations accumulate.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title id="7509A47928EAB1A35144DBFD8E2B0177">Two new taxa of brush-footed butterflies (Lepidoptera: Nymphalidae) from the Western Ghats biodiversity hotspot, southern India</mods:title>
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2.
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<taxonomicName id="2ECEC1C2DA1F783121E48617ADC0F85A" ID-CoL="5TYM5" authority="Fruhstorfer, 1899" authorityName="Fruhstorfer" authorityYear="1899" box="[199,510,1907,1934]" class="Insecta" family="Nymphalidae" genus="Amathusia" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="344" phylum="Arthropoda" rank="species" species="andamanensis">
<emphasis id="DBBA6653DA1F783121E48617ADC0F85A" bold="true" box="[199,510,1907,1934]" italics="true" pageId="1" pageNumber="344">Amathusia andamanensis</emphasis>
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:
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:
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(E) #984596 (from the collection of
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),
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:
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(E) #984589 (
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),
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specimens from
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,
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, now in
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dates unknown, presumably between mid-1800s and 1920s
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.
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<emphasis id="00AB17693207FFCBFB93FE6B78F354E2" bold="true" box="[1034,1266,427,453]" pageId="4">Phylogenetic analyses</emphasis>
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The MRBAYES and BEAST produced similar topologies. Bayesian inference recovered almost all genera with posterior probabilities&gt;0.95 (
<figureCitation id="AAE4D7FE3207FFCBFB90FDD07840570F" box="[1033,1089,528,552]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="4">Fig. 3</figureCitation>
). Most genera were monophyletic except the following: (i)
<taxonomicName id="F5DFB0F83207FFCBFBBCFDF0795B5760" authority="(Bean, 1887)" baseAuthorityName="Bean" baseAuthorityYear="1887" box="[1061,1370,559,583]" family="Goodeidae" genus="Xenotoca" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="variata">
<emphasis id="00AB17693207FFCBFBBCFDF078CD5760" box="[1061,1228,560,583]" italics="true" pageId="4">Xenotoca variata</emphasis>
(Bean, 1887)
</taxonomicName>
was sister to
<taxonomicName id="F5DFB0F83207FFCBFCCFFD8F79195740" authority="Miller &amp; Fitzsimons, 1971" authorityName="Miller &amp; Fitzsimons" authorityYear="1971" box="[854,1304,591,615]" family="Goodeidae" genus="Ameca" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="splendens">
<emphasis id="00AB17693207FFCBFCCFFD8F7FFD5740" box="[854,1020,591,615]" italics="true" pageId="4">Ameca splendens</emphasis>
Miller &amp; Fitzsimons, 1971
</taxonomicName>
and placed well outside genus
<taxonomicName id="F5DFB0F83207FFCBFC48FDAF792957A1" authority="Hubbs &amp; Turner, 1939" authorityName="Hubbs &amp; Turner" authorityYear="1939" box="[977,1320,622,646]" family="Goodeidae" genus="Xenotoca" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFC48FDAF782A57A1" box="[977,1067,623,646]" italics="true" pageId="4">Xenotoca</emphasis>
Hubbs &amp; Turner, 1939
</taxonomicName>
; (ii)
<taxonomicName id="F5DFB0F83207FFCBFAC6FDAE78275782" authority="(Hubbs, 1936)" baseAuthorityName="Hubbs" baseAuthorityYear="1936" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="pulchra">
<emphasis id="00AB17693207FFCBFAC6FDAE7F835782" italics="true" pageId="4">Jordanella pulchra</emphasis>
(Hubbs, 1936)
</taxonomicName>
was sister to
<taxonomicName id="F5DFB0F83207FFCBFB2DFD4D79BF5782" authority="Hubbs, 1926" authorityName="Hubbs" authorityYear="1926" box="[1204,1470,653,677]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFB2DFD4D79335782" box="[1204,1330,653,677]" italics="true" pageId="4">Floridichthys</emphasis>
Hubbs, 1926
</taxonomicName>
, while
<taxonomicName id="F5DFB0F83207FFCBFCE4FD6D792F57E2" authority="Goode &amp; Bean, 1879" authorityName="Goode &amp; Bean" authorityYear="1879" box="[893,1326,685,709]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="floridae">
<emphasis id="00AB17693207FFCBFCE4FD6D783757E2" box="[893,1078,685,709]" italics="true" pageId="4">Jordanella floridae</emphasis>
Goode &amp; Bean, 1879
</taxonomicName>
was sister to
<taxonomicName id="F5DFB0F83207FFCBFCA0FD0C78CA57C3" authority="Miller and Walters, 1972" authorityName="Miller and Walters" authorityYear="1972" box="[825,1227,716,740]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFCA0FD0C7FA857C3" box="[825,937,716,740]" italics="true" pageId="4">Megupsilon</emphasis>
Miller and Walters, 1972
</taxonomicName>
<emphasis id="00AB17693207FFCBFB52FD0C78D957C3" box="[1227,1240,716,740]" italics="true" pageId="4">–</emphasis>
<taxonomicName id="F5DFB0F83207FFCBFB41FD0C7F715624" authority="Lacepede, 1803" authorityName="Lacepede" authorityYear="1803" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFB41FD0C794C57C3" box="[1240,1357,716,740]" italics="true" pageId="4">Cyprinodon</emphasis>
Lacépède, 1803
</taxonomicName>
; and (iii)
<taxonomicName id="F5DFB0F83207FFCBFC42FD2C78B95624" authority="Girard, 1860" authorityName="Girard" authorityYear="1860" box="[987,1208,747,771]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFC42FD2C782B5624" box="[987,1066,748,771]" italics="true" pageId="4">Lucania</emphasis>
Girard, 1860
</taxonomicName>
was sister to
<taxonomicName id="F5DFB0F83207FFCBFAD1FD2B782C5604" authority="Ghedotti &amp; Davis, 2013" authorityName="Ghedotti &amp; Davis" authorityYear="2013" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="genus">
<emphasis id="00AB17693207FFCBFAD1FD2B79C35624" box="[1352,1474,747,771]" italics="true" pageId="4">Wileyichthys</emphasis>
<bibRefCitation id="564EB68A3207FFCBFCA0FCCB782C5604" author="Ghedotti MJ &amp; Davis MP" box="[825,1069,779,803]" pageId="4" pagination="1 - 65" refId="ref16579" refString="Ghedotti MJ, Davis MP. Phylogeny, classification, and evolution of salinity tolerance of the North American topminnows and killifishes, family Fundulidae (Teleostei: Cyprinodontiformes). Fieldiana Life and Earth Sciences 2013; 7: 1 - 65. https: // doi. org / 10.3158 / 2158 - 5520 - 12.7.1" type="journal article" year="2013">Ghedotti &amp; Davis, 2013</bibRefCitation>
</taxonomicName>
(
<taxonomicName id="F5DFB0F83207FFCBFBA4FCCB795A5604" authority="Vaillant, 1894" authorityName="Vaillant" authorityYear="1894" box="[1085,1371,779,803]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="lima">
<emphasis id="00AB17693207FFCBFBA4FCCB78C85604" box="[1085,1225,779,803]" italics="true" pageId="4">Fundulus lima</emphasis>
Vaillant, 1894
</taxonomicName>
<emphasis id="00AB17693207FFCBFAC2FCCB79695604" box="[1371,1384,779,803]" italics="true" pageId="4">–</emphasis>
<taxonomicName id="F5DFB0F83207FFCBFAFEFCCB78355665" authority="Girard, 1856" authorityName="Girard" authorityYear="1856" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="parvipinnis">
<emphasis id="00AB17693207FFCBFAFEFCCB7FA65665" italics="true" pageId="4">Fundulus parvipinnis</emphasis>
Girard, 1856
</taxonomicName>
).
</paragraph>
<paragraph id="3260CB7B3207FFC8FCCCFC8A7EB5565E" blockId="4.[825,1475,427,1147]" lastBlockId="7.[113,764,582,889]" lastPageId="7" pageId="4">
In our phylogeny, the families
<taxonomicName id="F5DFB0F83207FFCBFA94FC8979BF5646" box="[1293,1470,841,865]" pageId="4">Cubanichthyidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83207FFCBFCA0FCA97FED56A6" box="[825,1004,873,897]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83207FFCBFC63FCA9787156A6" box="[1018,1136,873,897]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83207FFCBFBE7FCA978F056A6" box="[1150,1265,873,897]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
, and
<taxonomicName id="F5DFB0F83207FFCBFAB6FCA979C356A6" box="[1327,1474,873,897]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
were recovered as composing a monophyletic group, in agreement with recent studies (
<bibRefCitation id="564EB68A3207FFCBFBD7FC6779445698" author="Ghezelayagh A &amp; Harrington RC &amp; Burress ED" box="[1102,1349,935,959]" pageId="4" pagination="1211 - 20" refId="ref16688" refString="Ghezelayagh A, Harrington RC, Burress ED et al. Prolonged morphological expansion of spiny-rayed fishes following the end- Cretaceous. Nature Ecology &amp; Evolution 2022; 6: 1211 - 20. https: // doi. org / 10.1038 / s 41559 - 022 - 01801 - 3" type="journal article" year="2022">
Ghezelayagh
<emphasis id="00AB17693207FFCBFB40FC68790B5698" box="[1241,1290,935,959]" italics="true" pageId="4">et al.</emphasis>
2022
</bibRefCitation>
,
<bibRefCitation id="564EB68A3207FFCBFACAFC677F6C56F8" author="Piller KR &amp; Parker E &amp; Lemmon AR" pageId="4" refId="ref19125" refString="Piller KR, Parker E, Lemmon AR et al. Investigating the utility of anchored hybrid enrichment data to investigate the relationships among the killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed group of fishes. Molecular Phylogenetics and Evolution 2022; 173: 107482. https: // doi. org / 10.1016 / j. ympev. 2022.107482" type="journal volume" year="2022">
Piller
<emphasis id="00AB17693207FFCBFA0BFC6879C25698" box="[1426,1475,935,959]" italics="true" pageId="4">et al.</emphasis>
2022
</bibRefCitation>
). The branching pattern among families in this clade was in agreement with genomic results provided by
<bibRefCitation id="564EB68A3207FFCBFA92FC267F85513A" author="Ghezelayagh A &amp; Harrington RC &amp; Burress ED" pageId="4" pagination="1211 - 20" refId="ref16688" refString="Ghezelayagh A, Harrington RC, Burress ED et al. Prolonged morphological expansion of spiny-rayed fishes following the end- Cretaceous. Nature Ecology &amp; Evolution 2022; 6: 1211 - 20. https: // doi. org / 10.1038 / s 41559 - 022 - 01801 - 3" type="journal article" year="2022">
Ghezelayagh
<emphasis id="00AB17693207FFCBFA0AFC2779C256D9" box="[1427,1475,998,1022]" italics="true" pageId="4">et al.</emphasis>
(2022)
</bibRefCitation>
and
<bibRefCitation id="564EB68A3207FFCBFC2AFBC5786A513A" author="Piller KR &amp; Parker E &amp; Lemmon AR" box="[947,1131,1029,1053]" pageId="4" refId="ref19125" refString="Piller KR, Parker E, Lemmon AR et al. Investigating the utility of anchored hybrid enrichment data to investigate the relationships among the killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed group of fishes. Molecular Phylogenetics and Evolution 2022; 173: 107482. https: // doi. org / 10.1016 / j. ympev. 2022.107482" type="journal volume" year="2022">
Piller
<emphasis id="00AB17693207FFCBFC76FBC6781C513A" box="[1007,1053,1029,1053]" italics="true" pageId="4">et al.</emphasis>
(2022)
</bibRefCitation>
. However, in contrast to their and our findings, which place Cubanichthyidae sister to
<taxonomicName id="F5DFB0F83207FFCBFADBFBE579B5511A" box="[1346,1460,1061,1085]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83207FFCBFCA0FB847899517B" authority=", Piller et al. (2022)" authorityName="Piller" authorityYear="2022" box="[825,1176,1092,1116]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">
Profundulidae,
<bibRefCitation id="564EB68A3207FFCBFC41FB847899517B" author="Piller KR &amp; Parker E &amp; Lemmon AR" box="[984,1176,1092,1116]" pageId="4" refId="ref19125" refString="Piller KR, Parker E, Lemmon AR et al. Investigating the utility of anchored hybrid enrichment data to investigate the relationships among the killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed group of fishes. Molecular Phylogenetics and Evolution 2022; 173: 107482. https: // doi. org / 10.1016 / j. ympev. 2022.107482" type="journal volume" year="2022">
Piller
<emphasis id="00AB17693207FFCBFB8FFB857846517B" box="[1046,1095,1092,1116]" italics="true" pageId="4">et al.</emphasis>
(2022)
</bibRefCitation>
</taxonomicName>
found that Cubanichthyidae was paraphyletic, with
<taxonomicName id="F5DFB0F83207FFC8FBDFFBA37CB05779" authority="(Eigenmann, 1903)" baseAuthorityName="Eigenmann" baseAuthorityYear="1903" family="Cyprinodontidae" genus="Cubanichthys" kingdom="Animalia" lastPageId="7" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="species" species="cubensis">
<emphasis id="00AB17693207FFCBFBDFFBA3792A515C" box="[1094,1323,1123,1147]" italics="true" pageId="4">Cubanichthys cubensis</emphasis>
(Eigenmann, 1903)
</taxonomicName>
sister to
<taxonomicName id="F5DFB0F83204FFC8FEAAFD867DE75779" box="[307,486,582,606]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
, and with
<taxonomicName id="F5DFB0F83204FFC8FDEFFD867D715759" authority="(Fowler, 1930)" baseAuthorityName="Fowler" baseAuthorityYear="1930" family="Cyprinodontidae" genus="Cubanichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="pengelleyi">
<emphasis id="00AB17693204FFC8FDEFFD867CCE575A" italics="true" pageId="7">Cubanichthys pengelleyi</emphasis>
(Fowler, 1930)
</taxonomicName>
sister to
<taxonomicName id="F5DFB0F83204FFC8FE52FDA67E34575A" baseAuthorityName="Eigenmann" baseAuthorityYear="1903" box="[459,565,613,637]" family="Cyprinodontidae" genus="Cubanichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="cubensis">
<emphasis id="00AB17693204FFC8FE52FDA67E34575A" box="[459,565,613,637]" italics="true" pageId="7">C. cubensis</emphasis>
</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FDDAFDA57EF6575A" box="[579,759,613,637]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
. Confidence intervals for divergence estimates of family originations in our tree (nodes 1–4;
<tableCitation id="7F5DFEC03204FFC8FE45FD647E27579B" box="[476,550,676,700]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="7">Table 2</tableCitation>
) overlapped broadly with confidence intervals for the same nodes in the study by
<bibRefCitation id="564EB68A3204FFC8FFE8FD237D7A57DC" author="Ghezelayagh A &amp; Harrington RC &amp; Burress ED" box="[113,379,739,763]" pageId="7" pagination="1211 - 20" refId="ref16688" refString="Ghezelayagh A, Harrington RC, Burress ED et al. Prolonged morphological expansion of spiny-rayed fishes following the end- Cretaceous. Nature Ecology &amp; Evolution 2022; 6: 1211 - 20. https: // doi. org / 10.1038 / s 41559 - 022 - 01801 - 3" type="journal article" year="2022">
Ghezelayagh
<emphasis id="00AB17693204FFC8FF62FD247D2A57DC" box="[251,299,739,763]" italics="true" pageId="7">et al.</emphasis>
(2022)
</bibRefCitation>
. The greatest difference was for node 4 (separation of
<taxonomicName id="F5DFB0F83204FFC8FEBCFCC27D96563D" box="[293,407,770,794]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
and
<taxonomicName id="F5DFB0F83204FFC8FE49FCC27E69563D" box="[464,616,770,794]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
), wherein our confidence intervals overlapped with theirs, but their range also extended into more recent time, with their point estimate of ~22 Mya being substantially younger than ours of ~30.0 Mya.
</paragraph>
<caption id="66A09BF33207FFCBFF18F88E7D53529C" pageId="4" startId="4.[129,194,1870,1894]" targetBox="[132,1470,1224,1839]" targetPageId="4" targetType="figure">
<paragraph id="3260CB7B3207FFCBFF18F88E7D53529C" blockId="4.[129,1469,1870,1979]" pageId="4">
<emphasis id="00AB17693207FFCBFF18F88E7CD95240" bold="true" box="[129,216,1870,1895]" pageId="4">Figure 2.</emphasis>
Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae,
<taxonomicName id="F5DFB0F83207FFCBFBC6F88F79045240" box="[1119,1285,1871,1895]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83207FFCBFA94F88F797B5240" box="[1293,1402,1871,1895]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83207FFCBFF18F8AB7CED52A4" box="[129,236,1899,1923]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
, and
<taxonomicName id="F5DFB0F83207FFCBFE85F8AB7DA752A4" box="[284,422,1899,1923]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
. A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both.
</paragraph>
</caption>
<caption id="66A09BF33206FFCAFFE8F8EB7E9A5294" pageId="5" startId="5.[113,178,1835,1859]" targetBox="[192,1381,152,1803]" targetPageId="5" targetType="figure">
<paragraph id="3260CB7B3206FFCAFFE8F8EB7E9A5294" blockId="5.[113,1418,1835,1971]" pageId="5">
<emphasis id="00AB17693206FFCAFFE8F8EB7CC85264" bold="true" box="[113,201,1835,1859]" pageId="5">Figure 3.</emphasis>
Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae,
<taxonomicName id="F5DFB0F83206FFCAFFE8F8877D165278" box="[113,279,1863,1887]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83206FFCAFE86F8877D8D5278" box="[287,396,1863,1887]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83206FFCAFE0CF8877E015278" box="[405,512,1863,1887]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
, and
<taxonomicName id="F5DFB0F83206FFCAFDA9F8877EB95278" box="[560,696,1863,1887]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species. Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2).
</paragraph>
</caption>
<caption id="66A09BF33205FFC9FF18FF4F7E0E55F8" pageId="6" startId="6.[129,184,143,167]" targetBox="[129,1472,244,1905]" targetIsTable="true" targetPageId="6" targetType="table">
<paragraph id="3260CB7B3205FFC9FF18FF4F7E0E55F8" blockId="6.[129,1412,142,223]" pageId="6">
<emphasis id="00AB17693205FFC9FF18FF4F7CCF5581" bold="true" box="[129,206,142,167]" pageId="6">Table 2.</emphasis>
Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig. 3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided. Further description is provided in the main text, with references.
</paragraph>
</caption>
<paragraph id="3260CB7B3205FFC9FF18FF3578A65256" pageId="6">
<table id="40DF39DB32050030FF18FF3479C15256" box="[129,1472,244,1905]" colsContinueIn="7.[113,1456,203,485]" gridcols="4" gridrows="16" pageId="6">
<tr id="8CEFC93932050030FF18FF3479C1540D" box="[129,1472,244,298]" gridrow="0" pageId="6">
<th id="CF3EA04532050030FF18FF347CB9540D" box="[129,184,244,298]" gridcol="0" gridrow="0" pageId="6">
<emphasis id="00AB17693205FFC9FF18FF357CB9542A" bold="true" box="[129,184,245,269]" pageId="6">Node</emphasis>
</th>
<th id="CF3EA04532050030FF69FF347E82540D" box="[240,643,244,298]" gridcol="1" gridrow="0" pageId="6">
<emphasis id="00AB17693205FFC9FF69FF347DAA542B" bold="true" box="[240,427,244,268]" pageId="6">Divergence (node)</emphasis>
</th>
<th id="CF3EA04532050030FD32FF347805540D" box="[683,1028,244,298]" gridcol="2" gridrow="0" pageId="6">
<emphasis id="00AB17693205FFC9FD32FF347805540E" bold="true" pageId="6">Age estimate (Mya) (95% highest posterior density CI)</emphasis>
</th>
<th id="CF3EA04532050030FBB1FF3479C1540D" box="[1064,1472,244,298]" gridcol="3" gridrow="0" pageId="6">
<emphasis id="00AB17693205FFC9FBB1FF347915542B" bold="true" box="[1064,1300,244,268]" pageId="6">Reciprocal illumination</emphasis>
</th>
</tr>
<tr id="8CEFC93932050030FF18FE8179C154E8" box="[129,1472,321,463]" gridrow="1" pageId="6">
<th id="CF3EA04532050030FF18FE817CB954E8" box="[129,184,321,463]" gridcol="0" gridrow="1" pageId="6">1</th>
<td id="CF3EA04532050030FF69FE817E8254E8" box="[240,643,321,463]" gridcol="1" gridrow="1" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FE827D97547D" box="[240,406,322,346]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FE3BFE827E10547D" box="[418,529,322,346]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
/ Cubanichthyidae–
<taxonomicName id="F5DFB0F83205FFC9FE2CFE9F7E1F5450" box="[437,542,351,375]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FE9CFEBC7D8C54B3" box="[261,397,380,404]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
</td>
<td id="CF3EA04532050030FD32FE81780554E8" box="[683,1028,321,463]" gridcol="2" gridrow="1" pageId="6">52.3 (58.62–45.79)</td>
<td id="CF3EA04532050030FBB1FE8179C154E8" box="[1064,1472,321,463]" gridcol="3" gridrow="1" pageId="6">59.2–47.8 Mya: Tropic of Cancer retracted to 27°N (Nearctic–Neotropical subdivision, ecological diversification) 55.8–55.0 Mya: Gulf of México drawdown (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FE1979C1576E" box="[129,1472,473,585]" gridrow="2" pageId="6">
<th id="CF3EA04532050030FF18FE197CB9576E" box="[129,184,473,585]" gridcol="0" gridrow="2" pageId="6">2</th>
<td id="CF3EA04532050030FF69FE197E82576E" box="[240,643,473,585]" gridcol="1" gridrow="2" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FE197D9754D6" box="[240,406,473,497]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FE04FE197E0D54D6" box="[413,524,473,497]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
</td>
<td id="CF3EA04532050030FD32FE197805576E" box="[683,1028,473,585]" gridcol="2" gridrow="2" pageId="6">46.2 (54.77–37.63)</td>
<td id="CF3EA04532050030FBB1FE1979C1576E" box="[1064,1472,473,585]" gridcol="3" gridrow="2" pageId="6">48.2 Mya: North American drainage reorganization (dispersal–vicariance cycle) 47.5, 46.9, and 45.6 Mya: Sea-level minima (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FD9379C157E4" box="[129,1472,595,707]" gridrow="3" pageId="6">
<th id="CF3EA04532050030FF18FD937CB957E4" box="[129,184,595,707]" gridcol="0" gridrow="3" pageId="6">3</th>
<td id="CF3EA04532050030FF69FD93780557E4" box="[240,1028,595,707]" colspan="2" colspanRight="1" gridcol="1" gridrow="3" pageId="6">
Cubanichthyidae/
<taxonomicName id="F5DFB0F83205FFC9FE05FD937E04574C" box="[412,517,595,619]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FD88FD937E9D574C" box="[529,668,595,619]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
45.9 (54.82–35.79)
</td>
<td id="CF3EA04532050030FBB1FD9379C157E4" box="[1064,1472,595,707]" gridcol="3" gridrow="3" pageId="6">48.0 Mya: maximum sea level (ice-free world) (dispersal) 47.5, 46.9, and 45.6 Mya: sea-level minima (glaciation) (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FD0C79C156B3" box="[129,1472,716,916]" gridrow="4" pageId="6">
<th id="CF3EA04532050030FF18FD0C7CB956B3" box="[129,184,716,916]" gridcol="0" gridrow="4" pageId="6">4</th>
<td id="CF3EA04532050030FF69FD0C7E8256B3" box="[240,643,716,916]" gridcol="1" gridrow="4" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FD0C7D5B57C3" box="[240,346,716,740]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FEF8FD0C7DEC57C3" box="[353,493,716,740]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
</td>
<td id="CF3EA04532050030FD32FD0C780556B3" box="[683,1028,716,916]" gridcol="2" gridrow="4" pageId="6">30.0 (39.19–21.38)</td>
<td id="CF3EA04532050030FBB1FD0C79C156B3" box="[1064,1472,716,916]" gridcol="3" gridrow="4" pageId="6">
35.0–25.0 Mya: Chiapas Massif uplift and drainage capture (
<taxonomicName id="F5DFB0F83205FFC9FB7CFD2A79715625" box="[1253,1392,746,770]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
vicariance) 33.9 Mya: Madrean River capture (
<taxonomicName id="F5DFB0F83205FFC9FBDAFC8278AE567D" box="[1091,1199,834,858]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
vicariance) 31.8 Mya: cyprinodontidae southward immigration (ecological replacement)
</td>
</tr>
<tr id="8CEFC93932050030FF18FC5E79C156D6" box="[129,1472,926,1009]" gridrow="5" pageId="6">
<th id="CF3EA04532050030FF18FC5E7CB956D6" box="[129,184,926,1009]" gridcol="0" gridrow="5" pageId="6">5</th>
<td id="CF3EA04532050030FF69FC5E7E8256D6" box="[240,643,926,1009]" gridcol="1" gridrow="5" pageId="6">
<emphasis id="00AB17693205FFC9FF69FC5E7DA35691" box="[240,418,926,950]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FC5E7D605691" authorityName="Hubbs" authorityYear="1924" box="[240,353,926,950]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Profundulus</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FEF2FC5E7DA35691" authorityName="Alvarez and Carranza" authorityYear="1951" box="[363,418,926,950]" family="Profundulidae" genus="Tlaloc" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Tlaloc</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32FC5E780556D6" box="[683,1028,926,1009]" gridcol="2" gridrow="5" pageId="6">22.1 (30.66–14.19)</td>
<td id="CF3EA04532050030FBB1FC5E79C156D6" box="[1064,1472,926,1009]" gridcol="3" gridrow="5" pageId="6">35.0–25.0 Mya: uplift and erosion of Chiapas Massif foreland captures Upper Río Grijalva (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FC3B79C1516A" box="[129,1472,1019,1101]" gridrow="6" pageId="6">
<th id="CF3EA04532050030FF18FC3B7CB9516A" box="[129,184,1019,1101]" gridcol="0" gridrow="6" pageId="6">6</th>
<td id="CF3EA04532050030FF69FC3B7E82516A" box="[240,643,1019,1101]" gridcol="1" gridrow="6" pageId="6">
Empetrichthyinae/
<taxonomicName id="F5DFB0F83205FFC9FE3FFC3B7E0E5134" box="[422,527,1019,1043]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
</td>
<td id="CF3EA04532050030FD32FC3B7805516A" box="[683,1028,1019,1101]" gridcol="2" gridrow="6" pageId="6">17.9 (23.32–12.90)</td>
<td id="CF3EA04532050030FBB1FC3B79C1516A" box="[1064,1472,1019,1101]" gridcol="3" gridrow="6" pageId="6">23.0 Mya: North American drainage reorganization, Great Basin tectonism (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FB9779C15025" box="[129,1472,1111,1282]" gridrow="7" pageId="6">
<th id="CF3EA04532050030FF18FB977CB95025" box="[129,184,1111,1282]" gridcol="0" gridrow="7" pageId="6">7</th>
<td id="CF3EA04532050030FF69FB977E825025" box="[240,643,1111,1282]" gridcol="1" gridrow="7" pageId="6">
Characodontini–Ilyodontini/crown
<taxonomicName id="F5DFB0F83205FFC9FE9CFBB57D6F51AA" box="[261,366,1141,1165]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
</td>
<td id="CF3EA04532050030FD32FB9778055025" box="[683,1028,1111,1282]" gridcol="2" gridrow="7" pageId="6">13.4 (16.78–10.25)</td>
<td id="CF3EA04532050030FBB1FB9779C15025" box="[1064,1472,1111,1282]" gridcol="3" gridrow="7" pageId="6">14.0–5.0 Mya: San Marcos Fault reactivation (vicariance) 13.6–10.6 Mya: Los Encinos volcanism (San Luis Potosí) (vicariance) 12.0 Mya: Metates Volcanism (Durango) (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FACC79C15066" box="[129,1472,1292,1345]" gridrow="8" pageId="6">
<th id="CF3EA04532050030FF18FACC7CB95066" box="[129,184,1292,1345]" gridcol="0" gridrow="8" pageId="6">8</th>
<td id="CF3EA04532050030FF69FACC7E825066" box="[240,643,1292,1345]" gridcol="1" gridrow="8" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FACC7D985003" box="[240,409,1292,1316]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
/
<emphasis id="00AB17693205FFC9FE3CFACC7D485066" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FE3CFACC7E185003" authorityName="Hubbs" authorityYear="1926" box="[421,537,1292,1316]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FDBDFACC7D485066" baseAuthorityName="Hubbs" baseAuthorityYear="1936" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="species" species="pulchra">Jordanella pulchra</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32FACC78055066" box="[683,1028,1292,1345]" gridcol="2" gridrow="8" pageId="6">31.8 (42.33–21.61)</td>
<td id="CF3EA04532050030FBB1FACC79C15066" box="[1064,1472,1292,1345]" gridcol="3" gridrow="8" pageId="6">33.9–26.0 Mya: Oligocene icehouse (dispersal–vicariance cycle)</td>
</tr>
<tr id="8CEFC93932050030FF18FA8B79C150B9" box="[129,1472,1355,1438]" gridrow="9" pageId="6">
<th id="CF3EA04532050030FF18FA8B7CB950B9" box="[129,184,1355,1438]" gridcol="0" gridrow="9" pageId="6">9</th>
<td id="CF3EA04532050030FF69FA8B7E8250B9" box="[240,643,1355,1438]" gridcol="1" gridrow="9" pageId="6">
<emphasis id="00AB17693205FFC9FF69FA8B7E825044" box="[240,643,1355,1379]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FA8B7D5B5044" authorityName="Lacepede" authorityYear="1803" box="[240,346,1355,1379]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Cyprinodon</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FEFCFA8B7DCD5044" authorityName="Miller and Walters" authorityYear="1972" box="[357,460,1355,1379]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Megupsilon</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FE41FA8B7E385044" authorityName="Goode &amp; Bean" authorityYear="1879" box="[472,569,1355,1379]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Jordanella</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FDDDFA8B7E825044" authorityName="Miller" authorityYear="1956" box="[580,643,1355,1379]" family="Cyprinodontidae" genus="Cualac" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Cualac</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32FA8B780550B9" box="[683,1028,1355,1438]" gridcol="2" gridrow="9" pageId="6">21.9 (29.87–14.61)</td>
<td id="CF3EA04532050030FBB1FA8B79C150B9" box="[1064,1472,1355,1438]" gridcol="3" gridrow="9" pageId="6">27.8–16.0 Mya: Late Oligocene–Early Miocene Chicontepec Basin uplift (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18FA6779C150FA" box="[129,1472,1447,1501]" gridrow="10" pageId="6">
<th id="CF3EA04532050030FF18FA677CB950FA" box="[129,184,1447,1501]" gridcol="0" gridrow="10" pageId="6">10</th>
<td id="CF3EA04532050030FF69FA677E8250FA" box="[240,643,1447,1501]" gridcol="1" gridrow="10" pageId="6">
<emphasis id="00AB17693205FFC9FF69FA687E8150E7" box="[240,640,1448,1472]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FA687D9A50E7" authorityName="Goode &amp; Bean" authorityYear="1879" box="[240,411,1448,1472]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="species" species="floridae">Jordanella floridae</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FE3FFA687E0D50E7" authorityName="Miller and Walters" authorityYear="1972" box="[422,524,1448,1472]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Megupsilon</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FD8EFA687E8150E7" authorityName="Lacepede" authorityYear="1803" box="[535,640,1448,1472]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Cyprinodon</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32FA67780550FA" box="[683,1028,1447,1501]" gridcol="2" gridrow="10" pageId="6">16.8 (23.27–10.76)</td>
<td id="CF3EA04532050030FBB1FA6779C150FA" box="[1064,1472,1447,1501]" gridcol="3" gridrow="10" pageId="6">17.0–13.8 Mya: Middle Miocene Climatic Optimum (dispersal–vicariance cycle)</td>
</tr>
<tr id="8CEFC93932050030FF18FA2779C1533B" box="[129,1472,1511,1564]" gridrow="11" pageId="6">
<th id="CF3EA04532050030FF18FA277CB9533B" box="[129,184,1511,1564]" gridcol="0" gridrow="11" pageId="6">11</th>
<td id="CF3EA04532050030FF69FA277E82533B" box="[240,643,1511,1564]" gridcol="1" gridrow="11" pageId="6">
<emphasis id="00AB17693205FFC9FF69FA277DCF50D8" box="[240,462,1511,1535]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69FA277D5850D8" authorityName="Miller and Walters" authorityYear="1972" box="[240,345,1511,1535]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Megupsilon</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FEFAFA277DCF50D8" authorityName="Lacepede" authorityYear="1803" box="[355,462,1511,1535]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Cyprinodon</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32FA277805533B" box="[683,1028,1511,1564]" gridcol="2" gridrow="11" pageId="6">10.9 (14.91–7.37)</td>
<td id="CF3EA04532050030FBB1FA2779C1533B" box="[1064,1472,1511,1564]" gridcol="3" gridrow="11" pageId="6">11.8 Mya: development of Río San Fernando (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18F9E679C1535E" box="[129,1472,1574,1657]" gridrow="12" pageId="6">
<th id="CF3EA04532050030FF18F9E67CB9535E" box="[129,184,1574,1657]" gridcol="0" gridrow="12" pageId="6">12</th>
<td id="CF3EA04532050030FF69F9E67E82535E" box="[240,643,1574,1657]" gridcol="1" gridrow="12" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69F9E67D615319" box="[240,352,1574,1598]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FEF1F9E67DDC5319" authorityName="Myers" authorityYear="1924" box="[360,477,1574,1598]" family="Fundulidae" genus="Leptolucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">
<emphasis id="00AB17693205FFC9FEF1F9E67DDC5319" box="[360,477,1574,1598]" italics="true" pageId="6">Leptolucania</emphasis>
</taxonomicName>
</td>
<td id="CF3EA04532050030FD32F9E67805535E" box="[683,1028,1574,1657]" gridcol="2" gridrow="12" pageId="6">30.4 (39.35–22.47)</td>
<td id="CF3EA04532050030FBB1F9E679C1535E" box="[1064,1472,1574,1657]" gridcol="3" gridrow="12" pageId="6">30.0 Mya: Oligocene oxygen isotope zone 2 (Oi2) sea-level fall (dispersal–vicariance cycle)</td>
</tr>
<tr id="8CEFC93932050030FF18F94279C153F2" box="[129,1472,1666,1749]" gridrow="13" pageId="6">
<th id="CF3EA04532050030FF18F9427CB953F2" box="[129,184,1666,1749]" gridcol="0" gridrow="13" pageId="6">13</th>
<td id="CF3EA04532050030FF69F9427E8253F2" box="[240,643,1666,1749]" gridcol="1" gridrow="13" pageId="6">
<emphasis id="00AB17693205FFC9FF69F9427E1753BD" box="[240,534,1666,1690]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69F9427D4453BD" authorityName="Lacepede" authorityYear="1803" box="[240,325,1666,1690]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Fundulus</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FED6F9427DC153BD" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[335,448,1666,1690]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F83205FFC9FE55F9427E1753BD" authorityName="Girard" authorityYear="1860" box="[460,534,1666,1690]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Lucania</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32F942780553F2" box="[683,1028,1666,1749]" gridcol="2" gridrow="13" pageId="6">24.8 (31.31–19.12)</td>
<td id="CF3EA04532050030FBB1F94279C153F2" box="[1064,1472,1666,1749]" gridcol="3" gridrow="13" pageId="6">25.1 Mya: Oligocene oxygen isotope zone 2c (Oi2c) sea-level fall (dispersal–vicariance cycle)</td>
</tr>
<tr id="8CEFC93932050030FF18F91F79C15233" box="[129,1472,1759,1812]" gridrow="14" pageId="6">
<th id="CF3EA04532050030FF18F91F7CB95233" box="[129,184,1759,1812]" gridcol="0" gridrow="14" pageId="6">14</th>
<td id="CF3EA04532050030FF69F91F7E825233" box="[240,643,1759,1812]" gridcol="1" gridrow="14" pageId="6">
<emphasis id="00AB17693205FFC9FF69F91F7DB753D0" box="[240,438,1759,1783]" italics="true" pageId="6">
<taxonomicName id="F5DFB0F83205FFC9FF69F91F7D3B53D0" authorityName="Girard" authorityYear="1860" box="[240,314,1759,1783]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Lucania</taxonomicName>
/
<taxonomicName id="F5DFB0F83205FFC9FEDCF91F7DB753D0" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[325,438,1759,1783]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532050030FD32F91F78055233" box="[683,1028,1759,1812]" gridcol="2" gridrow="14" pageId="6">15.29 (30.72–7.47)</td>
<td id="CF3EA04532050030FBB1F91F79C15233" box="[1064,1472,1759,1812]" gridcol="3" gridrow="14" pageId="6">23.0 Mya: Basin and Range drainages separate from Río Grande (vicariance)</td>
</tr>
<tr id="8CEFC93932050030FF18F8DE79C15256" box="[129,1472,1822,1905]" gridrow="15" pageId="6">
<th id="CF3EA04532050030FF18F8DE7CB95256" box="[129,184,1822,1905]" gridcol="0" gridrow="15" pageId="6">15</th>
<td id="CF3EA04532050030FF69F8DE7E825256" box="[240,643,1822,1905]" gridcol="1" gridrow="15" pageId="6">
Crown
<taxonomicName id="F5DFB0F83205FFC9FEAEF8DE7D8B5211" authorityName="Lacepede" authorityYear="1803" box="[311,394,1822,1846]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" phylum="Chordata" rank="genus">
<emphasis id="00AB17693205FFC9FEAEF8DE7D8B5211" box="[311,394,1822,1846]" italics="true" pageId="6">Fundulus</emphasis>
</taxonomicName>
main split
</td>
<td id="CF3EA04532050030FD32F8DE78055256" box="[683,1028,1822,1905]" gridcol="2" gridrow="15" pageId="6">20.47 (24.24–17.11)</td>
<td id="CF3EA04532050030FBB1F8DE79C15256" box="[1064,1472,1822,1905]" gridcol="3" gridrow="15" pageId="6">20.8 Mya: Miocene oxygen isotope zone 1a (Mi1a) sea-level fall (dispersal–vicariance cycle)</td>
</tr>
</table>
</paragraph>
<caption id="66A09BF33204FFC8FFE8FF4F7D2A5580" box="[113,299,142,167]" isContinuationCaption="true" pageId="7" startId="7.[113,168,143,167]" targetBox="[113,1456,203,485]" targetIsTable="true" targetPageId="7" targetType="table">
<paragraph id="3260CB7B3204FFC8FFE8FF4F7D2A5580" blockId="7.[113,299,142,167]" box="[113,299,142,167]" pageId="7">
<emphasis id="00AB17693204FFC8FFE8FF4F7CBE5581" bold="true" box="[113,191,142,167]" pageId="7">Table 2.</emphasis>
Continued
</paragraph>
</caption>
<paragraph id="3260CB7B3204FFC8FFE8FF0B793A54C2" pageId="7">
<table id="40DF39DB32040030FFE8FF0B79B154C2" box="[113,1456,203,485]" colsContinueFrom="6.[129,1472,244,1905]" gridcols="4" gridrows="3" pageId="7">
<tr id="8CEFC93932040030FFE8FF0B79B15427" box="[113,1456,203,256]" gridrow="0" pageId="7">
<th id="CF3EA04532040030FFE8FF0B7CA95427" box="[113,168,203,256]" gridcol="0" gridrow="0" pageId="7">
<emphasis id="00AB17693204FFC8FFE8FF0B7CA955C4" bold="true" box="[113,168,203,227]" pageId="7">Node</emphasis>
</th>
<th id="CF3EA04532040030FF78FF0B7E5F5427" box="[225,606,203,256]" gridcol="1" gridrow="0" pageId="7">
<emphasis id="00AB17693204FFC8FF78FF0B7D9A55C4" bold="true" box="[225,411,203,227]" pageId="7">Divergence (node)</emphasis>
</th>
<th id="CF3EA04532040030FD05FF0B7FF45427" box="[668,1013,203,256]" gridcol="2" gridrow="0" pageId="7">
<emphasis id="00AB17693204FFC8FD05FF0B7FF45427" bold="true" pageId="7">Age estimate (Mya) (95% highest posterior density CI)</emphasis>
</th>
<th id="CF3EA04532040030FB80FF0B79B15427" box="[1049,1456,203,256]" gridcol="3" gridrow="0" pageId="7">
<emphasis id="00AB17693204FFC8FB80FF0B790455C4" bold="true" box="[1049,1285,203,227]" pageId="7">Reciprocal illumination</emphasis>
</th>
</tr>
<tr id="8CEFC93932040030FFE8FED879B15481" box="[113,1456,280,422]" gridrow="1" pageId="7">
<th id="CF3EA04532040030FFE8FED87CA95481" box="[113,168,280,422]" gridcol="0" gridrow="1" pageId="7">16</th>
<td id="CF3EA04532040030FF78FED87E5F5481" box="[225,606,280,422]" gridcol="1" gridrow="1" pageId="7">
<emphasis id="00AB17693204FFC8FF78FED87DE65469" italics="true" pageId="7">
<taxonomicName id="F5DFB0F83204FFC8FF78FED87D345417" authorityName="Lacepede" authorityYear="1803" box="[225,309,280,304]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">Fundulus</taxonomicName>
/
<taxonomicName id="F5DFB0F83204FFC8FEA6FED87DDB5417" baseAuthorityName="Walbaum" baseAuthorityYear="1792" box="[319,474,280,304]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="majalis">Fundulus majalis</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FE7CFED87D4D5469" authorityName="Miller" authorityYear="1955" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="persimilis">Fundulus persimilis</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FECEFEF67DE65469" baseAuthorityName="Baird &amp; Girard" baseAuthorityYear="1853" box="[343,487,310,334]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="similis">Fundulus similis</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532040030FD05FED87FF45481" box="[668,1013,280,422]" gridcol="2" gridrow="1" pageId="7">17.9 (20.98–15.15)</td>
<td id="CF3EA04532040030FB80FED879B15481" box="[1049,1456,280,422]" gridcol="3" gridrow="1" pageId="7">19.7–17.0 Mya: sea-level rise culminating in Middle Miocene Climatic Optimum (dispersal) 15.6 Mya: disjunct Guadalupe River delta arises (vicariance and ecological isolation)</td>
</tr>
<tr id="8CEFC93932040030FFE8FE7079B154C2" box="[113,1456,432,485]" gridrow="2" pageId="7">
<th id="CF3EA04532040030FFE8FE707CA954C2" box="[113,168,432,485]" gridcol="0" gridrow="2" pageId="7">17</th>
<td id="CF3EA04532040030FF78FE707E5F54C2" box="[225,606,432,485]" gridcol="1" gridrow="2" pageId="7">
<emphasis id="00AB17693204FFC8FF78FE707E5F54EF" box="[225,606,432,456]" italics="true" pageId="7">
<taxonomicName id="F5DFB0F83204FFC8FF78FE707D4254EF" authorityName="Agassiz" authorityYear="1854" box="[225,323,432,456]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">Zygonectes</taxonomicName>
/
<taxonomicName id="F5DFB0F83204FFC8FED4FE707DF454EF" authorityName="Cope" authorityYear="1865" box="[333,501,432,456]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="sciadicus">Fundulus sciadicus</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FD98FE707E5F54EF" authorityName="Garman" authorityYear="1895" box="[513,606,432,456]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">Plancterus</taxonomicName>
</emphasis>
</td>
<td id="CF3EA04532040030FD05FE707FF454C2" box="[668,1013,432,485]" gridcol="2" gridrow="2" pageId="7">16.6 (20.38–12.98)</td>
<td id="CF3EA04532040030FB80FE7079B154C2" box="[1049,1456,432,485]" gridcol="3" gridrow="2" pageId="7">17.0 Mya: nascent Great Plains ~68% open habitat (ecological isolation)</td>
</tr>
</table>
</paragraph>
<paragraph id="3260CB7B3204FFC8FE86FC577E4C5696" blockId="7.[113,762,919,1233]" box="[287,589,919,946]" pageId="7">
<emphasis id="00AB17693204FFC8FE86FC577E4C5696" bold="true" box="[287,589,919,946]" pageId="7">Diversification through time</emphasis>
</paragraph>
<paragraph id="3260CB7B3204FFC8FFE8FC7E7D2751F6" blockId="7.[113,762,919,1233]" pageId="7">
The lineage-through-time plots (
<figureCitation id="AAE4D7FE3204FFC8FE60FC7E7E4556F1" box="[505,580,958,982]" captionStart="Figure 4" captionStartId="8.[127,192,1459,1483]" captionTargetBox="[132,1470,147,1429]" captionTargetId="figure-233@8.[129,1473,144,1432]" captionTargetPageId="8" captionText="Figure 4. A, BAMM phylorate plot showing speciation rates along each branch within the North American killifish families Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae. Cool colours represent slow rates; warm colours represent fast rates.B–E, speciation rate over time is shown graphically for Fundulidae (B), Cyprinodontidae (C), Profundulidae (D), and Goodeidae (E). λ = mean diversification rate." pageId="7">Fig. 4</figureCitation>
) indicate that
<taxonomicName id="F5DFB0F83204FFC8FFE8FC1D7D2556D2" box="[113,292,989,1013]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
is the only family that experienced a diversification-rate shift, with acceleration initiating ~10.9 Mya and continuing to the present (λ = 0.23 mean; range = 0.14– 0.38). The initial rate increase occurred when
<taxonomicName id="F5DFB0F83204FFC8FDFDFBFB7ED45174" authorityName="Miller and Walters" authorityYear="1972" box="[612,725,1083,1107]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FDFDFBFB7ED45174" box="[612,725,1083,1107]" italics="true" pageId="7">Megupsilon</emphasis>
</taxonomicName>
diverged from
<taxonomicName id="F5DFB0F83204FFC8FF6AFB9B7D695154" authorityName="Lacepede" authorityYear="1803" box="[243,360,1115,1139]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FF6AFB9B7D695154" box="[243,360,1115,1139]" italics="true" pageId="7">Cyprinodon</emphasis>
</taxonomicName>
. No shifts were detected in
<taxonomicName id="F5DFB0F83204FFC8FD1DFB9B7EFB5154" box="[644,762,1115,1139]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
(λ =.12 mean; range =.07–.21),
<taxonomicName id="F5DFB0F83204FFC8FD9BFBBA7E7451B5" box="[514,629,1146,1170]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
(λ = mean.13; range =.09–.29), or
<taxonomicName id="F5DFB0F83204FFC8FE3CFB597E385196" box="[421,569,1177,1201]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
(λ = mean.12; range =.07–.24).
</paragraph>
<paragraph id="3260CB7B3204FFC8FE9EFB397E645034" blockId="7.[112,763,1273,1962]" box="[263,613,1273,1299]" pageId="7">
<emphasis id="00AB17693204FFC8FE9EFB397E645034" bold="true" box="[263,613,1273,1299]" pageId="7">Reconstruction of ancestral areas</emphasis>
</paragraph>
<paragraph id="3260CB7B3204FFC8FFE8FADF7ECD5334" blockId="7.[112,763,1273,1962]" pageId="7">
The best-fitting model for reconstruction of ancestral areas was the BAYAREALIKE+J (Supporting Information,
<tableCitation id="7F5DFEC03204FFC8FD10FAFF7EE85070" box="[649,745,1343,1367]" captionStart="Table 3" captionStartId="17.[113,168,143,167]" captionTargetPageId="17" captionText="Table 3. Early-diverging lineages with peripheral geography and relatively limited diversification represent phylogenetic evidence for diversification by barrier displacement (Albert et al. 2018). Examples from Cyprinodontidae, Fundulidae, and Goodeidae are provided here. Sister clades are provided for comparison with each peripheral lineage.The chronogram node (Fig. 3) is given for each sister relationship" pageId="7">Table S3</tableCitation>
), consistent with the coarse level of analysis, which recognized only the Nearctic and Neotropical realms. Most divergence events occurred within one realm or the other, as required by the BAYAREALIKE model (Garcia-R and
<bibRefCitation id="564EB68A3204FFC8FE67FA7C7E8550F3" author="Matzke NJ" box="[510,644,1468,1492]" pageId="7" refId="ref16531" refString="Garcia-R JC, Matzke NJ. Trait-dependent dispersal in rails (Aves: Rallidae): historical biogeography of a cosmopolitan bird clade. Molecular Phylogenetics and Evolution 2021; 159: 107106. https: // doi. org / 10.1016 / j. ympev. 2021.107106" type="journal volume" year="2021">Matzke 2021</bibRefCitation>
). Inclusion of the +J portion of the model implies that immigrations between realms occurred via long-distance dispersal (
<bibRefCitation id="564EB68A3204FFC8FDAFFA3B7EBA5334" author="Matzke NJ" box="[566,699,1531,1555]" pageId="7" pagination="951 - 70" refId="ref18102" refString="Matzke NJ. Model selection in historical biogeography reveals that founder-event speciation is a crucial process in island clades. Systematic Biology 2014; 63: 951 - 70. https: // doi. org / 10.1093 / sysbio / syu 056" type="journal article" year="2014">Matzke 2014</bibRefCitation>
).
</paragraph>
<paragraph id="3260CB7B3204FFC8FF14F9DA78E757FB" blockId="7.[112,763,1273,1962]" lastBlockId="7.[809,1460,582,732]" pageId="7">
The ancestral distribution for this clade of North American killifishes is optimized as Nearctic–Neotropical, suggesting a widespread most recent common ancestor (MRCA) (
<figureCitation id="AAE4D7FE3204FFC8FD37F9997EE85356" box="[686,745,1625,1649]" captionStart="Figure 5" captionStartId="10.[129,194,1836,1860]" captionTargetBox="[212,1390,147,1805]" captionTargetId="figure-7@10.[209,1393,144,1808]" captionTargetPageId="10" captionText="Figure 5. Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (Rico et al.2022). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS." pageId="7">Fig. 5</figureCitation>
). This clade first split into a northern fork with primarily Nearctic affinities (
<taxonomicName id="F5DFB0F83204FFC8FF7EF9577D615388" box="[231,352,1687,1711]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FEF4F9577E235388" box="[365,546,1687,1711]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
) and a southern fork with Nearctic–Neotropical affiliation (Cubanichthyidae,
<taxonomicName id="F5DFB0F83204FFC8FFE8F9167D0453C9" box="[113,261,1750,1774]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
, and
<taxonomicName id="F5DFB0F83204FFC8FEDEF9167DBF53C9" box="[327,446,1750,1774]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
). Appearance of fundulids in the Neotropics is relatively recent (Late Neogene–Quaternary), limited to the
<taxonomicName id="F5DFB0F83204FFC8FE9EF8D57EAB520A" authority="Baird and Girard, 1853" authorityName="Baird and Girard" authorityYear="1853" box="[263,682,1813,1837]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="grandis">
<emphasis id="00AB17693204FFC8FE9EF8D57DB1520A" box="[263,432,1813,1837]" italics="true" pageId="7">Fundulus grandis</emphasis>
Baird and Girard, 1853
</taxonomicName>
species group. Nearctic
<taxonomicName id="F5DFB0F83204FFC8FEBFF8F47DD9526B" box="[294,472,1844,1868]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
made several Neotropical invasions (
<taxonomicName id="F5DFB0F83204FFC8FF6DF8937D73524C" authorityName="Hubbs" authorityYear="1926" box="[244,370,1875,1899]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FF6DF8937D73524C" box="[244,370,1875,1899]" italics="true" pageId="7">Floridichthys</emphasis>
</taxonomicName>
,
<collectingRegion id="F01B05993204FFC8FE08F8947DE3524B" box="[401,482,1876,1900]" country="Mexico" name="Yucatan" pageId="7">Yucatán</collectingRegion>
<taxonomicName id="F5DFB0F83204FFC8FE65F8937EFB524C" authority=", Caribbean" authorityName="Caribbean" box="[508,762,1875,1899]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FE65F8937E71524C" box="[508,624,1875,1899]" italics="true" pageId="7">Cyprinodon</emphasis>
, Caribbean
</taxonomicName>
<taxonomicName id="F5DFB0F83204FFC8FFE8F8B37CE452AC" box="[113,229,1907,1931]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FFE8F8B37CE452AC" box="[113,229,1907,1931]" italics="true" pageId="7">Cyprinodon</emphasis>
</taxonomicName>
, and
<taxonomicName id="F5DFB0F83204FFC8FEB8F8B37EAB52AC" authority="Lacepede, 1803" authorityName="Lacepede" authorityYear="1803" box="[289,682,1907,1931]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="variegatus">
<emphasis id="00AB17693204FFC8FEB8F8B37E0152AC" box="[289,512,1907,1931]" italics="true" pageId="7">Cyprinodon variegatus</emphasis>
Lacépède, 1803
</taxonomicName>
species group). The MRCAs of Cubanichthyidae and
<taxonomicName id="F5DFB0F83204FFC8FDFEF8527EFB528D" box="[615,762,1938,1962]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
were Neotropical, and both families are restricted to that realm.
<taxonomicName id="F5DFB0F83204FFC8FCB2FDA57F9C575A" box="[811,925,613,637]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
is most likely to have had a Nearctic origin, with representatives of
<taxonomicName id="F5DFB0F83204FFC8FC5AFD45785857BA" authorityName="Bleeker" authorityYear="1860" box="[963,1113,645,669]" family="Goodeidae" genus="Girardinichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FC5AFD45785857BA" box="[963,1113,645,669]" italics="true" pageId="7">Girardinichthys</emphasis>
</taxonomicName>
,
<taxonomicName id="F5DFB0F83204FFC8FBFCFD457F60579B" authority="Hubbs &amp; Turner, 1939" authorityName="Hubbs &amp; Turner" authorityYear="1939" family="Goodeidae" genus="Allodontichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FBFCFD4578FD57BA" box="[1125,1276,645,669]" italics="true" pageId="7">Allodontichthys</emphasis>
Hubbs &amp; Turner, 1939
</taxonomicName>
,
<taxonomicName id="F5DFB0F83204FFC8FCF1FD65785E579B" authority="Turner, 1946" authorityName="Turner" authorityYear="1946" box="[872,1119,676,700]" family="Goodeidae" genus="Xenotaenia" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FCF1FD657FD7579B" box="[872,982,677,700]" italics="true" pageId="7">Xenotaenia</emphasis>
Turner, 1946
</taxonomicName>
, and
<taxonomicName id="F5DFB0F83204FFC8FB16FD6478D9579B" authorityName="Eigenmann" authorityYear="1907" box="[1167,1240,676,700]" family="Goodeidae" genus="Ilyodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FB16FD6478D9579B" box="[1167,1240,676,700]" italics="true" pageId="7">Ilyodon</emphasis>
</taxonomicName>
reaching Neotropical drainages in the Late Neogene–Quaternary.
</paragraph>
<paragraph id="3260CB7B3204FFC8FC28FD3B7FE05008" blockId="7.[809,1460,763,1327]" pageId="7">
<emphasis id="00AB17693204FFC8FC28FD3B792D5632" bold="true" box="[945,1324,763,789]" pageId="7">Reconstruction of ancestral habitats</emphasis>
Ancestral-habitats reconstruction (
<figureCitation id="AAE4D7FE3204FFC8FB00FCE178E5561D" box="[1177,1252,801,826]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="7">Fig. 2B</figureCitation>
) indicates that the MRCA of North American killifishes was coastal. Inland invasions within
<taxonomicName id="F5DFB0F83204FFC8FC4BFCA07846565F" box="[978,1095,864,888]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
include
<taxonomicName id="F5DFB0F83204FFC8FB34FCA17FCB56B0" authority="Hubbs &amp; Miller, 1965" authorityName="Hubbs &amp; Miller" authorityYear="1965" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="interioris">
<emphasis id="00AB17693204FFC8FB34FCA17961565F" box="[1197,1376,865,888]" italics="true" pageId="7">Lucania interioris</emphasis>
Hubbs &amp; Miller, 1965
</taxonomicName>
, the
<taxonomicName id="F5DFB0F83204FFC8FB9BFCBF794156B0" authority="Cope, 1865" authorityName="Cope" authorityYear="1865" box="[1026,1344,895,920]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="species" species="sciadicus">
<emphasis id="00AB17693204FFC8FB9BFCBF78BA56B0" box="[1026,1211,895,919]" italics="true" pageId="7">Fundulus sciadicus</emphasis>
Cope, 1865
</taxonomicName>
<emphasis id="00AB17693204FFC8FAD9FCBF794C56B0" box="[1344,1357,895,919]" italics="true" pageId="7">–</emphasis>
<taxonomicName id="F5DFB0F83204FFC8FAD4FCBF7FC15690" authority="Garman, 1895" authorityName="Garman" authorityYear="1895" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FAD4FCBF79B256B0" box="[1357,1459,895,919]" italics="true" pageId="7">Plancterus</emphasis>
Garman, 1895
</taxonomicName>
group, and later-branching lineages within subgenera
<taxonomicName id="F5DFB0F83204FFC8FCEFFC7E787B56F1" authority="Lacepede, 1803" authorityName="Lacepede" authorityYear="1803" box="[886,1146,958,982]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FCEFFC7E7FD156F1" box="[886,976,958,982]" italics="true" pageId="7">Fundulus</emphasis>
Lacépède, 1803
</taxonomicName>
and
<taxonomicName id="F5DFB0F83204FFC8FB29FC7F79AE56F1" authority="Agassiz, 1854" authorityName="Agassiz" authorityYear="1854" box="[1200,1455,958,982]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FB29FC7F791856F1" box="[1200,1305,959,982]" italics="true" pageId="7">Zygonectes</emphasis>
Agassiz, 1854
</taxonomicName>
.
<taxonomicName id="F5DFB0F83204FFC8FCB3FC1D7F9E56D2" box="[810,927,989,1013]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
are the only family with species broadly distributed between coastal and upland habitats. In
<taxonomicName id="F5DFB0F83204FFC8FB65FC3D79AE5132" box="[1276,1455,1021,1045]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83204FFC8FCB5FBDC7FAB5113" authorityName="Hubbs" authorityYear="1926" box="[812,938,1052,1076]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FCB5FBDC7FAB5113" box="[812,938,1052,1076]" italics="true" pageId="7">Floridichthys</emphasis>
</taxonomicName>
and
<taxonomicName id="F5DFB0F83204FFC8FC4CFBDC79105113" authority="Goode &amp; Bean, 1879" authorityName="Goode &amp; Bean" authorityYear="1879" box="[981,1297,1052,1076]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FC4CFBDC78395113" box="[981,1080,1052,1076]" italics="true" pageId="7">Jordanella</emphasis>
Goode &amp; Bean, 1879
</taxonomicName>
retained coastal affinities, while the maritime branch of
<taxonomicName id="F5DFB0F83204FFC8FB5CFBFB79385174" box="[1221,1337,1083,1107]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FB5CFBFB79385174" box="[1221,1337,1083,1107]" italics="true" pageId="7">Cyprinodon</emphasis>
</taxonomicName>
gave rise to upland and coastal lineages. Other branches within
<taxonomicName id="F5DFB0F83204FFC8FAD9FB9B79B55154" box="[1344,1460,1115,1139]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="genus">
<emphasis id="00AB17693204FFC8FAD9FB9B79B55154" box="[1344,1460,1115,1139]" italics="true" pageId="7">Cyprinodon</emphasis>
</taxonomicName>
diverged to become upland lineages. Cubanichthyidae remained coastal, whereas
<taxonomicName id="F5DFB0F83204FFC8FC4BFB5978645196" box="[978,1125,1177,1201]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
and
<taxonomicName id="F5DFB0F83204FFC8FB0DFB5979075196" box="[1172,1286,1177,1201]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
are likely to have originated and diversified entirely in uplands. Overall, 58.6% of species had coastal distributions, 33.5% had upland distributions, and 7.9% had distributions spanning coastal and upland habitats (
<figureCitation id="AAE4D7FE3204FFC8FC13FAD77FD15008" box="[906,976,1303,1327]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="7">Fig. 2B</figureCitation>
).
</paragraph>
<paragraph id="3260CB7B3204FFC8FC7DFA8E78F8504F" blockId="7.[996,1273,1358,1384]" box="[996,1273,1358,1384]" pageId="7">
<emphasis id="00AB17693204FFC8FC7DFA8E78F8504F" bold="true" box="[996,1273,1358,1384]" pageId="7">Biogeographical synthesis</emphasis>
</paragraph>
<paragraph id="3260CB7B3204FFC8FC4DFAB8790850B5" blockId="7.[980,1289,1400,1426]" box="[980,1289,1400,1426]" pageId="7">
<emphasis id="00AB17693204FFC8FC4DFAB8790850B5" box="[980,1289,1400,1426]" italics="true" pageId="7">Early branching of major lineages</emphasis>
</paragraph>
<paragraph id="3260CB7B3204FFC8FCB3FA5F791353F6" blockId="7.[809,1462,1439,1964]" pageId="7">
<collectingCounty id="DB01B3F73204FFC8FCB3FA5F7F895090" box="[810,904,1439,1463]" pageId="7">Evidence</collectingCounty>
suggests that the MRCA of
<collectingMunicipality id="D20451013204FFC8FB3DFA5F794C5090" box="[1188,1357,1439,1463]" pageId="7">North American</collectingMunicipality>
killifishes arose in the
<location id="37009DA03204FFC8FC39FA7E785650F1" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FC39FA7E785650F1" box="[928,1111,1470,1494]" county="Evidence" latitude="3.0" municipality="North American" name="Early Eocene Gulf" pageId="7" stateProvince="Mexico">Early Eocene Gulf</location>
of
<collectingRegion id="F01B05993204FFC8FBEAFA7E78C150F1" box="[1139,1216,1470,1494]" country="Mexico" name="Mexico" pageId="7">México</collectingRegion>
(
<figureCitation id="AAE4D7FE3204FFC8FB57FA7E790950F1" box="[1230,1288,1470,1494]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="7">Figs 3</figureCitation>
,
<figureCitation id="AAE4D7FE3204FFC8FA88FA7E791F50F1" box="[1297,1310,1470,1494]" captionStart="Figure 4" captionStartId="8.[127,192,1459,1483]" captionTargetBox="[132,1470,147,1429]" captionTargetId="figure-233@8.[129,1473,144,1432]" captionTargetPageId="8" captionText="Figure 4. A, BAMM phylorate plot showing speciation rates along each branch within the North American killifish families Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae. Cool colours represent slow rates; warm colours represent fast rates.B–E, speciation rate over time is shown graphically for Fundulidae (B), Cyprinodontidae (C), Profundulidae (D), and Goodeidae (E). λ = mean diversification rate." pageId="7">4</figureCitation>
).
<location id="37009DA03204FFC8FAAAFA7E79AE50F1" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FAAAFA7E79AE50F1" box="[1331,1455,1470,1494]" county="Evidence" latitude="3.0" municipality="North American" name="Accordingly" pageId="7" stateProvince="Mexico">Accordingly</location>
, the MRCA is likely to have had high salinity tolerance (
<bibRefCitation id="564EB68A3204FFC8FACFFA1D7FD65332" author="Ghedotti MJ &amp; Davis MP" pageId="7" pagination="1 - 65" refId="ref16579" refString="Ghedotti MJ, Davis MP. Phylogeny, classification, and evolution of salinity tolerance of the North American topminnows and killifishes, family Fundulidae (Teleostei: Cyprinodontiformes). Fieldiana Life and Earth Sciences 2013; 7: 1 - 65. https: // doi. org / 10.3158 / 2158 - 5520 - 12.7.1" type="journal article" year="2013">Ghedotti and Davis 2013</bibRefCitation>
).
<location id="37009DA03204FFC8FC6DFA3D78215332" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FC6DFA3D78215332" box="[1012,1056,1533,1557]" county="Evidence" latitude="3.0" municipality="North American" name="This" pageId="7" stateProvince="Mexico">This</location>
timing corresponds precisely with a period of increased taxonomic and morphological diversification (
<bibRefCitation id="564EB68A3204FFC8FCFFF9FB785A5373" author="Ghezelayagh A &amp; Harrington RC &amp; Burress ED" box="[870,1115,1595,1620]" pageId="7" pagination="1211 - 20" refId="ref16688" refString="Ghezelayagh A, Harrington RC, Burress ED et al. Prolonged morphological expansion of spiny-rayed fishes following the end- Cretaceous. Nature Ecology &amp; Evolution 2022; 6: 1211 - 20. https: // doi. org / 10.1038 / s 41559 - 022 - 01801 - 3" type="journal article" year="2022">
Ghezelayagh
<emphasis id="00AB17693204FFC8FC69F9FC78205374" box="[1008,1057,1595,1619]" italics="true" pageId="7">et al.</emphasis>
2022
</bibRefCitation>
).
<location id="37009DA03204FFC8FBEAF9FB789B5374" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FBEAF9FB789B5374" box="[1139,1178,1595,1619]" county="Evidence" latitude="3.0" municipality="North American" name="The" pageId="7" stateProvince="Mexico">The</location>
distribution of the MRCA potentially spanned the present-day
<location id="37009DA03204FFC8FB35F99B79025354" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FB35F99B79025354" box="[1196,1283,1627,1651]" county="Evidence" latitude="3.0" municipality="North American" name="Nearctic" pageId="7" stateProvince="Mexico">Nearctic</location>
and
<location id="37009DA03204FFC8FAA2F99B79B75354" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FAA2F99B79B75354" box="[1339,1462,1627,1651]" county="Evidence" latitude="3.0" municipality="North American" name="Neotropical" pageId="7" stateProvince="Mexico">Neotropical</location>
realms (
<figureCitation id="AAE4D7FE3204FFC8FCE6F9BA7FB953B5" box="[895,952,1658,1682]" captionStart="Figure 5" captionStartId="10.[129,194,1836,1860]" captionTargetBox="[212,1390,147,1805]" captionTargetId="figure-7@10.[209,1393,144,1808]" captionTargetPageId="10" captionText="Figure 5. Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (Rico et al.2022). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS." pageId="7">Fig. 5</figureCitation>
).
<location id="37009DA03204FFC8FC49F9BB782F53B5" LSID="urn:lsid:plazi:treatment:BA767A6D3207FFC6FBD9FEBB78A85484:37009DA03204FFC8FC49F9BB782F53B5" box="[976,1070,1659,1682]" county="Evidence" latitude="3.0" municipality="North American" name="However" pageId="7" stateProvince="Mexico">However</location>
, it was likely to be tropical because at this time; the tropics extended to
<geoCoordinate id="57EBADBC3204FFC8FB1CF95A78BD5395" box="[1157,1212,1690,1714]" degrees="27" direction="north" orientation="latitude" pageId="7" precision="55555" value="27.0">27°N</geoCoordinate>
,
<geoCoordinate id="57EBADBC3204FFC8FB5FF95A791B5396" box="[1222,1306,1689,1714]" degrees="3" direction="north" orientation="latitude" pageId="7" precision="55555" value="3.0">3° north</geoCoordinate>
of the present-day boundary of the tropics (
<bibRefCitation id="564EB68A3204FFC8FBCAF979790053F6" author="Zhang L &amp; Hay WW &amp; Wang C" box="[1107,1281,1721,1745]" pageId="7" pagination="147 - 58" refId="ref21386" refString="Zhang L, Hay WW, Wang C et al. The evolution of latitudinal temperature gradients from the latest Cretaceous through the Present. Earth-Science Reviews 2019; 189: 147 - 58. https: // doi. org / 10.1016 / j. earscirev. 2019.01.025" type="journal article" year="2019">
Zhang
<emphasis id="00AB17693204FFC8FB03F97A78C853F6" box="[1178,1225,1721,1745]" italics="true" pageId="7">et al.</emphasis>
2019
</bibRefCitation>
).
</paragraph>
<paragraph id="3260CB7B3204FFC7FCDCF9187984538E" blockId="7.[809,1462,1439,1964]" lastBlockId="8.[823,1476,1587,1956]" lastPageId="8" pageId="7">
The primary phylogenetic split in North American killifishes separated the northern sister families
<taxonomicName id="F5DFB0F83204FFC8FCB3F8D77FDB5208" box="[810,986,1815,1839]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FC71F8D7785F5208" box="[1000,1118,1815,1839]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
from a southern subclade of Cubanichthyidae–
<taxonomicName id="F5DFB0F83204FFC8FC7EF8F678565269" box="[999,1111,1846,1870]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83204FFC8FBFDF8F678F85269" box="[1124,1273,1846,1870]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
58.6–45.8 Mya (
<figureCitation id="AAE4D7FE3204FFC8FCACF8957F735249" box="[821,882,1877,1902]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="7">Fig. 3</figureCitation>
, node 1;
<tableCitation id="7F5DFEC03204FFC8FC46F895782E5249" box="[991,1071,1877,1902]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="7">Table 2</tableCitation>
). The relative distributions of these families (
<figureCitation id="AAE4D7FE3204FFC8FC16F8B57FCC52AA" box="[911,973,1909,1933]" captionStart="Figure 1" captionStartId="1.[113,178,1926,1950]" captionTargetBox="[116,1455,809,1895]" captionTargetId="figure-368@1.[114,1458,806,1898]" captionTargetPageId="1" captionText="Figure 1. Map depicting the distribution of: A, North American killifish families; B, Fundulidae; C, Cyprinodontidae; D, Profundulidae; and E, Goodeidae." pageId="7">Fig. 1</figureCitation>
), our ancestral-areas reconstruction (
<figureCitation id="AAE4D7FE3204FFC8FAFDF8B579A352AA" box="[1380,1442,1909,1933]" captionStart="Figure 5" captionStartId="10.[129,194,1836,1860]" captionTargetBox="[212,1390,147,1805]" captionTargetId="figure-7@10.[209,1393,144,1808]" captionTargetPageId="10" captionText="Figure 5. Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (Rico et al.2022). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS." pageId="7">Fig. 5</figureCitation>
), and our ancestral-habitats reconstruction (
<figureCitation id="AAE4D7FE3204FFC8FB66F8547941528B" box="[1279,1344,1940,1964]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="7">Fig. 2</figureCitation>
) combine to suggest that separation occurred as a north–south division within the Gulf of
<collectingRegion id="F01B0599320BFFC7FEDBF9937D91534C" box="[322,400,1619,1643]" country="Mexico" name="Mexico" pageId="8">México</collectingRegion>
. The Tropic of Cancer was gradually retracting southwards through this period (
<bibRefCitation id="564EB68A320BFFC7FDDDF9B27EF853AD" author="Zhang L &amp; Hay WW &amp; Wang C" box="[580,761,1650,1674]" pageId="8" pagination="147 - 58" refId="ref21386" refString="Zhang L, Hay WW, Wang C et al. The evolution of latitudinal temperature gradients from the latest Cretaceous through the Present. Earth-Science Reviews 2019; 189: 147 - 58. https: // doi. org / 10.1016 / j. earscirev. 2019.01.025" type="journal article" year="2019">
Zhang
<emphasis id="00AB1769320BFFC7FD14F9B37EBF53AD" box="[653,702,1650,1674]" italics="true" pageId="8">et al.</emphasis>
2019
</bibRefCitation>
), which could imply that distinct subtropical ecosystems developed in the northern Gulf. We hypothesize that the northern MRCA of
<taxonomicName id="F5DFB0F8320BFFC7FF65F9107D7353CF" box="[252,370,1744,1768]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
–
<taxonomicName id="F5DFB0F8320BFFC7FE19F9107E3053CF" box="[384,561,1744,1768]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
adapted to inhabit subtropical estuaries, including those of the
<collectingRegion id="F01B0599320BFFC7FDDBF92F7EB75220" box="[578,694,1775,1799]" country="United States of America" name="Mississippi" pageId="8">Mississippi</collectingRegion>
, Brazos, and Río Grande (
<figureCitation id="AAE4D7FE320BFFC7FED8F8CF7D7C5200" box="[321,381,1807,1831]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="8">Fig. 6</figureCitation>
). Concurrently, the southern MRCA of Cubanichthyidae–
<taxonomicName id="F5DFB0F8320BFFC7FEF7F8EE7DDF5261" box="[366,478,1838,1862]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
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potentially occupied a tropical estuary in the southwestern Gulf (Bejuco, Chicontepec, and Nautla) (
<figureCitation id="AAE4D7FE320BFFC7FE31F8AD7DE452A2" box="[424,485,1901,1925]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="8">Fig. 6</figureCitation>
). Geological evidence suggests the Gulf of
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became isolated from the world ocean 55.8–55.0 Mya, accompanied by sea-level recession of
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(
<bibRefCitation id="564EB68A320BFFC7FC0FF993784A534C" author="Cossey SPJ &amp; Van Nieuwenhuise D &amp; Davis J" box="[918,1099,1619,1643]" pageId="8" pagination="63 - 80" refId="ref15238" refString="Cossey SPJ, Van Nieuwenhuise D, Davis J et al. Compelling evidence from eastern Mexico for a Late Paleocene / Early Eocene isolation, drawdown, and refill of the Gulf of Mexico. Interpretation 2016; 4: SC 63 - 80. https: // doi. org / 10.1190 / int- 2015 - 0107.1" type="journal article" year="2016">
Cossey
<emphasis id="00AB1769320BFFC7FC7CF9947812534C" box="[997,1043,1619,1643]" italics="true" pageId="8">et al.</emphasis>
2016
</bibRefCitation>
,
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). Because sea-level fall isolates estuaries (
<bibRefCitation id="564EB68A320BFFC7FC3BF9B2784C53AD" author="Dolby GA &amp; Bedolla AM &amp; Bennett SEK" box="[930,1101,1650,1674]" pageId="8" refId="ref15502" refString="Dolby GA, Bedolla AM, Bennett SEK et al. Global physical controls on estuarine habitat distribution during sea level change: consequences for genetic diversification through time. Global and Planetary Change 2020; 187: 103128. https: // doi. org / 10.1016 / j. gloplacha. 2020.103128" type="journal volume" year="2020">
Dolby
<emphasis id="00AB1769320BFFC7FC7EF9B3781453AD" box="[999,1045,1650,1674]" italics="true" pageId="8">et al.</emphasis>
2020
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), this extreme drawdown could have caused strong separation of northern and southern clades.
</paragraph>
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<paragraph id="3260CB7B320BFFC7FFE6FA7379685324" blockId="8.[127,1472,1459,1540]" pageId="8">
<emphasis id="00AB1769320BFFC7FFE6FA737CD650EC" bold="true" box="[127,215,1459,1483]" pageId="8">Figure 4.</emphasis>
A, BAMM phylorate plot showing speciation rates along each branch within the North American killifish families
<taxonomicName id="F5DFB0F8320BFFC7FA93FA7479B150EB" box="[1290,1456,1460,1484]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
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,
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, and
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. Cool colours represent slow rates; warm colours represent fast rates. B–E, speciation rate over time is shown graphically for
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(B),
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(C),
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(D), and
<taxonomicName id="F5DFB0F8320BFFC7FC25FA2C78245323" box="[956,1061,1516,1540]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
(E). λ = mean diversification rate.
</paragraph>
</caption>
<paragraph id="3260CB7B320BFFC6FCCCF9717CD5563C" blockId="8.[823,1476,1587,1956]" lastBlockId="9.[112,764,144,1954]" lastPageId="9" pageId="8">
We hypothesize divergence of
<taxonomicName id="F5DFB0F8320BFFC7FB50F971797A53EE" box="[1225,1403,1713,1737]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
from
<taxonomicName id="F5DFB0F8320BFFC7FCA0F9107FAE53CF" box="[825,943,1744,1768]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
, 54.8–37.6 Mya (
<figureCitation id="AAE4D7FE320BFFC7FBF4F91078A853CF" box="[1133,1193,1744,1768]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="8">Fig. 3</figureCitation>
, node 2;
<tableCitation id="7F5DFEC0320BFFC7FA8AF910796053CF" box="[1299,1377,1744,1768]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="8">Table 2</tableCitation>
) was via east–west vicariance along the northern Gulf Coast. Drainage reconfiguration occurred in the northern Gulf of
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48.2 Mya (Early–Middle Eocene transition), when reduced sediment supply caused alluvial deltas to retract and separate (
<bibRefCitation id="564EB68A320BFFC7FAA7F88D7FCE52A2" author="Snedden JW &amp; Galloway WE" pageId="8" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
). Shifting delta locations (
<bibRefCitation id="564EB68A320BFFC7FB43F8AD7F6D5283" author="Snedden JW &amp; Galloway WE" pageId="8" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
) might have initially dispersed and then subdivided killifish populations. To the northeast, the MRCA of
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was potentially isolated in the Middle Eocene Brazos River delta. From there, it is likely to have ranged east into the adjacent
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Embayment, which flooded at this time (
<figureCitation id="AAE4D7FE320AFFC6FDB0FF2E7E645421" box="[553,613,238,262]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="9">Fig. 6</figureCitation>
). The embayment refilled gradually through the Middle Eocene (
<bibRefCitation id="564EB68A320AFFC6FD39FECD7D3B5462" author="Snedden JW &amp; Galloway WE" pageId="9" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
), suggesting that it contained extensive estuarine habitats that could have been the cradle for
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. To the southwest, the MRCA of
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putatively originated in a separate delta region associated with the Río Grande and Río Bravo deltas (
<figureCitation id="AAE4D7FE320AFFC6FECEFE6A7D9154E5" box="[343,400,426,450]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="9">Fig. 6</figureCitation>
) that were comparatively sediment rich and wave swept (
<bibRefCitation id="564EB68A320AFFC6FED1FE097E6954C6" author="Snedden JW &amp; Galloway WE" box="[328,616,457,481]" pageId="9" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
). Such habitat is seemingly well suited for the ancestor of this group, which is hypothesized to have resembled
<taxonomicName id="F5DFB0F8320AFFC6FE5BFDC87D295718" authority="(Echelle &amp; Echelle, 2020)" baseAuthorityName="Echelle &amp; Echelle" baseAuthorityYear="2020" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="species" species="variegatus">
<emphasis id="00AB1769320AFFC6FE5BFDC87E9E5707" box="[450,671,520,544]" italics="true" pageId="9">Cyprinodon variegatus</emphasis>
(
<bibRefCitation id="564EB68A320AFFC6FD32FDC87D215718" author="Echelle AA &amp; Echelle AF" pageId="9" pagination="609 - 73" refId="ref15900" refString="Echelle AA, Echelle AF. Cyprinodontidae: Pupfishes. In: Warren ML Jr, Burr BM (eds), Freshwater fishes of North America, Vol. 2, Characidae to Poeciliidae. Baltimore: Johns Hopkins University Press, 2020, 609 - 73." type="book chapter" year="2020">Echelle &amp; Echelle, 2020</bibRefCitation>
)
</taxonomicName>
, a species well adapted for open coasts with fluctuating temperatures and salinities (
<bibRefCitation id="564EB68A320AFFC6FD82FD877CA45759" author="Simpson DG &amp; Gunter G" pageId="9" pagination="115 - 34" refId="ref20219" refString="Simpson DG, Gunter G. Notes on habitats, systematic characters and life histories of Texas salt water cyprinodonts. Tulane Studies in Zoology 1956; 116: 115 - 34." type="journal article" year="1956">Simpson and Gunter 1956</bibRefCitation>
, Martin 1968,
<bibRefCitation id="564EB68A320AFFC6FED1FDA67DD55759" author="Nordlie FG" box="[328,468,614,638]" pageId="9" pagination="51 - 106" refId="ref18725" refString="Nordlie FG. Physicochemical environments and tolerances of cyprinodontoid fishes found in estuaries and salt marshes of eastern North America. Reviews in Fish Biology and Fisheries 2006; 16: 51 - 106. https: // doi. org / 10.1007 / s 11160 - 006 - 9003 - 0" type="journal article" year="2006">Nordlie 2006</bibRefCitation>
). This period also included ~
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sea-level fall between 48.0 and 47.5 Mya (
<bibRefCitation id="564EB68A320AFFC6FD18FD457CB0579A" author="Miller KG &amp; Browning JV &amp; Schmelz WJ" pageId="9" refId="ref18255" refString="Miller KG, Browning JV, Schmelz WJ et al. Cenozoic sea-level and cryospheric evolution from deep-sea geochemical and continental margin records. Science Advances 2020 a; 6: eaaz 1346. https: // doi. org / 10.1126 / sciadv. aaz 1346" type="journal volume" year="2020">
Miller
<emphasis id="00AB1769320AFFC6FD50FD467EFA57BA" box="[713,763,645,669]" italics="true" pageId="9">et al.</emphasis>
2020a
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FF24FD657CF0579A" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[189,241,677,701]" pageId="9" refId="ref18344" refString="Miller KG, Schmelz WJ, Browning JV et al. Global mean and relative sea-level changes over the past 66 Myr: implications for early Eocene ice sheets. Earth Science, Systems and Society 2024; 3: 10091. https: // doi. org / 10.3389 / esss. 2023.10091" type="journal volume" year="2024">2024</bibRefCitation>
), which is likely to have isolated each delta region (
<bibRefCitation id="564EB68A320AFFC6FFE5FD047D3557FB" author="Dolby GA &amp; Bedolla AM &amp; Bennett SEK" box="[124,308,708,732]" pageId="9" refId="ref15502" refString="Dolby GA, Bedolla AM, Bennett SEK et al. Global physical controls on estuarine habitat distribution during sea level change: consequences for genetic diversification through time. Global and Planetary Change 2020; 187: 103128. https: // doi. org / 10.1016 / j. gloplacha. 2020.103128" type="journal volume" year="2020">
Dolby
<emphasis id="00AB1769320AFFC6FF5CFD057CF057FB" box="[197,241,708,732]" italics="true" pageId="9">et al</emphasis>
. 2020
</bibRefCitation>
). Subsequent sea-level recessions 46.9 and 45.6 Mya (
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Miller
<emphasis id="00AB1769320AFFC6FEB3FD247D5557DC" box="[298,340,739,763]" italics="true" pageId="9">et al</emphasis>
. 2020a
</bibRefCitation>
) might have amplified barriers to dispersal.
</paragraph>
<paragraph id="3260CB7B320AFFC6FF14FCE27DB651D6" blockId="9.[112,764,144,1954]" pageId="9">
Divergence of Cubanichthyidae from
<taxonomicName id="F5DFB0F8320AFFC6FFE8FC817D04567E" box="[113,261,833,857]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">Goodeidae-Profundulidae</taxonomicName>
, 54.8–35.8 Mya (
<figureCitation id="AAE4D7FE320AFFC6FE27FC817DF9567E" box="[446,504,833,857]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
, node 3;
<tableCitation id="7F5DFEC0320AFFC6FDC7FC817EA8567E" box="[606,681,833,857]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="9">Table 2</tableCitation>
), could also be linked to sea-level fluctuations. We hypothesize that maximum sea levels associated with ice-free conditions in the Early Eocene, including a peak at 48.0 Mya (
<bibRefCitation id="564EB68A320AFFC6FDA2FC5F7EE85690" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[571,745,927,951]" pageId="9" refId="ref18344" refString="Miller KG, Schmelz WJ, Browning JV et al. Global mean and relative sea-level changes over the past 66 Myr: implications for early Eocene ice sheets. Earth Science, Systems and Society 2024; 3: 10091. https: // doi. org / 10.3389 / esss. 2023.10091" type="journal volume" year="2024">
Miller
<emphasis id="00AB1769320AFFC6FD19FC607EA85690" box="[640,681,927,951]" italics="true" pageId="9">et al</emphasis>
. 2024
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), allowed an ancestral taxon to range across the
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platform, into the Caribbean Sea. If so, then subsequent sea-level falls in the Middle Eocene, along with failure to return to ice-free conditions until the end of the Eocene (
<bibRefCitation id="564EB68A320AFFC6FE6CFBDD7EA95112" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[501,680,1053,1077]" pageId="9" refId="ref18344" refString="Miller KG, Schmelz WJ, Browning JV et al. Global mean and relative sea-level changes over the past 66 Myr: implications for early Eocene ice sheets. Earth Science, Systems and Society 2024; 3: 10091. https: // doi. org / 10.3389 / esss. 2023.10091" type="journal volume" year="2024">
Miller
<emphasis id="00AB1769320AFFC6FDA2FBDD7E6C5112" box="[571,621,1053,1077]" italics="true" pageId="9">et al.</emphasis>
2024
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), could have made the
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platform a more formidable barrier, isolating the ancestor of Cubanichthyidae in the Caribbean (
<figureCitation id="AAE4D7FE320AFFC6FFE5FBBB7CBB51B4" box="[124,186,1147,1171]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="9">Fig. 6</figureCitation>
). However, as already noted, phylogenetic placement of Cubanichthyidae is inconsistent among recent studies (e.g.
<bibRefCitation id="564EB68A320AFFC6FFE8FB797D6651F6" author="Ghezelayagh A &amp; Harrington RC &amp; Burress ED" box="[113,359,1209,1233]" pageId="9" pagination="1211 - 20" refId="ref16688" refString="Ghezelayagh A, Harrington RC, Burress ED et al. Prolonged morphological expansion of spiny-rayed fishes following the end- Cretaceous. Nature Ecology &amp; Evolution 2022; 6: 1211 - 20. https: // doi. org / 10.1038 / s 41559 - 022 - 01801 - 3" type="journal article" year="2022">
Ghezelayagh
<emphasis id="00AB1769320AFFC6FF65FB7A7D2C51F6" box="[252,301,1209,1233]" italics="true" pageId="9">et al.</emphasis>
2022
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FEECFB797E1E51F6" author="Piller KR &amp; Parker E &amp; Lemmon AR" box="[373,543,1209,1233]" pageId="9" refId="ref19125" refString="Piller KR, Parker E, Lemmon AR et al. Investigating the utility of anchored hybrid enrichment data to investigate the relationships among the killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed group of fishes. Molecular Phylogenetics and Evolution 2022; 173: 107482. https: // doi. org / 10.1016 / j. ympev. 2022.107482" type="journal volume" year="2022">
Piller
<emphasis id="00AB1769320AFFC6FE2AFB7A7DE551F6" box="[435,484,1209,1233]" italics="true" pageId="9">et al.</emphasis>
2022
</bibRefCitation>
), indicating that this family requires additional study.
</paragraph>
<paragraph id="3260CB7B320AFFC6FF14FB3878A85484" blockId="9.[112,764,144,1954]" lastBlockId="9.[809,1461,144,419]" pageId="9">
Divergence of
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from
<taxonomicName id="F5DFB0F8320AFFC6FE5EFB387E5A5037" box="[455,603,1272,1296]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
, 39.2–21.4 Mya (
<figureCitation id="AAE4D7FE320AFFC6FFE5FAD77CB55008" box="[124,180,1303,1327]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
, node 4), presumably reflects upland isolation of lineages in separate river drainages (
<tableCitation id="7F5DFEC0320AFFC6FE1EFAF77DD35068" box="[391,466,1335,1359]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="9">Table 2</tableCitation>
;
<figureCitation id="AAE4D7FE320AFFC6FE47FAF77E175068" box="[478,534,1335,1359]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
). We hypothesize that
<taxonomicName id="F5DFB0F8320AFFC6FFE8FA967CE25049" box="[113,227,1366,1390]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
descend from a population that ranged inland into a Madrean River (
<figureCitation id="AAE4D7FE320AFFC6FEB5FAB57D6450AA" box="[300,357,1397,1421]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="9">Fig. 6</figureCitation>
), which joined the Bejuco delta system in the Early Eocene, after uplift of the
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Arch diverted it southwards (
<bibRefCitation id="564EB68A320AFFC6FE96FA747E3A50EB" author="Snedden JW &amp; Galloway WE" box="[271,571,1460,1484]" pageId="9" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
). As the Madrean River drainage coalesced among emerging tectonic basins and colliding terranes of southwestern North America (
<bibRefCitation id="564EB68A320AFFC6FDE0FA337CA4530D" author="Lawton TF &amp; Bradford IA &amp; Vega FJ" pageId="9" pagination="820 - 36" refId="ref17594" refString="Lawton TF, Bradford IA, Vega FJ et al. Provenance of Upper Cretaceous - Paleogene sandstones in the foreland basin system of the Sierra Madre Oriental, northeastern Mexico, and its bearing on fluvial dispersal systems of the Mexican Laramide Province. Geological Society of America Bulletin 2009; 121: 820 - 36. https: // doi. org / 10.1130 / b 26450.1" type="journal article" year="2009">
Lawton
<emphasis id="00AB1769320AFFC6FD54FA337EFA532C" box="[717,763,1523,1547]" italics="true" pageId="9">et al.</emphasis>
2009
</bibRefCitation>
), not necessarily incorporating any pre-existing rivers, this drainage might have been depauperate in freshwater fishes or lacked them entirely. If so, the Madrean River system might have provided open niches for coastal killifishes transitioning into freshwater habitats. In the Middle–Late Eocene, the outlet of the Madrean River shifted between the
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Diversion and Río Bravo delta, but at the end of the Eocene (33.9 Mya) the Río Grande captured the inland drainage (
<bibRefCitation id="564EB68A320AFFC6FE68F92D7EBB5222" author="Galloway WE &amp; Whiteaker TL &amp; Ganey-Curry P" box="[497,698,1773,1797]" pageId="9" pagination="938 - 73" refId="ref16432" refString="Galloway WE, Whiteaker TL, Ganey-Curry P. History of Cenozoic North American drainage basin evolution, sediment yield, and accumulation in the Gulf of Mexico basin. Geosphere 2011; 7: 938 - 73. https: // doi. org / 10.1130 / ges 00647.1" type="journal article" year="2011">
Galloway
<emphasis id="00AB1769320AFFC6FDCCF92E7E825222" box="[597,643,1773,1797]" italics="true" pageId="9">et al.</emphasis>
2011
</bibRefCitation>
) (
<figureCitation id="AAE4D7FE320AFFC6FD4DF92D7C7F5203" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="9">Fig. 7</figureCitation>
). A resulting influx of freshwater fishes might have constrained the distributions of goodeids to more marginal habitats isolated by distance upstream, barriers, or habitat conditions (i.e. salinity), presumably isolating upland populations from coastal relatives. For
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, the timing of divergence corresponds with uplift of the
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Massif (40–25 Mya) (
<bibRefCitation id="564EB68A320AFFC6FA8CFF507FAA55E0" author="Villagomez D &amp; Pindell J" pageId="9" pagination="421 - 38" refId="ref20712" refString="Villagomez D, Pindell J. Cooling and uplift history of the Chiapas Massif and its influence on sedimentation and deformation in the adjacent Sierra de Chiapas Basin. In: Martens UC, Garza RSM (eds.), The Geological Society of America Special Paper, 546. Boulder: Geological Society of America, 2020, 421 - 38." type="book chapter" year="2020">Villagómez and Pindell 2020</bibRefCitation>
). Drainage from the Massif to the Bay of
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was localized at this time, not reaching the Gulf of
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basin (
<bibRefCitation id="564EB68A320AFFC6FCACFF2E783C5421" author="Beltran-Trivino A &amp; Martens UC &amp; von Quadt A" box="[821,1085,238,262]" pageId="9" pagination="1 - 34" refId="ref14685" refString="Beltran-Trivino A, Martens UC, von Quadt A. Siliciclastic provenance of the Cenozoic stratigraphic succession in the southern Gulf of Mexico: insights from U-Pb detrital zircon geochronology and heavy minerals analysis. In: Martens UC, Molina Garza RS, (eds), Southern and Central Mexico: Basement Framework, Tectonic Evolution, and Provenance of Mesozoic - Cenozoic Basins. Geological Society of America Special Paper 546. Boulder: Geological Society of America, 2021, 1 - 34." type="book chapter" year="2021">
Beltrán-Triviño
<emphasis id="00AB1769320AFFC6FC4EFF2F78045421" box="[983,1029,238,262]" italics="true" pageId="9">et al.</emphasis>
2021
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FBD1FF2E79275421" author="Villagomez D &amp; Steffensen C &amp; Pindell J" box="[1096,1318,238,262]" pageId="9" refId="ref20776" refString="Villagomez D, Steffensen C, Pindell J et al. Tectono-sedimentary evolution of Southern Mexico. Implications for Cretaceous and younger source-to-sink systems in the Mexican foreland basins and the Gulf of Mexico. Earth-Science Reviews 2022; 231: 104066. https: // doi. org / 10.1016 / j. earscirev. 2022.104066" type="journal volume" year="2022">
Villagómez
<emphasis id="00AB1769320AFFC6FB26FF2F78EF5421" box="[1215,1262,238,262]" italics="true" pageId="9">et al.</emphasis>
2022
</bibRefCitation>
), isolating inland populations (
<figureCitation id="AAE4D7FE320AFFC6FC70FECD78225402" box="[1001,1059,269,293]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="9">Fig. 7</figureCitation>
). Given that cyprinodontids evidently inhabited the southwestern Gulf by ~31.8 Mya (see below;
<tableCitation id="7F5DFEC0320AFFC6FCB3FE8C7F775443" box="[810,886,332,356]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="9">Table 2</tableCitation>
;
<figureCitation id="AAE4D7FE320AFFC6FC1AFE8C7FBD5444" box="[899,956,332,356]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="9">Fig. 7</figureCitation>
), we cannot rule out the possibility that a southward cyprinodontid invasion contributed to the disappearance of coastal
<taxonomicName id="F5DFB0F8320AFFC6FC08FE4B78035484" box="[913,1026,395,419]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
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.
</paragraph>
</subSubSection>
</treatment>
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<mods:namePart id="6331091A3B2BE31BC086D731AC2655C3">Hoagstrom, Christopher W.</mods:namePart>
<mods:affiliation id="88819C0C28360208DD2840B876BBFA82">Department of Zoology, Weber State University, 1415 Edvalson, Ogden, UT, USA</mods:affiliation>
</mods:name>
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<mods:namePart id="10C4247EB44C45750ECC0CB483DE2749">Matamoros, Wilfredo A.</mods:namePart>
<mods:affiliation id="6340E971ABC2512362400B0ED07CF1BC">Programa de Licenciatura, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Libramiento Norte Poniente No. 1150, Colonia Lajas Maciel, C. P. 29039, Tuxtla Gutiérrez, Chiapas, México &amp; Maestría en Ciencias en Biodiversidad y Conservación de Ecosistemas Tropicales, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Tuxtla Gutiérrez, Chiapas, México</mods:affiliation>
<mods:nameIdentifier id="5582B738745480DE4CD1AD9D3EF48A31" type="email">wilmatamoros@yahoo.com</mods:nameIdentifier>
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<mods:typeOfResource id="3097CC799AF267BA9175289956D7B622">text</mods:typeOfResource>
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<mods:titleInfo id="68DEE32367DBD6995008DF767535659C">
<mods:title id="765F2255A9E1E5DBD3A6A04AC0BCBE08">Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part id="90328077904E104F2ADD16931D0B49D6">
<mods:date id="B0DE30235BD9A0D79FC5D59074F93C0D">2024</mods:date>
<mods:detail id="57016F8F6C0318BFA7BB0D1A403098CE" type="pubDate">
<mods:number id="DC5C5DC1DF52B09F443A99FF47818B3E">2024-10-26</mods:number>
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<mods:number id="F0C2D01FDB0BA550951163FD5FC56852">202</mods:number>
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<mods:number id="23AC87D9E855A0AB088B5D5AA0108247">2</mods:number>
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<mods:url id="8B3CEEF975DBFD1A4D59EAC7785C9EA3">https://doi.org/10.1093/zoolinnean/zlae105</mods:url>
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<mods:identifier id="53EA97691F02F68120153B3A540B88C1" type="DOI">10.1093/zoolinnean/zlae105</mods:identifier>
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<subSubSection id="7AC598F0320AFFC6FBA5FA8F78A0504E" box="[1084,1185,1359,1385]" pageId="9" type="nomenclature">
<paragraph id="3260CB7B320AFFC6FBA5FA8F78A0504E" blockId="9.[810,1461,1359,1954]" box="[1084,1185,1359,1385]" pageId="9">
<taxonomicName id="F5DFB0F8320AFFC6FBA5FA8F78A0504E" ID-CoL="623LX" box="[1084,1185,1359,1385]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">
<emphasis id="00AB1769320AFFC6FBA5FA8F78A0504E" box="[1084,1185,1359,1385]" italics="true" pageId="9">Goodeidae</emphasis>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="7AC598F0320AFFC2FCB3FAB57CB75718" lastPageId="13" pageId="9" type="discussion">
<paragraph id="3260CB7B320AFFC4FCB3FAB578E9529B" blockId="9.[810,1461,1359,1954]" lastBlockId="11.[810,1460,1769,1981]" lastPageId="11" pageId="9">
Through the Late Eocene and Oligocene, reorganization and drainage expansion within the Madrean River (
<bibRefCitation id="564EB68A320AFFC6FAACFA557F8350EB" author="Galloway WE &amp; Whiteaker TL &amp; Ganey-Curry P" pageId="9" pagination="938 - 73" refId="ref16432" refString="Galloway WE, Whiteaker TL, Ganey-Curry P. History of Cenozoic North American drainage basin evolution, sediment yield, and accumulation in the Gulf of Mexico basin. Geosphere 2011; 7: 938 - 73. https: // doi. org / 10.1130 / ges 00647.1" type="journal article" year="2011">
Galloway
<emphasis id="00AB1769320AFFC6FA3BFA557F4250EB" italics="true" pageId="9">et al.</emphasis>
2011
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FC0AFA7478CD50EB" author="Snedden JW &amp; Galloway WE" box="[915,1228,1460,1484]" pageId="9" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
) could have allowed goodeids to disperse along the front of the Sierra Madre Occidental to the southern Great Basin (
<figureCitation id="AAE4D7FE320AFFC6FB4BFA33790D532D" box="[1234,1292,1523,1547]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="9">Fig. 7</figureCitation>
). Divergence of Great Basin Empetrichthyinae from
<taxonomicName id="F5DFB0F8320AFFC6FB36F9D27920530D" box="[1199,1313,1554,1578]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
of the Trans-Mexican Volcanic Belt and Sierra Madre 23.3–12.9 Mya (
<figureCitation id="AAE4D7FE320AFFC6FA14F9F17F36534E" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
, node 6;
<tableCitation id="7F5DFEC0320AFFC6FC02F9917FE7534E" box="[923,998,1617,1641]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="9">Table 2</tableCitation>
) agrees with break-up of this drainage at the end of the Oligocene (
<bibRefCitation id="564EB68A320AFFC6FB8DF9B0793D53AF" author="Snedden JW &amp; Galloway WE" box="[1044,1340,1648,1672]" pageId="9" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
). The great distance between northern (Empetrichthyinae) and southern (
<taxonomicName id="F5DFB0F8320AFFC6FCA8F96F7FA953E0" box="[817,936,1711,1735]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
) goodeids is a biogeographical anomaly (
<bibRefCitation id="564EB68A320AFFC6FAEEF96F7F5F53C1" author="Webb S" pageId="9" pagination="508 - 48" refId="ref20984" refString="Webb S. Goodeids. In: Warren L., Burr B (eds), Freshwater Fishes of North America. Baltimore: Johns Hopkins University Press, 2020, 508 - 48." type="book chapter" year="2020">Webb 2020</bibRefCitation>
).
<bibRefCitation id="564EB68A320AFFC6FCE6F90E782353C1" author="Parenti LR" box="[895,1058,1742,1766]" pageId="9" pagination="335 - 557" refId="ref18881" refString="Parenti LR. A phylogenetic and biogeographic analysis of Cyprinodontiformes fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 1981; 168: 335 - 557." type="journal article" year="1981">Parenti (1981)</bibRefCitation>
proposed that regional desiccation eliminated intervening populations (
<bibRefCitation id="564EB68A320AFFC6FB28F92D79AC5222" author="Grant EC &amp; Riddle BR" box="[1201,1453,1773,1797]" pageId="9" pagination="209 - 12" refId="ref16794" refString="Grant EC, Riddle BR. Are the endangered springfish (Crenichthys Hubbs) and poolfish (Empetrichthys Gilbert) fundulines or goodeids?: a mitochondrial DNA assessment. Copeia 1995; 1995: 209 - 12. https: // doi. org / 10.2307 / 1446817" type="journal article" year="1995">Grant and Riddle 1995</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FCB3F8CD7FE15202" author="Webb SA &amp; Graves JA &amp; Macias-Garcia C" box="[810,992,1805,1829]" pageId="9" pagination="527 - 44" refId="ref21020" refString="Webb SA, Graves JA, Macias-Garcia C et al. Molecular phylogeny of the livebearing Goodeidae (Cyprinodontiformes). Molecular Phylogenetics and Evolution 2004; 30: 527 - 44. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00257 - 4" type="journal article" year="2004">
Webb
<emphasis id="00AB1769320AFFC6FCE9F8CD7FA25202" box="[880,931,1805,1829]" italics="true" pageId="9">et al.</emphasis>
2004
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FC69F8CD78A85203" author="Miller RR &amp; Minckley WL &amp; Norris SM" box="[1008,1193,1804,1829]" pageId="9" refId="ref18488" refString="Miller RR, Minckley WL, Norris SM. Freshwater Fishes of Mexico. Chicago: The University of Chicago Press, 2005." type="book" year="2005">
Miller
<emphasis id="00AB1769320AFFC6FBA0F8CD786D5202" box="[1081,1132,1805,1829]" italics="true" pageId="9">et al.</emphasis>
2005
</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FB20F8CD79325202" author="Webb S" box="[1209,1331,1805,1829]" pageId="9" pagination="508 - 48" refId="ref20984" refString="Webb S. Goodeids. In: Warren L., Burr B (eds), Freshwater Fishes of North America. Baltimore: Johns Hopkins University Press, 2020, 508 - 48." type="book chapter" year="2020">Webb 2020</bibRefCitation>
), but living Empetrichthyinae occupy the driest region of North America (i.e. aridity is associated with survival). Fragmentation of the Madrean River might have been an additional cause of extirpations (
<bibRefCitation id="564EB68A320AFFC6FC13F84A78405285" author="Fagan WF &amp; Unmack PJ &amp; Burgess C" box="[906,1089,1930,1954]" pageId="9" pagination="3250 - 6" refId="ref16238" refString="Fagan WF, Unmack PJ, Burgess C et al. Rarity, fragmentation, and extinction risk in desert fishes. Ecology 2002; 83: 3250 - 6. https: // doi. org / 10.2307 / 3072075" type="journal article" year="2002">
Fagan
<emphasis id="00AB1769320AFFC6FC48F84B78055285" box="[977,1028,1930,1954]" italics="true" pageId="9">et al.</emphasis>
2002
</bibRefCitation>
), and climatic cooling might have reduced habitat suitability at higher elevations and latitudes, perhaps explaining why Empetrichthyinae are associated with warm-water springs. Loss of goodeids from the Gulf of
<collectingRegion id="F01B05993208FFC4FD34F8E77EFB5218" box="[685,762,1831,1855]" country="Mexico" name="Mexico" pageId="11">México</collectingRegion>
drainage could reflect Late Miocene integration of the ancestral Río Nazas with the Río Grande, which facilitated an influx of fishes (
<bibRefCitation id="564EB68A3208FFC4FF26F8467E0252BA" author="Hoagstrom CW &amp; Osborne MJ" box="[191,515,1925,1950]" pageId="11" pagination="20 - 76" refId="ref17354" refString="Hoagstrom CW, Osborne MJ. Biogeography of Cyprinodon across the Great Plains-Chihuahuan Desert Region and Adjacent Areas. Proceedings of the Desert Fishes Council Special Publication 2021; 2021: 20 - 76." type="journal article" year="2021">Hoagstrom and Osborne 2021</bibRefCitation>
).
<taxonomicName id="F5DFB0F83208FFC4FD84F8457CA4529A" authority="Gunther, 1866" authorityName="Gunther" authorityYear="1866" family="Goodeidae" genus="Characodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="genus">
<emphasis id="00AB17693208FFC4FD84F8457E9552BA" box="[541,660,1925,1949]" italics="true" pageId="11">Characodon</emphasis>
Günther, 1866
</taxonomicName>
persisted only in remote basins (†
<taxonomicName id="F5DFB0F83208FFC4FDB8F86578005226" authority=", Mayran-Parras Basin" authorityName="Mayran-Parras Basin" box="[545,1025,1769,1981]" family="Goodeidae" genus="Characodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="species" species="garmani">
<emphasis id="00AB17693208FFC4FDB8F8657EF4529B" box="[545,757,1957,1981]" italics="true" pageId="11">Characodon garmani</emphasis>
, Mayran-Parras Basin
</taxonomicName>
;
<taxonomicName id="F5DFB0F83208FFC4FB92F92979A45226" authority="Smith &amp; Miller, 1986" authorityName="Smith &amp; Miller" authorityYear="1986" box="[1035,1445,1769,1793]" family="Goodeidae" genus="Characodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="species" species="audax">
<emphasis id="00AB17693208FFC4FB92F92978C75226" box="[1035,1222,1769,1793]" italics="true" pageId="11">Characodon audax</emphasis>
<bibRefCitation id="564EB68A3208FFC4FB52F92979A45226" author="Miller RR &amp; Smith ML" box="[1227,1445,1769,1793]" pageId="11" pagination="487 - 517" refId="ref18512" refString="Miller RR, Smith ML. Origin and geography of the fishes of central Mexico. In: Hocutt CH, Wiley EO (eds), The Zoogeography of North American Freshwater Fishes. New York: John Wiley &amp; Sons, 1986, 487 - 517." type="book chapter" year="1986">Smith &amp; Miller, 1986</bibRefCitation>
</taxonomicName>
<emphasis id="00AB17693208FFC4FA3CF92979B25227" box="[1445,1459,1769,1792]" italics="true" pageId="11">–</emphasis>
<taxonomicName id="F5DFB0F83208FFC4FCB3F8C878955207" authority="Gunther, 1866" authorityName="Gunther" authorityYear="1866" box="[810,1172,1800,1824]" family="Goodeidae" genus="Characodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="species" species="lateralis">
<emphasis id="00AB17693208FFC4FCB3F8C87FF55207" box="[810,1012,1800,1824]" italics="true" pageId="11">Characodon lateralis</emphasis>
Günther, 1866
</taxonomicName>
, Río Tunal) (
<bibRefCitation id="564EB68A3208FFC4FAB8F8C87F935218" author="Beltran-Lopez RG &amp; Perez-Rodriguez R &amp; Montanez-Garcia OC" pageId="11" refId="ref14641" refString="Beltran-Lopez RG, Perez-Rodriguez R, Montanez-Garcia OC et al. Genetic differentiation in the genus Characodon: implications for conservation and taxonomy. PeerJ 2021; 9: e 11492. https: // doi. org / 10.7717 / peerj. 11492" type="journal volume" year="2021">
Beltrán-López
<emphasis id="00AB17693208FFC4FCB3F8E87F5B5218" box="[810,858,1831,1855]" italics="true" pageId="11">et al.</emphasis>
2021
</bibRefCitation>
), potentially protected from invading fishes. It is also possible that remnant goodeid populations were present in the uninhabited region before European settlement, but disappeared owing to habitat destruction, dewatering, or invasive species associated with early European contact.
</paragraph>
<caption id="66A09BF33209FFC5FF18F8EC7D2052BF" pageId="10" startId="10.[129,194,1836,1860]" targetBox="[212,1390,147,1805]" targetPageId="10" targetType="figure">
<paragraph id="3260CB7B3209FFC5FF18F8EC7D2052BF" blockId="10.[129,1461,1836,1944]" pageId="10">
<emphasis id="00AB17693209FFC5FF18F8EC7CD95263" bold="true" box="[129,216,1836,1860]" pageId="10">Figure 5.</emphasis>
Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae,
<taxonomicName id="F5DFB0F83209FFC5FBDDF8EC78EB5263" box="[1092,1258,1836,1860]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83209FFC5FB6BF8EC795E5263" box="[1266,1375,1836,1860]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
,
<taxonomicName id="F5DFB0F83209FFC5FF18F8887CED5247" box="[129,236,1864,1888]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
, and
<taxonomicName id="F5DFB0F83209FFC5FE85F8887DA75247" box="[284,422,1864,1888]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
. A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (
<bibRefCitation id="564EB68A3209FFC5FD0DF8A47F25525B" author="Rico CN &amp; Hoagstrom CW &amp; Elias DJ" box="[660,804,1892,1916]" pageId="10" refId="ref19830" refString="Rico CN, Hoagstrom CW, Elias DJ et al. Biotic regionalization of freshwater fishes in Northern Middle America highlights high beta diversity created by prominent biogeographic barriers. Frontiers of Biogeography 2022; 14: e 58095. https: // doi. org / 10.21425 / F 5 FBG 58095" type="journal volume" year="2022">
Rico
<emphasis id="00AB17693209FFC5FD5CF8A57EF0525B" box="[709,753,1892,1916]" italics="true" pageId="10">et al.</emphasis>
2022
</bibRefCitation>
). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS.
</paragraph>
</caption>
<caption id="66A09BF33208FFC4FFE8FAAD7EFD539E" pageId="11" startId="11.[113,178,1389,1413]" targetBox="[114,1458,144,1361]" targetPageId="11" targetType="figure">
<paragraph id="3260CB7B3208FFC4FFE8FAAD7EFD539E" blockId="11.[113,1456,1389,1721]" pageId="11">
<emphasis id="00AB17693208FFC4FFE8FAAD7CC850A2" bold="true" box="[113,201,1389,1413]" pageId="11">Figure 6.</emphasis>
Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. =
<taxonomicName id="F5DFB0F83208FFC4FE64FA497EE75086" authority=", Good." authorityName="Good." box="[509,742,1417,1441]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Cyprinodontidae, Good.</taxonomicName>
-Profund. =
<taxonomicName id="F5DFB0F83208FFC4FCC5FA497FC75086" box="[860,966,1417,1441]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83208FFC4FC4AFA49785E5086" box="[979,1119,1417,1441]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
). Phylogenetic nodes (Fig. 2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (
<bibRefCitation id="564EB68A3208FFC4FFE2FA397D225336" author="Cossey SPJ &amp; Van Nieuwenhuise D &amp; Davis J" box="[123,291,1529,1553]" pageId="11" pagination="63 - 80" refId="ref15238" refString="Cossey SPJ, Van Nieuwenhuise D, Davis J et al. Compelling evidence from eastern Mexico for a Late Paleocene / Early Eocene isolation, drawdown, and refill of the Gulf of Mexico. Interpretation 2016; 4: SC 63 - 80. https: // doi. org / 10.1190 / int- 2015 - 0107.1" type="journal article" year="2016">
Cossey
<emphasis id="00AB17693208FFC4FF5DFA3A7CEE5336" box="[196,239,1529,1553]" italics="true" pageId="11">et al.</emphasis>
2016
</bibRefCitation>
), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of
<bibRefCitation id="564EB68A3208FFC4FDFCF9D57F86530A" author="Snedden JW &amp; Galloway WE" box="[613,903,1557,1581]" pageId="11" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway (2019)</bibRefCitation>
. Details on southwestern Gulf tributaries are from studies by
<bibRefCitation id="564EB68A3208FFC4FF4CF9F17DA1536E" author="Snedden JW &amp; Galloway WE &amp; Milliken KT" box="[213,416,1585,1609]" pageId="11" pagination="768 - 84" refId="ref20398" refString="Snedden JW, Galloway WE, Milliken KT et al. Validation of empirical source-to-sink scaling relationships in a continental-scale system: the Gulf of Mexico basin Cenozoic record. Geosphere 2018; 14: 768 - 84. https: // doi. org / 10.1130 / ges 01452.1" type="journal article" year="2018">
Snedden
<emphasis id="00AB17693208FFC4FEB4F9F27D59536E" box="[301,344,1585,1609]" italics="true" pageId="11">et al.</emphasis>
(2018)
</bibRefCitation>
and
<bibRefCitation id="564EB68A3208FFC4FE57F9F17E8B536E" author="Cossey SPJ &amp; Rosenfeld J &amp; Bitter M" box="[462,650,1585,1609]" pageId="11" pagination="123 - 41" refId="ref15205" refString="Cossey SPJ, Rosenfeld J, Bitter M et al. Update on the Paleogene water-level drawdown hypothesis, Gulf of Mexico. GCAGS Journal 2021; 10: 123 - 41." type="journal article" year="2021">
Cossey
<emphasis id="00AB17693208FFC4FD8EF9F27E43536E" box="[535,578,1585,1609]" italics="true" pageId="11">et al.</emphasis>
(2021)
</bibRefCitation>
. Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of
<taxonomicName id="F5DFB0F83208FFC4FCB9F98D7F885342" box="[800,905,1613,1637]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route. High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated
<taxonomicName id="F5DFB0F83208FFC4FCC4F945780353BA" box="[861,1026,1669,1693]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
from
<taxonomicName id="F5DFB0F83208FFC4FBA3F94578A753BA" box="[1082,1190,1669,1693]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Fundulidae</taxonomicName>
in the north (N2) and Cubanichthyidae from
<taxonomicName id="F5DFB0F83208FFC4FED5F9617DB5539E" box="[332,436,1697,1721]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
–
<taxonomicName id="F5DFB0F83208FFC4FE59F9617E48539E" box="[448,585,1697,1721]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
in the south (N3).
</paragraph>
</caption>
<caption id="66A09BF3320FFFC3FF18FA8679255305" pageId="12" startId="12.[129,194,1350,1374]" targetBox="[129,1473,144,1323]" targetPageId="12" targetType="figure">
<paragraph id="3260CB7B320FFFC3FF18FA8679255305" blockId="12.[129,1468,1350,1571]" pageId="12">
<emphasis id="00AB1769320FFFC3FF18FA867CD95078" bold="true" box="[129,216,1350,1375]" pageId="12">Figure 7.</emphasis>
Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled. Phylogenetic nodes (Fig. 3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig. 6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by
<bibRefCitation id="564EB68A320FFFC3FB0AFA5A79B75094" author="Snedden JW &amp; Galloway WE" box="[1171,1462,1434,1459]" pageId="12" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway (2019)</bibRefCitation>
. Details on the Chiapas region are from the study by
<bibRefCitation id="564EB68A320FFFC3FDF3FA767F4C50E8" author="Villagomez D &amp; Steffensen C &amp; Pindell J" box="[618,845,1462,1487]" pageId="12" refId="ref20776" refString="Villagomez D, Steffensen C, Pindell J et al. Tectono-sedimentary evolution of Southern Mexico. Implications for Cretaceous and younger source-to-sink systems in the Mexican foreland basins and the Gulf of Mexico. Earth-Science Reviews 2022; 231: 104066. https: // doi. org / 10.1016 / j. earscirev. 2022.104066" type="journal volume" year="2022">
Villagómez
<emphasis id="00AB1769320FFFC3FD40FA777F0550E9" box="[729,772,1462,1486]" italics="true" pageId="12">et al.</emphasis>
(2022)
</bibRefCitation>
. Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding
<emphasis id="00AB1769320FFFC3FE26FA127EE450CD" box="[447,741,1490,1514]" italics="true" pageId="12">
<taxonomicName id="F5DFB0F8320FFFC3FE26FA127E3250CD" authorityName="Hubbs" authorityYear="1926" box="[447,563,1490,1514]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F8320FFFC3FDA7FA127EE450CD" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[574,741,1490,1514]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="species" species="pulchra">Jordanella pulchra</taxonomicName>
</emphasis>
on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding
<taxonomicName id="F5DFB0F8320FFFC3FD63FA2E7F6E5321" authorityName="Myers" authorityYear="1924" box="[762,879,1518,1542]" family="Fundulidae" genus="Leptolucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320FFFC3FD63FA2E7F6E5321" box="[762,879,1518,1542]" italics="true" pageId="12">Leptolucania</emphasis>
</taxonomicName>
on the Ocala High; and N13, fundulid westward immigration founding
<emphasis id="00AB1769320FFFC3FF45F9CA7DA25305" box="[220,419,1546,1570]" italics="true" pageId="12">
<taxonomicName id="F5DFB0F8320FFFC3FF45F9CA7D275305" authorityName="Girard" authorityYear="1860" box="[220,294,1546,1570]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="genus">Lucania</taxonomicName>
–
<taxonomicName id="F5DFB0F8320FFFC3FEABF9CA7DA25305" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[306,419,1546,1570]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
</emphasis>
in the Río Grande–Río Bravo estuary, where it presumably co-occurred with
<taxonomicName id="F5DFB0F8320FFFC3FBE0F9CA791E5305" box="[1145,1311,1546,1570]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="3260CB7B320FFFC3FF05F9937F965261" blockId="12.[126,779,1618,1987]" lastBlockId="12.[825,1476,1618,1987]" pageId="12">
Because this scenario agrees with reconstructed hydrography, it is unnecessary to invoke the popular hypothesis (
<bibRefCitation id="564EB68A320FFFC3FD05F9B27CF6538E" author="Miller RR &amp; Smith ML" pageId="12" pagination="487 - 517" refId="ref18512" refString="Miller RR, Smith ML. Origin and geography of the fishes of central Mexico. In: Hocutt CH, Wiley EO (eds), The Zoogeography of North American Freshwater Fishes. New York: John Wiley &amp; Sons, 1986, 487 - 517." type="book chapter" year="1986">Miller and Smith 1986</bibRefCitation>
,
<bibRefCitation id="564EB68A320FFFC3FE98F9517DCD538E" author="Minckley WL &amp; Hendrickson DA &amp; Bond CE" box="[257,460,1681,1705]" pageId="12" pagination="519 - 613" refId="ref18561" refString="Minckley WL, Hendrickson DA, Bond CE. Geography of western North American freshwater fishes: description and relations to intracontinental tectonism. In: Hocutt CH, Wiley EO (eds), Zoogeography of Western North American Freshwater Fishes. New York: John Wiley and Sons, 1986, 519 - 613." type="book chapter" year="1986">
Minckley
<emphasis id="00AB1769320FFFC3FEFFF9527D95538E" box="[358,404,1681,1705]" italics="true" pageId="12">et al.</emphasis>
1986
</bibRefCitation>
,
<bibRefCitation id="564EB68A320FFFC3FE4FF9517EC1538E" author="Grant EC &amp; Riddle BR" box="[470,704,1681,1705]" pageId="12" pagination="209 - 12" refId="ref16794" refString="Grant EC, Riddle BR. Are the endangered springfish (Crenichthys Hubbs) and poolfish (Empetrichthys Gilbert) fundulines or goodeids?: a mitochondrial DNA assessment. Copeia 1995; 1995: 209 - 12. https: // doi. org / 10.2307 / 1446817" type="journal article" year="1995">Grant and Riddle 1995</bibRefCitation>
,
<bibRefCitation id="564EB68A320FFFC3FD53F9517CE853EF" author="Miller RR &amp; Minckley WL &amp; Norris SM" pageId="12" refId="ref18488" refString="Miller RR, Minckley WL, Norris SM. Freshwater Fishes of Mexico. Chicago: The University of Chicago Press, 2005." type="book" year="2005">
Miller
<emphasis id="00AB1769320FFFC3FF18F9717CB153EF" box="[129,176,1712,1736]" italics="true" pageId="12">et al.</emphasis>
2005
</bibRefCitation>
,
<bibRefCitation id="564EB68A320FFFC3FF6DF9707D6B53EE" author="Webb S" box="[244,362,1712,1737]" pageId="12" pagination="508 - 48" refId="ref20984" refString="Webb S. Goodeids. In: Warren L., Burr B (eds), Freshwater Fishes of North America. Baltimore: Johns Hopkins University Press, 2020, 508 - 48." type="book chapter" year="2020">Webb 2020</bibRefCitation>
) that northward drift of the Pacific Plate created the gap between Empetrichthyinae and
<taxonomicName id="F5DFB0F8320FFFC3FD0AF9107F0653CF" box="[659,775,1744,1768]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="family">Goodeidae</taxonomicName>
. Furthermore,thehypothesisthatGoodeidaeusedawesternroute from the Great Basin into central
<collectingCountry id="4AC88BEB320FFFC3FE57F8CE7E1A5201" box="[462,539,1806,1830]" name="Mexico" pageId="12">México</collectingCountry>
(
<bibRefCitation id="564EB68A320FFFC3FDB4F8CE7CB45261" author="Perez-Rodriguez R &amp; Dominguez-Dominguez O &amp; Doadrio I" pageId="12" pagination="993 - 1015" refId="ref19076" refString="Perez-Rodriguez R, Dominguez-Dominguez O, Doadrio I et al. Comparative historical biogeography of three groups of Nearctic freshwater fishes across central Mexico. Journal of Fish Biology 2015; 86: 993 - 1015. https: // doi. org / 10.1111 / jfb. 12611" type="journal article" year="2015">
Pérez-Rodríguez
<emphasis id="00AB1769320FFFC3FD45F8CF7F0B5201" box="[732,778,1806,1830]" italics="true" pageId="12">et al.</emphasis>
2015
</bibRefCitation>
) is unnecessary because goodeids dispersing inland from the Gulf of
<collectingCountry id="4AC88BEB320FFFC3FF65F88D7D485242" box="[252,329,1869,1893]" name="Mexico" pageId="12">México</collectingCountry>
via the Madrean River would have reached central
<collectingCountry id="4AC88BEB320FFFC3FF54F8AC7D1B52A3" box="[205,282,1900,1924]" name="Mexico" pageId="12">México</collectingCountry>
en route to the Great Basin (
<figureCitation id="AAE4D7FE320FFFC3FDA5F8AC7E7552A3" box="[572,628,1900,1924]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="12">Fig. 7</figureCitation>
), not from the Great Basin. Separation of Empetrichthyinae from
<taxonomicName id="F5DFB0F8320FFFC3FD1BF84C7EF55283" box="[642,756,1932,1956]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
is attributable to vicariance during fragmentation of the Madrean River drainage, with Empetrichthyinae representing a relict lineage that has persisted in the Mojave Desert after reaching the region when the Madrean River extended from the Gulf Coast to the Basin and Range geomorphic province.
<taxonomicName id="F5DFB0F8320FFFC3FA89F970798353EF" box="[1296,1410,1712,1736]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
represent lineages descending from populations further downstream that persisted within the Sierra Madre Occidental and Trans-Mexican Volcanic Belt after separation from the Gulf of
<collectingCountry id="4AC88BEB320FFFC3FAECF8CE79C35201" box="[1397,1474,1806,1830]" name="Mexico" pageId="12">México</collectingCountry>
drainage.
</paragraph>
<paragraph id="3260CB7B320FFFC2FCCCF88D7CB75718" blockId="12.[825,1476,1618,1987]" lastBlockId="13.[113,763,144,575]" lastPageId="13" pageId="12">
Within
<taxonomicName id="F5DFB0F8320FFFC3FC25F88D782E5242" box="[956,1071,1869,1893]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
, divergence of CharacodontiniIllyodontini from crown
<taxonomicName id="F5DFB0F8320FFFC3FBD0F8AC78BA52A3" box="[1097,1211,1900,1924]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
16.8–10.3 Mya (
<figureCitation id="AAE4D7FE320FFFC3FAE7F8AC79BD52A2" box="[1406,1468,1900,1925]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="12">Fig. 3</figureCitation>
, node 7;
<tableCitation id="7F5DFEC0320FFFC3FC16F84C7FD85283" box="[911,985,1932,1956]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="12">Table 2</tableCitation>
) appears to have been an east–west divergence, possibly between ancestral ríos Nazas and Aguanaval (
<figureCitation id="AAE4D7FE320FFFC3FAEFF86B79B052E4" box="[1398,1457,1963,1987]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="12">Fig. 8</figureCitation>
). Reactivation of the San Marcos Fault 14.0–5.0 Mya and contemporary volcanism on the borders of the Mesa Central (
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Aranda-Gómez
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, 2007,
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Chávez-Cabello
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2007
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,
<bibRefCitation id="564EB68A320EFFC2FFE8FF2E7D985421" author="Nieto-Samaniego AF &amp; Alaniz-Alvarez SA &amp; Camprubi A" box="[113,409,238,262]" pageId="13" pagination="41 - 70" refId="ref18673" refString="Nieto-Samaniego AF, Alaniz-Alvarez SA, Camprubi A. Mesa Central of Mexico: stratigraphy, structure, and Cenozoic tectonic evolution. Special Paper of the Geological Society of America 2007; 422: 41 - 70. https: // doi. org / 10.1130 / 2007.2422 (02)" type="journal article" year="2007">
Nieto-Samaniego
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) are likely to have contributed to drainage fragmentation. Positioning of crown
<taxonomicName id="F5DFB0F8320EFFC2FDFBFECD7ED55402" box="[610,724,269,293]" family="Goodeidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="subFamily" subFamily="Goodeinae">Goodeinae</taxonomicName>
on the Trans-Mexican Volcanic Belt throughout its tectonic evolution, which initiated ~20 Mya and lasted 17 Myr (
<bibRefCitation id="564EB68A320EFFC2FDE7FE8C7CA454A4" author="Ferrari L &amp; Orozco-Esquivel T &amp; Manea V" pageId="13" pagination="122 - 49" refId="ref16330" refString="Ferrari L, Orozco-Esquivel T, Manea V et al. The dynamic history of the Trans-Mexican Volcanic Belt and the Mexico subduction zone. Tectonophysics 2012; 522 - 523: 122 - 49. https: // doi. org / 10.1016 / j. tecto. 2011.09.018" type="book chapter" year="2012">
Ferrari
<emphasis id="00AB1769320EFFC2FD53FE8D7EFA5443" box="[714,763,332,356]" italics="true" pageId="13">et al.</emphasis>
2012
</bibRefCitation>
), ensured that goodeids experienced frequent drainage reorganizations and shifting hydrographic barriers (
<bibRefCitation id="564EB68A320EFFC2FD3FFE4B7CA454E5" author="Barbour CD" pageId="13" pagination="533 - 56" refId="ref14550" refString="Barbour CD. A biogeographical history of Chirostoma (Pisces: Atherinidae): a species flock from the Mexican Plateau. Copeia 1973; 1973: 533 - 56. https: // doi. org / 10.2307 / 1443118" type="journal article" year="1973">Barbour 1973</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FF29FE6A7E1254E5" author="Dominguez-Dominguez O &amp; Pedraza-Lara C &amp; Gurrola-Sanchez N" box="[176,531,426,450]" pageId="13" pagination="13 - 30" refId="ref15712" refString="Dominguez-Dominguez O, Pedraza-Lara C, Gurrola-Sanchez N et al. Historical biogeography of the Goodeinae (Cyprinodontiforms). In: Uribe MC, Grier HJ (eds), Viviparous Fishes II. Homestead: New Life Publications, 2010, 13 - 30." type="book chapter" year="2010">
Domínguez‐Domínguez
<emphasis id="00AB1769320EFFC2FE37FE6B7DDD54E5" box="[430,476,426,450]" italics="true" pageId="13">et al.</emphasis>
2010
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,
<bibRefCitation id="564EB68A320EFFC2FD87FE6A7CA454C6" author="Perez-Rodriguez R &amp; Dominguez-Dominguez O &amp; Doadrio I" pageId="13" pagination="993 - 1015" refId="ref19076" refString="Perez-Rodriguez R, Dominguez-Dominguez O, Doadrio I et al. Comparative historical biogeography of three groups of Nearctic freshwater fishes across central Mexico. Journal of Fish Biology 2015; 86: 993 - 1015. https: // doi. org / 10.1111 / jfb. 12611" type="journal article" year="2015">
Pérez-Rodríguez
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</bibRefCitation>
). Barrier displacement via stream capture was probably the main mode of range expansion and diversification (
<bibRefCitation id="564EB68A320EFFC2FD11FE297CA45707" author="Webb SA &amp; Graves JA &amp; Macias-Garcia C" pageId="13" pagination="527 - 44" refId="ref21020" refString="Webb SA, Graves JA, Macias-Garcia C et al. Molecular phylogeny of the livebearing Goodeidae (Cyprinodontiformes). Molecular Phylogenetics and Evolution 2004; 30: 527 - 44. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00257 - 4" type="journal article" year="2004">
Webb
<emphasis id="00AB1769320EFFC2FD52FE297EFA5726" box="[715,763,489,513]" italics="true" pageId="13">et al.</emphasis>
2004
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FF2AFDC87E205707" author="Dominguez-Dominguez O &amp; Pedraza-Lara C &amp; Gurrola-Sanchez N" box="[179,545,520,544]" pageId="13" pagination="13 - 30" refId="ref15712" refString="Dominguez-Dominguez O, Pedraza-Lara C, Gurrola-Sanchez N et al. Historical biogeography of the Goodeinae (Cyprinodontiforms). In: Uribe MC, Grier HJ (eds), Viviparous Fishes II. Homestead: New Life Publications, 2010, 13 - 30." type="book chapter" year="2010">
Domínguez‐Domínguez
<emphasis id="00AB1769320EFFC2FE2CFDC97DE75707" box="[437,486,520,544]" italics="true" pageId="13">et al.</emphasis>
2010
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FDB6FDC87CA45718" author="Beltran-Lopez RG &amp; Perez-Rodriguez R &amp; Montanez-Garcia OC" pageId="13" refId="ref14641" refString="Beltran-Lopez RG, Perez-Rodriguez R, Montanez-Garcia OC et al. Genetic differentiation in the genus Characodon: implications for conservation and taxonomy. PeerJ 2021; 9: e 11492. https: // doi. org / 10.7717 / peerj. 11492" type="journal volume" year="2021">
Beltrán-López
<emphasis id="00AB1769320EFFC2FD50FDC97EFA5707" box="[713,763,520,544]" italics="true" pageId="13">et al.</emphasis>
2021
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

171
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<paragraph id="3260CB7B320AFFC6FBB5FE0278B354FB" blockId="9.[1068,1202,450,476]" box="[1068,1202,450,476]" pageId="9">
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<emphasis id="00AB1769320AFFC6FBB5FE0278B354FB" box="[1068,1202,450,476]" italics="true" pageId="9">Profundulidae</emphasis>
</taxonomicName>
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<subSubSection id="7AC598F0320AFFC6FCB3FE297FF25008" pageId="9" type="discussion">
<paragraph id="3260CB7B320AFFC6FCB3FE2979695690" blockId="9.[809,1461,489,1327]" pageId="9">
The major node within
<taxonomicName id="F5DFB0F8320AFFC6FBBFFE2978B85726" box="[1062,1209,489,513]" family="Profundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="family">Profundulidae</taxonomicName>
30.7–14.2 Mya (
<figureCitation id="AAE4D7FE320AFFC6FAEBFE2979AC5726" box="[1394,1453,489,513]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
, node 5) separates
<taxonomicName id="F5DFB0F8320AFFC6FC7CFDC878E75707" authority="Hubbs, 1924" authorityName="Hubbs" authorityYear="1924" box="[997,1254,520,544]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FC7CFDC8785D5707" box="[997,1116,520,544]" italics="true" pageId="9">Profundulus</emphasis>
Hubbs, 1924
</taxonomicName>
from
<taxonomicName id="F5DFB0F8320AFFC6FABDFDC87FF25718" authority="Alvarez and Carranza, 1951" authorityName="Alvarez and Carranza" authorityYear="1951" family="Profundulidae" genus="Tlaloc" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FABDFDC879635707" box="[1316,1378,520,544]" italics="true" pageId="9">Tlaloc</emphasis>
Álvarez and Carranza, 1951
</taxonomicName>
.
<taxonomicName id="F5DFB0F8320AFFC6FC63FDE778395718" authorityName="Alvarez and Carranza" authorityYear="1951" box="[1018,1080,551,575]" family="Profundulidae" genus="Tlaloc" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FC63FDE778395718" box="[1018,1080,551,575]" italics="true" pageId="9">Tlaloc</emphasis>
</taxonomicName>
occurs across the
<collectingRegion id="F01B0599320AFFC6FB77FDE779405718" box="[1262,1345,551,575]" country="Mexico" name="Chiapas" pageId="9">Chiapas</collectingRegion>
Massif and adjacent highlands within the Río Grijalva drainage (
<bibRefCitation id="564EB68A320AFFC6FAAEFD8779AC5779" author="Miller RR" box="[1335,1453,582,607]" pageId="9" pagination="1 - 64" refId="ref18453" refString="Miller RR. A systematic review of the Middle American fishes of the genus Profundulus. Miscellaneous Publications of the Museum of Zoology, University of Michigan 1955; 92: 1 - 64." type="journal article" year="1955">Miller 1955</bibRefCitation>
,
<bibRefCitation id="564EB68A320AFFC6FCB3FDA678335759" author="Cashner MF &amp; Echelle AA" box="[810,1074,614,638]" pageId="9" pagination="493 - 502" refId="ref15008" refString="Cashner MF, Echelle AA. Profundulidae: Middle American killifishes. North America, Vol. 2, Characidae to Poeciliidae. Baltimore: Johns Hopkins University Press, 2020, 493 - 502." type="book chapter" year="2020">Cashner and Echelle 2020</bibRefCitation>
). The Upper Río Grijalva flows northwest until making an abrupt turn northeast, suggesting capture from a north-flowing river, possibly the ancestral Río Uxpanapa or Tonalá. This capture could have been caused by uplift of the northwestern
<collectingRegion id="F01B0599320AFFC6FC24FD23781157DC" box="[957,1040,739,763]" country="Mexico" name="Chiapas" pageId="9">Chiapas</collectingRegion>
Massif 30–25 Mya (
<bibRefCitation id="564EB68A320AFFC6FB70FD23798B57DC" author="Witt C &amp; Brichau S &amp; Carter A" box="[1257,1418,739,763]" pageId="9" pagination="1 - 15" refId="ref21156" refString="Witt C, Brichau S, Carter A. New constraints on the origin of the Sierra Madre de Chiapas (south Mexico) from sediment provenance and apatite thermochronometry. Tectonics 2012; 31: 1 - 15." type="journal article" year="2012">
Witt
<emphasis id="00AB1769320AFFC6FA87FD24794E57DC" box="[1310,1359,739,763]" italics="true" pageId="9">et al.</emphasis>
2012
</bibRefCitation>
) or head-cutting of the lower Río Grijalva during a period of extensive erosion 35–25 Mya (
<bibRefCitation id="564EB68A320AFFC6FBAFFCE27905561D" author="Abdullin F &amp; Sole J &amp; Meneses-Rocha J de" box="[1078,1284,802,826]" pageId="9" pagination="32 - 48" refId="ref14204" refString="Abdullin F, Sole J, Meneses-Rocha J de J et al. LA-ICP-MS-based apatite fission track dating of the Todos Santos Formation sandstones from the Sierra de Chiapas (SE Mexico) and its tectonic significance. International Geology Review 2016; 58: 32 - 48. https: // doi. org / 10.108 0 / 00206814.2015.1055596" type="journal article" year="2016">
Abdullin
<emphasis id="00AB1769320AFFC6FB01FCE378CB561D" box="[1176,1226,802,826]" italics="true" pageId="9">et al.</emphasis>
2016
</bibRefCitation>
) and potentially explains the distinctiveness of the Upper Grijalva fish community (
<bibRefCitation id="564EB68A320AFFC6FCFCFCA17809565E" author="Elias DJ &amp; McMahan CD &amp; Matamoros WA" box="[869,1032,865,889]" pageId="9" pagination="2483 - 501" refId="ref16098" refString="Elias DJ, McMahan CD, Matamoros WA et al. Scale (s) matter: deconstructing an area of endemism for Middle American freshwater fishes. Journal of Biogeography 2020; 47: 2483 - 501. https: // doi. org / 10.1111 / jbi. 13941" type="journal article" year="2020">
Elías
<emphasis id="00AB1769320AFFC6FC04FCA17FCF565E" box="[925,974,865,889]" italics="true" pageId="9">et al.</emphasis>
2020
</bibRefCitation>
). Divergence of
<taxonomicName id="F5DFB0F8320AFFC6FB21FCA178F7565E" authorityName="Alvarez and Carranza" authorityYear="1951" box="[1208,1270,865,889]" family="Profundulidae" genus="Tlaloc" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FB21FCA178F7565E" box="[1208,1270,865,889]" italics="true" pageId="9">Tlaloc</emphasis>
</taxonomicName>
from
<taxonomicName id="F5DFB0F8320AFFC6FAAEFCA179AF565E" authorityName="Hubbs" authorityYear="1924" box="[1335,1454,865,889]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FAAEFCA179AF565E" box="[1335,1454,865,889]" italics="true" pageId="9">Profundulus</emphasis>
</taxonomicName>
, which probably arose in an ancestral Río Coatzacoalcos (
<figureCitation id="AAE4D7FE320AFFC6FAF3FC4079A356BF" box="[1386,1442,896,920]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="9">Fig. 8</figureCitation>
), provides an estimate for timing of this capture (
<tableCitation id="7F5DFEC0320AFFC6FA94FC5F79565690" box="[1293,1367,927,951]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="9">Table 2</tableCitation>
).
</paragraph>
<paragraph id="3260CB7B320AFFC6FCDCFC7F7FF25008" blockId="9.[809,1461,489,1327]" pageId="9">
Notably,
<taxonomicName id="F5DFB0F8320AFFC6FC3AFC7F781B56F0" authorityName="Hubbs" authorityYear="1924" box="[931,1050,959,983]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FC3AFC7F781B56F0" box="[931,1050,959,983]" italics="true" pageId="9">Profundulus</emphasis>
</taxonomicName>
is now largely restricted to Pacific slope drainages (
<bibRefCitation id="564EB68A320AFFC6FC39FC1E78F956D1" author="Dominguez-Cisneros S &amp; Dominguez-Dominguez O &amp; Velazquez-Velazquez E" box="[928,1272,990,1014]" pageId="9" refId="ref15657" refString="Dominguez-Cisneros S, Dominguez-Dominguez O, Velazquez-Velazquez E et al. Redescription and diagnoses of the genera Profundulus and Tlaloc (Cyprinodontiformes: Profundulidae), Mesoamerican endemic fishes. Neotropical Ichthyology 2023; 21: e 220089. https: // doi. org / 10.1590 / 1982 - 0224 - 2022 - 0089" type="journal volume" year="2023">
Domínguez-Cisneros
<emphasis id="00AB1769320AFFC6FB1FFC1F78BB56D1" box="[1158,1210,990,1014]" italics="true" pageId="9">et al.</emphasis>
2023
</bibRefCitation>
). Presuming that Miocene, upland
<taxonomicName id="F5DFB0F8320AFFC6FC40FC3D78515132" authorityName="Hubbs" authorityYear="1924" box="[985,1104,1021,1045]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FC40FC3D78515132" box="[985,1104,1021,1045]" italics="true" pageId="9">Profundulus</emphasis>
</taxonomicName>
no longer tolerated high-salinity or coastal environments, rising sea levels could explain the rarity of
<taxonomicName id="F5DFB0F8320AFFC6FCB3FBFC7FA05173" authorityName="Hubbs" authorityYear="1924" box="[810,929,1084,1108]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FCB3FBFC7FA05173" box="[810,929,1084,1108]" italics="true" pageId="9">Profundulus</emphasis>
</taxonomicName>
on the Gulf slope, as proposed for
<taxonomicName id="F5DFB0F8320AFFC6FA97FBFC7FC95154" authority="Baird &amp; Girard, 1854" authorityName="Baird &amp; Girard" authorityYear="1854" family="Cichlidae" genus="Herichthys" kingdom="Animalia" order="Perciformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FA97FBFC79775173" box="[1294,1398,1084,1108]" italics="true" pageId="9">Herichthys</emphasis>
Baird &amp; Girard, 1854
</taxonomicName>
(
<bibRefCitation id="564EB68A320AFFC6FC4EFB9B78DA5154" author="Perez-Miranda F &amp; Mejia O &amp; Lopez B" box="[983,1243,1115,1139]" pageId="9" refId="ref19005" refString="Perez-Miranda F, Mejia O, Lopez B et al. Molecular clocks, biogeography and species diversity in Herichthys with evaluation of the role of Punta del Morro as a vicariant brake along the Mexican Transition Zone in the context of local and global time frame of cichlid diversification. PeerJ 2020; 8: e 8818. https: // doi. org / 10.7717 / peerj. 8818" type="journal volume" year="2020">
Pérez-Miranda
<emphasis id="00AB1769320AFFC6FBEAFB9C78A25154" box="[1139,1187,1115,1139]" italics="true" pageId="9">et al.</emphasis>
2020
</bibRefCitation>
). During the Middle Miocene Climatic Optimum (MMCO) 17.0–13.8 Mya (
<bibRefCitation id="564EB68A320AFFC6FAEDFBBB7FA05195" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" pageId="9" pagination="32 - 41" refId="ref18302" refString="Miller KG, Schmelz WJ, Browning JV et al. Ancient sea level as key to the future. Oceanography 2020 b; 33: 32 - 41. https: // doi. org / 10.5670 / oceanog. 2020.224" type="journal article" year="2020">
Miller
<emphasis id="00AB1769320AFFC6FCB3FB5B7F5A5195" box="[810,859,1178,1202]" italics="true" pageId="9">et al.</emphasis>
2020b
</bibRefCitation>
), seas inundated broad areas north and east of the Sierra Madre del Sur and
<collectingRegion id="F01B0599320AFFC6FBA0FB79788D51F6" box="[1081,1164,1209,1233]" country="Mexico" name="Chiapas" pageId="9">Chiapas</collectingRegion>
Massif (
<bibRefCitation id="564EB68A320AFFC6FB73FB797F5F51D6" author="Blakey RC &amp; Ranney WD" pageId="9" refId="ref14773" refString="Blakey RC, Ranney WD. Ancient Landscapes of Western North America: A Geologic History with Paleogeographic Maps. Cham: Springer, 2018." type="book" year="2018">Blakey and Ranney 2018</bibRefCitation>
). Our chronogram indicates that diversification within
<taxonomicName id="F5DFB0F8320AFFC6FCB3FB387FA05037" authorityName="Hubbs" authorityYear="1924" box="[810,929,1272,1296]" family="Profundulidae" genus="Profundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320AFFC6FCB3FB387FA05037" box="[810,929,1272,1296]" italics="true" pageId="9">Profundulus</emphasis>
</taxonomicName>
occurred thereafter (
<figureCitation id="AAE4D7FE320AFFC6FB1DFB3878C15037" box="[1156,1216,1272,1296]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="9">Fig. 3</figureCitation>
), potentially from one refugial population.
</paragraph>
</subSubSection>
</treatment>
</document>

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<taxonomicName id="F5DFB0F8320CFFC0FE18FD1C7DEA57D1" ID-CoL="623TQ" box="[385,491,732,758]" family="Fundulidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="family">
<emphasis id="00AB1769320CFFC0FE18FD1C7DEA57D1" box="[385,491,732,758]" italics="true" pageId="15">Fundulidae</emphasis>
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<paragraph id="3260CB7B320CFFC0FFE8FCC37CCC5049" blockId="15.[113,764,771,1985]" pageId="15">
As described above (Early branching of major lineages), Middle Eocene fundulids theoretically inhabited the Brazos River delta and flooded
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Embayment (
<figureCitation id="AAE4D7FE320CFFC0FD97FC817E4D567E" box="[526,588,833,857]" captionStart="Figure 6" captionStartId="11.[113,178,1389,1413]" captionTargetBox="[114,1458,144,1361]" captionTargetId="figure-132@11.[114,1458,144,1361]" captionTargetPageId="11" captionText="Figure 6. Early–Middle Eocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Eocene lineage are labelled (Cypr. = Cyprinodontidae, Good.-Profund. = Goodeidae–Profundulidae).Phylogenetic nodes (Fig.2) are indicated as N1–N3 (node 1–node 3; for details, see main text and Table 2). The base map is a reconstruction for 50 Mya (late Early Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The dashed line approximates the shore of an inland sea that is hypothesized to have formed 55.8–55.0 Mya (Early Eocene), at which time land would have surrounded the entire Gulf, separating it from the world ocean (Cossey et al. 2016), theoretically causing vicariance between northern and southern clades of North American killifishes (N1). Superimposed drainages and estuaries are adapted from the work of Snedden and Galloway (2019). Details on southwestern Gulf tributaries are from studies by Snedden et al. (2018) and Cossey et al. (2021). Alternate routes to the Gulf are shown for the Madrean River, a Tamaulipas route (Early Eocene), and a Río Bravo route (Middle Eocene). The ancestor of Goodeidae is hypothesized to have populated this drainage via the Early Eocene Tamaulipas route.High sea levels of the Early Eocene presumably allowed widespread dispersal within northern and southern regions, whereas sea-level falls in the early Middle Eocene potentially separated Cyprinodontidae from Fundulidae in the north (N2) and Cubanichthyidae from Goodeidae–Profundulidae in the south (N3)." pageId="15">Fig. 6</figureCitation>
). Late Eocene– Oligocene divergence of
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<emphasis id="00AB1769320CFFC0FEE4FCA17DFA565E" box="[381,507,865,889]" italics="true" pageId="15">Leptolucania</emphasis>
Myers, 1924
</taxonomicName>
39.4–22.5 Mya (
<figureCitation id="AAE4D7FE320CFFC0FF29FC407CEB56BF" box="[176,234,896,920]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
, node 12) suggests eastward immigration of an ancestral fundulid across the Suwannee Channel to the Ocala High (
<tableCitation id="7F5DFEC0320CFFC0FFE5FC7F7CCB56F0" box="[124,202,959,983]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="15">Table 2</tableCitation>
;
<figureCitation id="AAE4D7FE320CFFC0FF40FC7F7D1556F1" box="[217,276,959,983]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="15">Fig. 7</figureCitation>
), which developed peritidal landforms by this time (Avon Park and Ocala formations;
<bibRefCitation id="564EB68A320CFFC0FDBAFC1E7CA45132" author="Randazzo AF &amp; Jones DS" pageId="15" pagination="39 - 56" refId="ref19538" refString="Randazzo AF, Jones DS. The sedimentary platform of Florida: Mesozoic to Cenozoic. In: Randazzo AF, Jones DS (eds), The Geology of Florida. Gainesville: University Press of Florida, 1997, 39 - 56." type="book chapter" year="1997">Randazzo and Jones 1997</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FF2BFC3D7D685132" author="Maliva RG &amp; Budd DA &amp; Clayton EA" box="[178,361,1021,1045]" pageId="15" pagination="218 - 32" refId="ref17788" refString="Maliva RG, Budd DA, Clayton EA et al. Insights into the dolomitization process and porosity modification in sucrosic dolostones, Avon Park Formation (Middle Eocene), East-Central Florida, U. S. A. Journal of Sedimentary Research 2011; 81: 218 - 32. https: // doi. org / 10.2110 / jsr. 2011.18" type="journal article" year="2011">
Maliva
<emphasis id="00AB1769320CFFC0FF67FC3E7D2E5132" box="[254,303,1021,1045]" italics="true" pageId="15">et al.</emphasis>
2011
</bibRefCitation>
). During the Oligocene, any of several sea-level fluctuations could have facilitated lineage separation if a sea-level fall (detected as an oxygen isotope event) facilitated dispersal across the Suwannee Channel to the Ocala High, with subsequent sea-level rise causing vicariance of the Ocala population owing to widening and deepening of the ocean gap (as proposed above for
<taxonomicName id="F5DFB0F8320CFFC0FEB9FB7A7D7D51F6" authorityName="Goode &amp; Bean" authorityYear="1879" box="[288,380,1209,1233]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="species" species="floridae">
<emphasis id="00AB1769320CFFC0FEB9FB7A7D7D51F6" box="[288,380,1209,1233]" italics="true" pageId="15">J. floridae</emphasis>
</taxonomicName>
). The Oi2 event ~30.0 Mya (
<bibRefCitation id="564EB68A320CFFC0FD28FB797CE051D6" author="Boulila S &amp; Galbrun B &amp; Miller KG" pageId="15" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320CFFC0FFE8FB197C9F51D6" box="[113,158,1241,1265]" italics="true" pageId="15">et al</emphasis>
. 2011
</bibRefCitation>
) appears best aligned in time with our divergence estimate for
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<emphasis id="00AB1769320CFFC0FF61FB387D775037" box="[248,374,1272,1296]" italics="true" pageId="15">Leptolucania</emphasis>
</taxonomicName>
. Although modern
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<emphasis id="00AB1769320CFFC0FDD4FB387ECA5037" box="[589,715,1272,1296]" italics="true" pageId="15">Leptolucania</emphasis>
</taxonomicName>
is a freshwater genus, the ancestor is likely to have had high salinity tolerance (
<bibRefCitation id="564EB68A320CFFC0FF7BFAF77DED5068" author="Ghedotti MJ &amp; Davis MP" box="[226,492,1335,1359]" pageId="15" pagination="1 - 65" refId="ref16579" refString="Ghedotti MJ, Davis MP. Phylogeny, classification, and evolution of salinity tolerance of the North American topminnows and killifishes, family Fundulidae (Teleostei: Cyprinodontiformes). Fieldiana Life and Earth Sciences 2013; 7: 1 - 65. https: // doi. org / 10.3158 / 2158 - 5520 - 12.7.1" type="journal article" year="2013">Ghedotti and Davis 2013</bibRefCitation>
), consistent with this hypothesis.
</paragraph>
<paragraph id="3260CB7B320CFFC0FF14FAB578FB55C0" blockId="15.[113,764,771,1985]" lastBlockId="15.[809,1461,144,1986]" pageId="15">
Our phylogeny places
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<emphasis id="00AB1769320CFFC0FEEFFAB67DC450AA" box="[374,453,1398,1421]" italics="true" pageId="15">Lucania</emphasis>
</taxonomicName>
within
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<emphasis id="00AB1769320CFFC0FD8EFAB57E7050AA" box="[535,625,1397,1421]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
, on an earlydiverging branch that includes
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<emphasis id="00AB1769320CFFC0FE2FFA557E11508A" box="[438,528,1429,1453]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
subgenus
<taxonomicName id="F5DFB0F8320CFFC0FD19FA557EFB508A" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[640,762,1429,1453]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FD19FA557EFB508A" box="[640,762,1429,1453]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
(
<figureCitation id="AAE4D7FE320CFFC0FFE5FA747CB850EB" box="[124,185,1460,1484]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
). This is contrary to the work of
<bibRefCitation id="564EB68A320CFFC0FDB0FA747CBD50CC" author="Ghedotti MJ &amp; Davis MP" pageId="15" pagination="577 - 86" refId="ref16637" refString="Ghedotti MJ, Davis MP. The taxonomic placement of three fossil Fundulus species and the timing of divergence within the North American topminnows (Teleostei: Fundulidae). Zootaxa 2017; 4250: 577 - 86. https: // doi. org / 10.11646 / zootaxa. 4250.6.5" type="journal article" year="2017">Ghedotti and Davis (2017)</bibRefCitation>
, who recovered
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<emphasis id="00AB1769320CFFC0FEFEFA147DB750CC" box="[359,438,1492,1515]" italics="true" pageId="15">Lucania</emphasis>
</taxonomicName>
outside
<taxonomicName id="F5DFB0F8320CFFC0FD89FA137E6850CC" authorityName="Lacepede" authorityYear="1803" box="[528,617,1491,1515]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FD89FA137E6850CC" box="[528,617,1491,1515]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
, but in partial agreement with the fish tree of life (
<bibRefCitation id="564EB68A320CFFC0FE46FA337EA4532C" author="Rabosky DL &amp; Chang J &amp; Title PO" box="[479,677,1523,1547]" pageId="15" pagination="392 - 5" refId="ref19373" refString="Rabosky DL, Chang J, Title PO et al. An inverse latitudinal gradient in speciation rate for marine fishes. Nature 2018; 559: 392 - 5. https: // doi. org / 10.1038 / s 41586 - 018 - 0273 - 1" type="journal article" year="2018">
Rabosky
<emphasis id="00AB1769320CFFC0FDA4FA337E67532C" box="[573,614,1523,1547]" italics="true" pageId="15">et al</emphasis>
. 2018
</bibRefCitation>
), which places
<taxonomicName id="F5DFB0F8320CFFC0FF2CF9D37D05530D" authorityName="Girard" authorityYear="1860" box="[181,260,1555,1578]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FF2CF9D37D05530D" box="[181,260,1555,1578]" italics="true" pageId="15">Lucania</emphasis>
</taxonomicName>
inside
<taxonomicName id="F5DFB0F8320CFFC0FED4F9D27DA6530D" authorityName="Lacepede" authorityYear="1803" box="[333,423,1554,1578]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FED4F9D27DA6530D" box="[333,423,1554,1578]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
, albeit on a separate branch from
<taxonomicName id="F5DFB0F8320CFFC0FFEBF9F17CED536E" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[114,236,1585,1609]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FFEBF9F17CED536E" box="[114,236,1585,1609]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
(this tree did not include
<taxonomicName id="F5DFB0F8320CFFC0FE63F9F17E76536E" authorityName="Myers" authorityYear="1924" box="[506,631,1585,1609]" family="Fundulidae" genus="Leptolucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FE63F9F17E76536E" box="[506,631,1585,1609]" italics="true" pageId="15">Leptolucania</emphasis>
</taxonomicName>
). Given that
<taxonomicName id="F5DFB0F8320CFFC0FFEBF9917CED534E" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[114,236,1617,1641]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FFEBF9917CED534E" box="[114,236,1617,1641]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
includes only the Pacific Coast taxa
<taxonomicName id="F5DFB0F8320CFFC0FDF4F9917EFB534E" authorityName="Vaillant" authorityYear="1894" box="[621,762,1617,1641]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="species" species="lima">
<emphasis id="00AB1769320CFFC0FDF4F9917EFB534E" box="[621,762,1617,1641]" italics="true" pageId="15">Fundulus lima</emphasis>
</taxonomicName>
and
<taxonomicName id="F5DFB0F8320CFFC0FF07F9B07D6D53AF" authorityName="Girard" authorityYear="1856" box="[158,364,1648,1672]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="species" species="parvipinnis">
<emphasis id="00AB1769320CFFC0FF07F9B07D6D53AF" box="[158,364,1648,1672]" italics="true" pageId="15">Fundulus parvipinnis</emphasis>
</taxonomicName>
and given that the Coahuilan endemic
<taxonomicName id="F5DFB0F8320CFFC0FFEBF9507D225380" authorityName="Hubbs &amp; Miller" authorityYear="1965" box="[114,291,1680,1703]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="species" species="interioris">
<emphasis id="00AB1769320CFFC0FFEBF9507D225380" box="[114,291,1680,1703]" italics="true" pageId="15">Lucania interioris</emphasis>
</taxonomicName>
is the earliest diverging lineage of
<taxonomicName id="F5DFB0F8320CFFC0FD3CF9507EF55380" authorityName="Girard" authorityYear="1860" box="[677,756,1680,1703]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FD3CF9507EF55380" box="[677,756,1680,1703]" italics="true" pageId="15">Lucania</emphasis>
</taxonomicName>
, we propose (following
<figureCitation id="AAE4D7FE320CFFC0FEF6F96F7DAF53E0" box="[367,430,1711,1735]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
) that the MRCA of
<emphasis id="00AB1769320CFFC0FD04F96F7CEA53C1" italics="true" pageId="15">
<taxonomicName id="F5DFB0F8320CFFC0FD04F96F7EEC53E1" authorityName="Girard" authorityYear="1860" box="[669,749,1711,1734]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Lucania</taxonomicName>
–
<taxonomicName id="F5DFB0F8320CFFC0FFE8F90E7CEA53C1" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[113,235,1742,1766]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
</emphasis>
originated from range expansion of fundulids westwards from the Brazos delta to the Río Grande-Río Bravo delta 31.3–19.1 Mya (
<figureCitation id="AAE4D7FE320CFFC0FE85F8CD7D545202" box="[284,341,1805,1829]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
, node 13;
<figureCitation id="AAE4D7FE320CFFC0FE5BF8CD7DFB5203" box="[450,506,1805,1829]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="15">Fig. 7</figureCitation>
). During this time, there were sea-level falls of
<quantity id="F527669E320CFFC0FED7F8EC7DAA5264" box="[334,427,1836,1860]" metricMagnitude="1" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="15" unit="m" value="35.0" valueMax="40.0" valueMin="30.0">30–40 m</quantity>
below modern sea level approximately every 1.2 Mya (
<bibRefCitation id="564EB68A320CFFC0FECFF88B7E085244" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[342,521,1867,1891]" pageId="15" pagination="32 - 41" refId="ref18302" refString="Miller KG, Schmelz WJ, Browning JV et al. Ancient sea level as key to the future. Oceanography 2020 b; 33: 32 - 41. https: // doi. org / 10.5670 / oceanog. 2020.224" type="journal article" year="2020">
Miller
<emphasis id="00AB1769320CFFC0FE00F88C7DC75244" box="[409,454,1867,1891]" italics="true" pageId="15">et al.</emphasis>
2020b
</bibRefCitation>
). Following
<bibRefCitation id="564EB68A320CFFC0FD11F88B7CB052A4" author="Dolby GA &amp; Hechinger R &amp; Ellingson RA" pageId="15" refId="ref15608" refString="Dolby GA, Hechinger R, Ellingson RA et al. Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study. Proceedings Biological Sciences 2016; 283: 20161571. https: // doi. org / 10.1098 / rspb. 2016.1571" type="journal volume" year="2016">
Dolby
<emphasis id="00AB1769320CFFC0FD57F88C7EFA5244" box="[718,763,1867,1891]" italics="true" pageId="15">et al.</emphasis>
(2016
</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FF25F8AB7CF152A4" author="Dolby GA &amp; Ellingson RA &amp; Findley LT" box="[188,240,1899,1923]" pageId="15" pagination="994 - 1011" refId="ref15556" refString="Dolby GA, Ellingson RA, Findley LT et al. How sea level change mediates genetic divergence in coastal species across regions with varying tectonic and sediment processes. Molecular Ecology 2018; 27: 994 - 1011. https: // doi. org / 10.1111 / mec. 14487" type="journal article" year="2018">2018</bibRefCitation>
), we postulate that sea-level fluctuations promoted cycles of dispersal and vicariance. Any one or a combination of recessions might account for vicariance of fundulids between the Brazos and Río Grande-Río Bravo deltas, but the Oi2c event dated ~25.1 Mya (
<bibRefCitation id="564EB68A320CFFC0FC77FF6F78AA55E0" author="Boulila S &amp; Galbrun B &amp; Miller KG" box="[1006,1195,175,199]" pageId="15" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320CFFC0FBA6FF70787155E0" box="[1087,1136,175,199]" italics="true" pageId="15">et al.</emphasis>
2011
</bibRefCitation>
) best aligns in time with divergence of
<emphasis id="00AB1769320CFFC0FC21FF0F788E55C0" box="[952,1167,207,231]" italics="true" pageId="15">
<taxonomicName id="F5DFB0F8320CFFC0FC21FF0F780955C0" authorityName="Girard" authorityYear="1860" box="[952,1032,207,231]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Lucania</taxonomicName>
–
<taxonomicName id="F5DFB0F8320CFFC0FB8CFF0F788E55C0" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[1045,1167,207,231]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
</emphasis>
(
<tableCitation id="7F5DFEC0320CFFC0FB07FF0F78E855C0" box="[1182,1257,207,231]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="15">Table 2</tableCitation>
).
</paragraph>
<paragraph id="3260CB7B320CFFC0FCDCFF2E78ED5112" blockId="15.[809,1461,144,1986]" pageId="15">
In the Oligocene, the common ancestor of
<emphasis id="00AB1769320CFFC0FACFFF2F7FA55402" italics="true" pageId="15">
<taxonomicName id="F5DFB0F8320CFFC0FACFFF2F79A75421" authorityName="Girard" authorityYear="1860" box="[1366,1446,239,262]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Lucania</taxonomicName>
–
<taxonomicName id="F5DFB0F8320CFFC0FCB3FECD7FA55402" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[810,932,269,293]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Wileyichthys</taxonomicName>
</emphasis>
presumably immigrated into the ancestral Río Grande drainage. This massive river system extended far across southern North America to what would become the Basin and Range geomorphic province (
<bibRefCitation id="564EB68A320CFFC0FBC3FEAB797C54A4" author="Snedden JW &amp; Galloway WE" box="[1114,1405,363,387]" pageId="15" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
) and appears to have been the only Cenozoic route to the southern Basin and Range from the western Gulf of
<collectingRegion id="F01B0599320CFFC0FB45FE6A792854E5" box="[1244,1321,426,450]" country="Mexico" name="Mexico" pageId="15">México</collectingRegion>
(
<figureCitation id="AAE4D7FE320CFFC0FAA1FE6A797054E6" box="[1336,1393,426,450]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="15">Fig. 7</figureCitation>
). This geographical scenario comes with relatively strict time constraints. Divergence of
<taxonomicName id="F5DFB0F8320CFFC0FB8EFE2978905726" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[1047,1169,489,513]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB8EFE2978905726" box="[1047,1169,489,513]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
from
<taxonomicName id="F5DFB0F8320CFFC0FB49FE29791E5727" authorityName="Girard" authorityYear="1860" box="[1232,1311,489,512]" family="Fundulidae" genus="Lucania" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB49FE29791E5727" box="[1232,1311,489,512]" italics="true" pageId="15">Lucania</emphasis>
</taxonomicName>
must have occurred when Basin and Range drainages became disconnected from the Río Grande, dated ~23.0 Mya (
<bibRefCitation id="564EB68A320CFFC0FB51FDE77F5F5778" author="Snedden JW &amp; Galloway WE" pageId="15" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
). This timing is compatible with our 30.7–7.5 Mya estimate for divergence of
<taxonomicName id="F5DFB0F8320CFFC0FB85FDA678975759" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[1052,1174,614,638]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB85FDA678975759" box="[1052,1174,614,638]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
(
<figureCitation id="AAE4D7FE320CFFC0FB31FDA678E25759" box="[1192,1251,614,638]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
, node 14;
<tableCitation id="7F5DFEC0320CFFC0FACCFDA679A35759" box="[1365,1442,614,638]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="15">Table 2</tableCitation>
). Fossil
<taxonomicName id="F5DFB0F8320CFFC0FCF5FD457FC757BA" authorityName="Lacepede" authorityYear="1803" box="[876,966,645,669]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FCF5FD457FC757BA" box="[876,966,645,669]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
in the Great Basin (
<bibRefCitation id="564EB68A320CFFC0FB01FD45794757BA" author="Smith GR &amp; Dowling T &amp; Gobalet KW" box="[1176,1350,645,669]" pageId="15" pagination="175 - 234" refId="ref20339" refString="Smith GR, Dowling T, Gobalet KW et al. 2002. Great Basin aquatic systems history. Smithsonian Contributions to Earth Sciences 2002; 33: 175 - 234." type="journal article" year="2002">
Smith
<emphasis id="00AB1769320CFFC0FB45FD46790D57BA" box="[1244,1292,645,669]" italics="true" pageId="15">et al.</emphasis>
2002
</bibRefCitation>
) also support this hypothesis. Finally, distribution of
<taxonomicName id="F5DFB0F8320CFFC0FB68FD65796A579A" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[1265,1387,677,701]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB68FD65796A579A" box="[1265,1387,677,701]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
on the Pacific Coast of southern
<collectingRegion id="F01B0599320CFFC0FBA4FD0478A557FB" box="[1085,1188,708,732]" country="United States of America" name="California" pageId="15">California</collectingRegion>
and south along the Baja Peninsula agrees with the Río Grande as the dispersal corridor, because the headwaters of the Oligocene Río Grande extended to the Continental Divide in what would become southern
<collectingRegion id="F01B0599320CFFC0FCB3FC817F90567E" box="[810,913,833,857]" country="United States of America" name="California" pageId="15">California</collectingRegion>
(
<bibRefCitation id="564EB68A320CFFC0FC3BFC817878567E" author="Karlstrom KE &amp; Jacobson CE &amp; Sundell KE" box="[930,1145,833,857]" pageId="15" pagination="1425 - 56" refId="ref17499" refString="Karlstrom KE, Jacobson CE, Sundell KE et al. Evaluating the Shinumo- Sespe drainage connection: Arguments against the &quot; old &quot; (70 - 17 Ma) Grand Canyon models for Colorado Plateau drainage evolution. Geosphere 2020; 16: 1425 - 56." type="journal article" year="2020">
Karlstrom
<emphasis id="00AB1769320CFFC0FB89FC827841567E" box="[1040,1088,833,857]" italics="true" pageId="15">et al.</emphasis>
2020
</bibRefCitation>
). Tectonism in this region potentially resulted in stream transfers that introduced
<taxonomicName id="F5DFB0F8320CFFC0FAA1FCA179B3565E" authorityName="Ghedotti &amp; Davis" authorityYear="2013" box="[1336,1458,865,889]" family="Fundulidae" genus="Wileyichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FAA1FCA179B3565E" box="[1336,1458,865,889]" italics="true" pageId="15">Wileyichthys</emphasis>
</taxonomicName>
to the Pacific slope. Likewise, Empetrichthyinae must have been living in Basin and Range drainages along the continental divide, after break-up of the Madrean River (above). Fossil evidence from Empetrichthyinae (
<bibRefCitation id="564EB68A320CFFC0FBB2FC1E791B56D1" author="Uyeno T &amp; Miller RR" box="[1067,1306,990,1014]" pageId="15" pagination="520 - 32" refId="ref20623" refString="Uyeno T, Miller RR. Relationships of Empetrichthys erdisi, a Pliocene cyprinodontid fish from California, with remarks on the Fundulinae and Cyprinodontinae. Copeia 1962; 1962: 520 - 32. https: // doi. org / 10.2307 / 1441173" type="journal article" year="1962">Uyeno and Miller 1962</bibRefCitation>
) indicates that this group was also transferred to the Pacific slope, although no representatives survived to the present time.
</paragraph>
<paragraph id="3260CB7B320CFFC0FCDCFBFC7FCA530E" blockId="15.[809,1461,144,1986]" pageId="15">
The major split within the
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<emphasis id="00AB1769320CFFC0FBFBFBFC78BD5173" box="[1122,1212,1084,1108]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
crown group separated subgenus
<taxonomicName id="F5DFB0F8320CFFC0FC08FB9B7FEA5154" authorityName="Lacepede" authorityYear="1803" box="[913,1003,1115,1139]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FC08FB9B7FEA5154" box="[913,1003,1115,1139]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
as a clade 24.2–17.1 Mya (
<figureCitation id="AAE4D7FE320CFFC0FA97FB9B79495154" box="[1294,1352,1115,1139]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
, node 15;
<tableCitation id="7F5DFEC0320CFFC0FCB3FBBB7F7451B4" box="[810,885,1147,1171]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="15">Table 2</tableCitation>
). An eastward shift of the Oligocene Brazos River at the Oligocene–Miocene transition (23 Mya) from its existing delta to form a new Red River delta relatively close to the
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River delta (present Sabine River delta;
<bibRefCitation id="564EB68A320CFFC0FB4BFB1979A951D6" author="Galloway WE &amp; Whiteaker TL &amp; Ganey-Curry P" box="[1234,1448,1241,1265]" pageId="15" pagination="938 - 73" refId="ref16432" refString="Galloway WE, Whiteaker TL, Ganey-Curry P. History of Cenozoic North American drainage basin evolution, sediment yield, and accumulation in the Gulf of Mexico basin. Geosphere 2011; 7: 938 - 73. https: // doi. org / 10.1130 / ges 00647.1" type="journal article" year="2011">
Galloway
<emphasis id="00AB1769320CFFC0FAA3FB19796751D6" box="[1338,1382,1241,1265]" italics="true" pageId="15">et al</emphasis>
. 2011
</bibRefCitation>
) might have provided a catalyst for this event by providing an uninhabited delta open to immigrants. During this time, there were sea-level falls of
<quantity id="F527669E320CFFC0FC54FAF778285069" box="[973,1065,1335,1359]" metricMagnitude="1" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="15" unit="m" value="35.0" valueMax="40.0" valueMin="30.0">30–40 m</quantity>
below modern sea level approximately every 1.2 Mya (
<bibRefCitation id="564EB68A320CFFC0FC56FA96788B5049" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[975,1162,1366,1390]" pageId="15" pagination="32 - 41" refId="ref18302" refString="Miller KG, Schmelz WJ, Browning JV et al. Ancient sea level as key to the future. Oceanography 2020 b; 33: 32 - 41. https: // doi. org / 10.5670 / oceanog. 2020.224" type="journal article" year="2020">
Miller
<emphasis id="00AB1769320CFFC0FB8DFA9778445049" box="[1044,1093,1366,1390]" italics="true" pageId="15">et al.</emphasis>
2020b
</bibRefCitation>
). Any one or a combination of these fluctuations might account for a dispersal–vicariance sequence in fundulids between the
<collectingRegion id="F01B0599320CFFC0FB39FA557913508A" box="[1184,1298,1429,1453]" country="United States of America" name="Mississippi" pageId="15">Mississippi</collectingRegion>
River and Red River deltas (
<bibRefCitation id="564EB68A320CFFC0FC2FFA74786950EB" author="Dolby GA &amp; Hechinger R &amp; Ellingson RA" box="[950,1128,1460,1484]" pageId="15" refId="ref15608" refString="Dolby GA, Hechinger R, Ellingson RA et al. Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study. Proceedings Biological Sciences 2016; 283: 20161571. https: // doi. org / 10.1098 / rspb. 2016.1571" type="journal volume" year="2016">
Dolby
<emphasis id="00AB1769320CFFC0FC64FA75782650EB" box="[1021,1063,1460,1484]" italics="true" pageId="15">et al</emphasis>
. 2016
</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FBEDFA7478A950EB" author="Dolby GA &amp; Ellingson RA &amp; Findley LT" box="[1140,1192,1460,1484]" pageId="15" pagination="994 - 1011" refId="ref15556" refString="Dolby GA, Ellingson RA, Findley LT et al. How sea level change mediates genetic divergence in coastal species across regions with varying tectonic and sediment processes. Molecular Ecology 2018; 27: 994 - 1011. https: // doi. org / 10.1111 / mec. 14487" type="journal article" year="2018">2018</bibRefCitation>
). However, the Mi1a isotope zone (sea-level recession) dated ~20.8 Mya (
<bibRefCitation id="564EB68A320CFFC0FAABFA137F5F532C" author="Boulila S &amp; Galbrun B &amp; Miller KG" pageId="15" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320CFFC0FA1AFA1479B550CC" box="[1411,1460,1491,1515]" italics="true" pageId="15">et al.</emphasis>
2011
</bibRefCitation>
) best aligns in time with our divergence estimate for subgenus
<taxonomicName id="F5DFB0F8320CFFC0FCF2F9D27FC4530D" authorityName="Lacepede" authorityYear="1803" box="[875,965,1554,1578]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FCF2F9D27FC4530D" box="[875,965,1554,1578]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="3260CB7B320CFFDFFCDCF9F27CC455C0" blockId="15.[809,1461,144,1986]" lastBlockId="16.[129,778,144,231]" lastPageId="16" pageId="15">
We hypothesize that the
<emphasis id="00AB1769320CFFC0FBC1F9F279B2536E" box="[1112,1459,1585,1609]" italics="true" pageId="15">
<taxonomicName id="F5DFB0F8320CFFC0FBC1F9F278C1536E" authorityName="Agassiz" authorityYear="1854" box="[1112,1216,1586,1609]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Zygonectes</taxonomicName>
–
<taxonomicName id="F5DFB0F8320CFFC0FB54F9F27940536E" authorityName="Cope" authorityYear="1865" box="[1229,1345,1585,1609]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="species" species="sciadicus">F. sciadicus</taxonomicName>
–
<taxonomicName id="F5DFB0F8320CFFC0FAD7F9F179B2536E" authorityName="Garman" authorityYear="1895" box="[1358,1459,1585,1609]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">Plancterus</taxonomicName>
</emphasis>
clade descends from a lineage that originated in the
<collectingRegion id="F01B0599320CFFC0FAD8F99179B2534E" box="[1345,1459,1617,1641]" country="United States of America" name="Mississippi" pageId="15">Mississippi</collectingRegion>
River delta, because
<taxonomicName id="F5DFB0F8320CFFC0FB9CF9B0786A53AF" authorityName="Garman" authorityYear="1895" box="[1029,1131,1648,1672]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB9CF9B0786A53AF" box="[1029,1131,1648,1672]" italics="true" pageId="15">Plancterus</emphasis>
</taxonomicName>
(node 18) has a Mississippian distribution and is the westernmost lineage on this branch (
<figureCitation id="AAE4D7FE320CFFC0FA14F94F7F3653E0" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="15">Fig. 8</figureCitation>
). By default, this implies that subgenus
<taxonomicName id="F5DFB0F8320CFFC0FB68F96F794A53E0" authorityName="Lacepede" authorityYear="1803" box="[1265,1355,1711,1735]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FB68F96F794A53E0" box="[1265,1355,1711,1735]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
descends from fundulids that immigrated into the
<collectingCounty id="DB01B3F7320CFFC0FB4DF90E793C53C1" box="[1236,1341,1742,1766]" pageId="15">Red River</collectingCounty>
delta, then diverged into a separate lineage during sea-level recession. Later abandonment of the
<collectingMunicipality id="D2045101320CFFC0FB95F8CD78765202" box="[1036,1143,1805,1829]" pageId="15">Red River</collectingMunicipality>
delta (
<figureCitation id="AAE4D7FE320CFFC0FB5EF8CD79025202" box="[1223,1283,1805,1829]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="15">Fig. 8</figureCitation>
), which merged with the
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River delta 15 Mya (
<bibRefCitation id="564EB68A320CFFC0FB50F8EC7F5F5244" author="Snedden JW &amp; Galloway WE" pageId="15" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
), mixed
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and
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River fishes (
<bibRefCitation id="564EB68A320CFFC0FAA6F88C7FDA52A4" author="Hoagstrom CW &amp; Echelle A" pageId="15" pagination="261 - 87" refId="ref17298" refString="Hoagstrom CW, Echelle A. Biogeography of the Macrhybopsis aestivalis complex (Teleostei: Cyprinidae): emphasis on speciation and ancient heterospecific mitochondrial transfer. Environmental Biology of Fishes 2022; 105: 261 - 87. https: // doi. org / 10.1007 / s 10641 - 022 - 01220 - 0" type="journal article" year="2022">Hoagstrom and Echelle 2022</bibRefCitation>
). This would have brought subgenus
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<emphasis id="00AB1769320CFFC0FAC0F8AB79B252A4" box="[1369,1459,1899,1923]" italics="true" pageId="15">Fundulus</emphasis>
</taxonomicName>
into sympatry with
<taxonomicName id="F5DFB0F8320CFFC0FC63F84B78625285" authorityName="Agassiz" authorityYear="1854" box="[1018,1123,1931,1954]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FC63F84B78625285" box="[1018,1123,1931,1954]" italics="true" pageId="15">Zygonectes</emphasis>
</taxonomicName>
. Thereafter, sea-level rise 19–17 Mya leading up to the MMCO 17.0–13.8 Mya, during which high sea levels were sustained (
<bibRefCitation id="564EB68A3213FFDFFE24FF507E75558F" author="Miller KG &amp; Browning JV &amp; Schmelz WJ" box="[445,628,144,168]" pageId="16" refId="ref18255" refString="Miller KG, Browning JV, Schmelz WJ et al. Cenozoic sea-level and cryospheric evolution from deep-sea geochemical and continental margin records. Science Advances 2020 a; 6: eaaz 1346. https: // doi. org / 10.1126 / sciadv. aaz 1346" type="journal volume" year="2020">
Miller
<emphasis id="00AB17693213FFDFFD98FF517E31558F" box="[513,560,144,168]" italics="true" pageId="16">et al.</emphasis>
2020a
</bibRefCitation>
), theoretically aided range expansion of all coastal fundulids (
<bibRefCitation id="564EB68A3213FFDFFDC0FF6F7F0555E0" author="Dolby GA &amp; Hechinger R &amp; Ellingson RA" box="[601,772,175,199]" pageId="16" refId="ref15608" refString="Dolby GA, Hechinger R, Ellingson RA et al. Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study. Proceedings Biological Sciences 2016; 283: 20161571. https: // doi. org / 10.1098 / rspb. 2016.1571" type="journal volume" year="2016">
Dolby
<emphasis id="00AB17693213FFDFFD07FF707ECD55E0" box="[670,716,175,199]" italics="true" pageId="16">et al.</emphasis>
2016
</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFF18FF0F7CB455C0" author="Dolby GA &amp; Ellingson RA &amp; Findley LT" box="[129,181,207,231]" pageId="16" pagination="994 - 1011" refId="ref15556" refString="Dolby GA, Ellingson RA, Findley LT et al. How sea level change mediates genetic divergence in coastal species across regions with varying tectonic and sediment processes. Molecular Ecology 2018; 27: 994 - 1011. https: // doi. org / 10.1111 / mec. 14487" type="journal article" year="2018">2018</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:namePart id="8D89332335C59E3483EAA629373ECA59">Matamoros, Wilfredo A.</mods:namePart>
<mods:affiliation id="67A6A9B896B4129C9201F941DFC42322">Programa de Licenciatura, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Libramiento Norte Poniente No. 1150, Colonia Lajas Maciel, C. P. 29039, Tuxtla Gutiérrez, Chiapas, México &amp; Maestría en Ciencias en Biodiversidad y Conservación de Ecosistemas Tropicales, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Tuxtla Gutiérrez, Chiapas, México</mods:affiliation>
<mods:nameIdentifier id="42400273EA502185062C5ACEE41C2094" type="email">wilmatamoros@yahoo.com</mods:nameIdentifier>
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<mods:typeOfResource id="6B87EBE966AD66609253A74DA7815886">text</mods:typeOfResource>
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<mods:title id="ED61E8133FCEAD1B28D1952D2F092B45">Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part id="DDAEA777435229809CCBDDAC944F7529">
<mods:date id="F5A8298CB919F09987167A76FFB8CE0B">2024</mods:date>
<mods:detail id="786E87A443F405CDC7226132E666518D" type="pubDate">
<mods:number id="31A597A7F09F6513E8D27C6A2F4C62A8">2024-10-26</mods:number>
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<mods:detail id="B78049FA55A047C325C08B911867BAEA" type="volume">
<mods:number id="AE3A78C0811DC714589FF5CE8C7B8871">202</mods:number>
</mods:detail>
<mods:detail id="A864C40CEA610BD4834B19A87942E9FB" type="issue">
<mods:number id="7BC83DC9D317D43D485BFF09102A056B">2</mods:number>
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<mods:location id="66D4776642859A044149D4413E054A8D">
<mods:url id="2EADFA860B390BF16A7949EE104B3113">https://doi.org/10.1093/zoolinnean/zlae105</mods:url>
</mods:location>
<mods:classification id="7271554D19F9DFA4F6B9B4FA30CFE5F6">journal volume</mods:classification>
<mods:identifier id="CC50AA02A2EC395DBC11D90C1B14E9C6" type="DOI">10.1093/zoolinnean/zlae105</mods:identifier>
<mods:identifier id="FD6A2127A04708DD13E1612A3094969D" type="ISSN">0024-4082</mods:identifier>
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<subSubSection id="7AC598F0320EFFC2FEFFFD9F7E07575E" box="[358,518,607,633]" pageId="13" type="nomenclature">
<paragraph id="3260CB7B320EFFC2FEFFFD9F7E07575E" blockId="13.[113,764,607,1985]" box="[358,518,607,633]" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FEFFFD9F7E07575E" ID-CoL="8WF" box="[358,518,607,633]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="family">
<emphasis id="00AB1769320EFFC2FEFFFD9F7E07575E" box="[358,518,607,633]" italics="true" pageId="13">Cyprinodontidae</emphasis>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="7AC598F0320EFFC0FFE8FD457DF3579A" lastPageId="15" pageId="13" type="discussion">
<paragraph id="3260CB7B320EFFC2FFE8FD457D9B508A" blockId="13.[113,764,607,1985]" pageId="13">
Divergence of the MRCA of
<emphasis id="00AB1769320EFFC2FE3BFD457E8A57BA" box="[418,651,645,669]" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FE3BFD457E1E57BA" authorityName="Hubbs" authorityYear="1926" box="[418,543,645,669]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F8320EFFC2FDB2FD457E8A57BA" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[555,651,645,669]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">J. pulchra</taxonomicName>
</emphasis>
42.3–21.6 Mya (
<figureCitation id="AAE4D7FE320EFFC2FF2AFD657CEF579A" box="[179,238,677,701]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
, node 8;
<tableCitation id="7F5DFEC0320EFFC2FEC1FD657DA7579A" box="[344,422,677,701]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="13">Table 2</tableCitation>
) is consistent with our hypothesis that
<taxonomicName id="F5DFB0F8320EFFC2FF57FD047D7E57FB" box="[206,383,708,732]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
originated in the northwestern Gulf (above). This divergence estimate spans a dramatic cooling event at the Eocene–Oligocene boundary 33.9 Mya, which initiated the Oligocene icehouse climate that lasted until ~26 Mya (
<bibRefCitation id="564EB68A320EFFC2FFE5FC817D43567E" author="Zachos J &amp; Pagani H &amp; Sloan L" box="[124,322,833,857]" pageId="13" pagination="686 - 93" refId="ref21340" refString="Zachos J, Pagani H, Sloan L et al. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 2001; 292: 686 - 93. https: // doi. org / 10.1126 / science. 1059412" type="journal article" year="2001">
Zachos
<emphasis id="00AB1769320EFFC2FF48FC827D04567E" box="[209,261,833,857]" italics="true" pageId="13">et al.</emphasis>
2001
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FECBFC817E16567E" author="Miller KG &amp; Schmelz WJ &amp; Browning JV" box="[338,535,833,857]" pageId="13" pagination="32 - 41" refId="ref18302" refString="Miller KG, Schmelz WJ, Browning JV et al. Ancient sea level as key to the future. Oceanography 2020 b; 33: 32 - 41. https: // doi. org / 10.5670 / oceanog. 2020.224" type="journal article" year="2020">
Miller
<emphasis id="00AB1769320EFFC2FE02FC827DCE567E" box="[411,463,833,857]" italics="true" pageId="13">et al.</emphasis>
2020b
</bibRefCitation>
). Sustained icehouse conditions possibly facilitated southward invasion of Nearctic
<taxonomicName id="F5DFB0F8320EFFC2FFE8FC407D2356BF" box="[113,290,896,920]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
(
<figureCitation id="AAE4D7FE320EFFC2FEA8FC407D6656B0" box="[305,359,896,920]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="13">Fig. 7</figureCitation>
), consistent with predictions of the intothe-tropics paradigm (
<bibRefCitation id="564EB68A320EFFC2FEC0FC5F7DC25690" author="Vrba ES" box="[345,451,927,951]" pageId="13" pagination="24 - 45" refId="ref20833" refString="Vrba ES. On the connections between paleoclimate and evolution. In: Vrba ES, Denton GH, Partridge TC, et al. (eds), Paleoclimate and Evolution, with Emphasis on Human Origins. New Haven: Yale University Press, 1995, 24 - 45." type="book chapter" year="1995">Vrba 1995</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FE56FC607E8B5690" author="Eronen JT &amp; Fortelius M &amp; Micheels A" box="[463,650,927,951]" pageId="13" pagination="823 - 6" refId="ref16200" refString="Eronen JT, Fortelius M, Micheels A et al. Neogene aridification of the Northern Hemisphere. Geology 2012; 40: 823 - 6. https: // doi. org / 10.1130 / g 33147.1" type="journal article" year="2012">
Eronen
<emphasis id="00AB1769320EFFC2FDB8FC607E505690" box="[545,593,927,951]" italics="true" pageId="13">et al.</emphasis>
2012
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FD0FFC607D4E56F0" author="Meseguer AS &amp; Condamine FL" pageId="13" pagination="1966 - 87" refId="ref18191" refString="Meseguer AS, Condamine FL. Ancient tropical extinctions at high latitudes contributed to the latitudinal diversity gradient. Evolution 2020; 74: 1966 - 87." type="journal article" year="2020">Meseguer and Condamine 2020</bibRefCitation>
) and with our ancestral-areas analysis that included the potential for jump dispersal between realms (
<figureCitation id="AAE4D7FE320EFFC2FD4DFC1E7C7F5132" captionStart="Figure 5" captionStartId="10.[129,194,1836,1860]" captionTargetBox="[212,1390,147,1805]" captionTargetId="figure-7@10.[209,1393,144,1808]" captionTargetPageId="10" captionText="Figure 5. Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (Rico et al.2022). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS." pageId="13">Fig. 5</figureCitation>
). The point estimate for this divergence (31.8 Mya) is aligned with Oligocene isotope zone 1b (abbreviated Oi1b), dated 31.8 Mya (
<bibRefCitation id="564EB68A320EFFC2FF29FBFC7D6B5173" author="Boulila S &amp; Galbrun B &amp; Miller KG" box="[176,362,1084,1108]" pageId="13" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320EFFC2FE99FBFD7D315173" box="[256,304,1084,1108]" italics="true" pageId="13">et al.</emphasis>
2011
</bibRefCitation>
). Oxygen isotope zones are periods of increased global glaciation (at which times, sea level is lowered), indicated by maximum relative abundance of δ
<superScript id="C5AA6633320EFFC2FDF8FBB97E7151A0" attach="right" box="[609,624,1145,1159]" fontSize="6" pageId="13">18</superScript>
O (
<bibRefCitation id="564EB68A320EFFC2FD0EFBBB7CC25195" author="Boulila S &amp; Galbrun B &amp; Miller KG" pageId="13" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320EFFC2FD70FBBB7C8B5195" italics="true" pageId="13">et al.</emphasis>
2011
</bibRefCitation>
). Concurrent timing of sea-level fall with divergence of the
<emphasis id="00AB1769320EFFC2FF03FB797D8E51F6" box="[154,399,1209,1233]" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FF03FB797D1951F6" authorityName="Hubbs" authorityYear="1926" box="[154,280,1209,1233]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–J.a
<taxonomicName id="F5DFB0F8320EFFC2FEDFFB797D8E51F6" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[326,399,1209,1233]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">pulchra</taxonomicName>
</emphasis>
lineage agrees with the hypothesis that sea-level fall isolates coastal fishes among disjunct estuaries (
<bibRefCitation id="564EB68A320EFFC2FFE5FB387D2C5037" author="Dolby GA &amp; Hechinger R &amp; Ellingson RA" box="[124,301,1272,1296]" pageId="13" refId="ref15608" refString="Dolby GA, Hechinger R, Ellingson RA et al. Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study. Proceedings Biological Sciences 2016; 283: 20161571. https: // doi. org / 10.1098 / rspb. 2016.1571" type="journal volume" year="2016">
Dolby
<emphasis id="00AB1769320EFFC2FF5AFB397CF25037" box="[195,243,1272,1296]" italics="true" pageId="13">et al.</emphasis>
2016
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FEA3FB387D6F5037" author="Dolby GA &amp; Ellingson RA &amp; Findley LT" box="[314,366,1272,1296]" pageId="13" pagination="994 - 1011" refId="ref15556" refString="Dolby GA, Ellingson RA, Findley LT et al. How sea level change mediates genetic divergence in coastal species across regions with varying tectonic and sediment processes. Molecular Ecology 2018; 27: 994 - 1011. https: // doi. org / 10.1111 / mec. 14487" type="journal article" year="2018">2018</bibRefCitation>
). Hence, we hypothesize that climatic cooling and periodic sea-level rise during Oligocene oscillations (
<bibRefCitation id="564EB68A320EFFC2FFE5FAF77D395068" author="Boulila S &amp; Galbrun B &amp; Miller KG" box="[124,312,1335,1359]" pageId="13" pagination="94 - 112" refId="ref14904" refString="Boulila S, Galbrun B, Miller KG et al. On the origin of Cenozoic and Mesozoic ' third-order' eustatic sequences. Earth-Science Reviews 2011; 109: 94 - 112. https: // doi. org / 10.1016 / j. earscirev. 2011.09.003" type="journal article" year="2011">
Boulila
<emphasis id="00AB1769320EFFC2FF55FAF77CFC5068" box="[204,253,1335,1359]" italics="true" pageId="13">et al.</emphasis>
2011
</bibRefCitation>
) allowed southward expansion of ancestral cyprinodontids, with one or more periods of sea-level fall (potentially Oi1b) subsequently isolating populations that had settled on the
<collectingRegion id="F01B0599320EFFC2FF7DFA557D34508A" box="[228,309,1429,1453]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
platform.
</paragraph>
<paragraph id="3260CB7B320EFFC2FF14FA74785F55E0" blockId="13.[113,764,607,1985]" lastBlockId="13.[809,1462,144,1985]" pageId="13">
Because living
<taxonomicName id="F5DFB0F8320EFFC2FEB8FA747D9E50EB" authorityName="Hubbs" authorityYear="1926" box="[289,415,1460,1484]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FEB8FA747D9E50EB" box="[289,415,1460,1484]" italics="true" pageId="13">Floridichthys</emphasis>
</taxonomicName>
have a disjunct distribution across the Gulf of
<collectingRegion id="F01B0599320EFFC2FF77FA137D3A50CC" box="[238,315,1491,1515]" country="Mexico" name="Mexico" pageId="13">México</collectingRegion>
between
<collectingRegion id="F01B0599320EFFC2FE3AFA137E7850CC" box="[419,633,1491,1515]" isEnumeration="true" pageId="13">Yucatán and Florida</collectingRegion>
, the MRCA of
<emphasis id="00AB1769320EFFC2FF09FA337D7F532C" box="[144,382,1523,1547]" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FF09FA337D0F532C" authorityName="Hubbs" authorityYear="1926" box="[144,270,1523,1547]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F8320EFFC2FE82FA337D7F532C" authorityName="Goode &amp; Bean" authorityYear="1879" box="[283,382,1523,1547]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Jordanella</taxonomicName>
</emphasis>
could be from
<collectingRegion id="F01B0599320EFFC2FDB0FA337ED7532C" box="[553,726,1523,1547]" isEnumeration="true" pageId="13">Florida, Yucatán</collectingRegion>
, or points between. Presence of the sister taxon
<taxonomicName id="F5DFB0F8320EFFC2FDA9F9D37E8B530D" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[560,650,1554,1578]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">
<emphasis id="00AB1769320EFFC2FDA9F9D37E8B530D" box="[560,650,1554,1578]" italics="true" pageId="13">J. pulchra</emphasis>
</taxonomicName>
in
<collectingRegion id="F01B0599320EFFC2FD30F9D27EFB530D" box="[681,762,1554,1578]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
favours
<collectingRegion id="F01B0599320EFFC2FF59F9F17D10536E" box="[192,273,1585,1609]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
as the ancestral area for
<taxonomicName id="F5DFB0F8320EFFC2FD9AF9F17E80536E" authorityName="Hubbs" authorityYear="1926" box="[515,641,1585,1609]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FD9AF9F17E80536E" box="[515,641,1585,1609]" italics="true" pageId="13">Floridichthys</emphasis>
</taxonomicName>
and is compatible with the hypothesis (above) that
<taxonomicName id="F5DFB0F8320EFFC2FD85F9917ECC534E" box="[540,717,1617,1641]" family="Cyprinodontidae" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="family">Cyprinodontidae</taxonomicName>
originated in the
<collectingCounty id="DB01B3F7320EFFC2FE9BF9B07D8453AF" box="[258,389,1648,1672]" pageId="13">western Gulf</collectingCounty>
of
<collectingRegion id="F01B0599320EFFC2FE3DF9B07DF053AF" box="[420,497,1648,1672]" country="Mexico" name="Mexico" pageId="13">México</collectingRegion>
(closer to
<collectingRegion id="F01B0599320EFFC2FDC2F9B07EAD53AF" box="[603,684,1648,1672]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
than to
<collectingRegion id="F01B0599320EFFC2FFE8F94F7CC05380" box="[113,193,1679,1703]" country="United States of America" name="Florida" pageId="13">Florida</collectingRegion>
). Furthermore, the Caribbean Loop Current could have facilitated later oceanic dispersal of
<taxonomicName id="F5DFB0F8320EFFC2FE71F96F7E6753E0" authorityName="Hubbs" authorityYear="1926" box="[488,614,1711,1735]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FE71F96F7E6753E0" box="[488,614,1711,1735]" italics="true" pageId="13">Floridichthys</emphasis>
</taxonomicName>
from
<collectingRegion id="F01B0599320EFFC2FD30F96F7EFB53E0" box="[681,762,1711,1735]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
to
<collectingRegion id="F01B0599320EFFC2FF16F90E7CDB53C1" box="[143,218,1742,1766]" country="United States of America" name="Florida" pageId="13">Florida</collectingRegion>
(as seen in several cyprinodontiform taxa; MassipVeloso
<emphasis id="00AB1769320EFFC2FF24F92E7CE95222" box="[189,232,1773,1797]" italics="true" pageId="13">et al</emphasis>
. 2024), whereas the current would have hampered immigration in the reverse direction. Hence, we suggest that the Oligocene ancestor of
<emphasis id="00AB1769320EFFC2FEFBF8EC7E4C5263" box="[354,589,1836,1860]" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FEFBF8EC7DE15263" authorityName="Hubbs" authorityYear="1926" box="[354,480,1836,1860]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F8320EFFC2FE77F8ED7E4C5263" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[494,589,1836,1860]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">J. pulchra</taxonomicName>
</emphasis>
dispersed along the
<collectingMunicipality id="D2045101320EFFC2FF02F88C7D655244" box="[155,356,1867,1891]" pageId="13">western Gulf Coast</collectingMunicipality>
from the
<location id="37009DA0320EFFC2FE49F88B7E4A5244" LSID="urn:lsid:plazi:treatment:BA767A6D320EFFC0FEFFFD9F7DF3579A:37009DA0320EFFC2FE49F88B7E4A5244" box="[464,587,1867,1891]" county="western Gulf" municipality="western Gulf Coast" name="Rio Grande" pageId="13" stateProvince="Mexico">Río Grande</location>
delta to the Bay of
<collectingRegion id="F01B0599320EFFC2FF08F8AB7D0052A4" box="[145,257,1899,1923]" country="Mexico" name="Campeche" pageId="13">Campeche</collectingRegion>
, where presence of coastal and shallow-marine areas ((
<bibRefCitation id="564EB68A320EFFC2FF5CF84A7DB25285" author="Villagomez D &amp; Steffensen C &amp; Pindell J" box="[197,435,1930,1954]" pageId="13" refId="ref20776" refString="Villagomez D, Steffensen C, Pindell J et al. Tectono-sedimentary evolution of Southern Mexico. Implications for Cretaceous and younger source-to-sink systems in the Mexican foreland basins and the Gulf of Mexico. Earth-Science Reviews 2022; 231: 104066. https: // doi. org / 10.1016 / j. earscirev. 2022.104066" type="journal volume" year="2022">
Villagómez
<emphasis id="00AB1769320EFFC2FEDBF84B7D775285" box="[322,374,1930,1954]" italics="true" pageId="13">et al.</emphasis>
2022
</bibRefCitation>
) could have provided suitable habitats (
<figureCitation id="AAE4D7FE320EFFC2FF48F8697D0852E6" box="[209,265,1961,1985]" captionStart="Figure 7" captionStartId="12.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1323]" captionTargetId="figure-237@12.[129,1473,144,1323]" captionTargetPageId="12" captionText="Figure 7. Oligocene diversification of North American killifishes in the Gulf of México region. Putative estuaries of origin for each Oligocene lineage are labelled.Phylogenetic nodes (Fig.3) are indicated as N4, N8, N12, and N13, showing hypothetical separations between lineages (for details, see main text, Table 2, and Fig.6). The base map is a reconstruction for 35 Mya (ending Late Eocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). Superimposed river drainages and estuaries are adapted from the study by Snedden and Galloway (2019). Details on the Chiapas region are from the study by Villagómez et al. (2022). Three putative dispersal events are illustrated: N8, cyprinodontid southward immigration founding Floridichthys–Jordanella pulchra on the Yucatán platform; N12, fundulid eastward immigration, including crossing the Suwanee Channel during sea-level recession, founding Leptolucania on the Ocala High; and N13, fundulid westward immigration founding Lucania–Wileyichthys in the Río Grande–Río Bravo estuary, where it presumably co-occurred with Cyprinodontidae." pageId="13">Fig. 7</figureCitation>
). However,
<emphasis id="00AB1769320EFFC2FE1CF8697E6D52E6" box="[389,620,1961,1985]" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FE1CF8697E0252E6" authorityName="Hubbs" authorityYear="1926" box="[389,515,1961,1985]" family="Cyprinodontidae" genus="Floridichthys" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Floridichthys</taxonomicName>
–
<taxonomicName id="F5DFB0F8320EFFC2FD88F86A7E6D52E6" baseAuthorityName="Hubbs" baseAuthorityYear="1936" box="[529,620,1961,1985]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">J. pulchra</taxonomicName>
</emphasis>
has uncertain phylogenetic placement (compare
<figureCitation id="AAE4D7FE320EFFC2FB15FF5078C4558F" box="[1164,1221,144,168]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
with
<bibRefCitation id="564EB68A320EFFC2FB66FF5079A3558F" author="Piller KR &amp; Parker E &amp; Lemmon AR" box="[1279,1442,144,168]" pageId="13" refId="ref19125" refString="Piller KR, Parker E, Lemmon AR et al. Investigating the utility of anchored hybrid enrichment data to investigate the relationships among the killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed group of fishes. Molecular Phylogenetics and Evolution 2022; 173: 107482. https: // doi. org / 10.1016 / j. ympev. 2022.107482" type="journal volume" year="2022">
Piller
<emphasis id="00AB1769320EFFC2FAA3FF51796B558F" box="[1338,1386,144,168]" italics="true" pageId="13">et al.</emphasis>
2022
</bibRefCitation>
), hence further study is needed.
</paragraph>
<paragraph id="3260CB7B320EFFC2FCDCFF0F7FD754C6" blockId="13.[809,1462,144,1985]" pageId="13">
Our estimate for vicariance of
<taxonomicName id="F5DFB0F8320EFFC2FB1DFF0F79B255C1" authority="Miller, 1956" authorityName="Miller" authorityYear="1956" box="[1156,1459,206,231]" family="Cyprinodontidae" genus="Cualac" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="tessellatus">
<emphasis id="00AB1769320EFFC2FB1DFF0F792E55C0" box="[1156,1327,207,231]" italics="true" pageId="13">Cualac tessellatus</emphasis>
Miller,
<quantity id="F527669E320EFFC2FAE6FF0E79B255C1" box="[1407,1459,206,230]" metricMagnitude="1" metricUnit="m" metricValue="4.96824" pageId="13" unit="in" value="1956.0">1956</quantity>
</taxonomicName>
in the Río Pánuco 29.9–14.6 Mya (
<figureCitation id="AAE4D7FE320EFFC2FB0AFF2E78CD5421" box="[1171,1228,238,262]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
, node 9) provides further support to the hypothesis that ancestral cyprinodontids inhabited the western Gulf Coast in the Oligocene. Presence of a nascent Río Pánuco (
<bibRefCitation id="564EB68A320EFFC2FB92FE8C791D5443" author="Beltran-Trivino A &amp; Martens UC &amp; von Quadt A" box="[1035,1308,332,356]" pageId="13" pagination="1 - 34" refId="ref14685" refString="Beltran-Trivino A, Martens UC, von Quadt A. Siliciclastic provenance of the Cenozoic stratigraphic succession in the southern Gulf of Mexico: insights from U-Pb detrital zircon geochronology and heavy minerals analysis. In: Martens UC, Molina Garza RS, (eds), Southern and Central Mexico: Basement Framework, Tectonic Evolution, and Provenance of Mesozoic - Cenozoic Basins. Geological Society of America Special Paper 546. Boulder: Geological Society of America, 2021, 1 - 34." type="book chapter" year="2021">
Beltrán-Triviño
<emphasis id="00AB1769320EFFC2FB29FE8D78E05443" box="[1200,1249,332,356]" italics="true" pageId="13">et al.</emphasis>
2021
</bibRefCitation>
) suggests that ancestral
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<emphasis id="00AB1769320EFFC2FC15FEAB783854A4" box="[908,1081,363,387]" italics="true" pageId="13">Cualac tessellatus</emphasis>
</taxonomicName>
ranged inland along this route (
<figureCitation id="AAE4D7FE320EFFC2FA14FEAB7F365484" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="13">Fig. 8</figureCitation>
). Late Oligocene–Early Miocene uplift of the Chicontepec Basin (
<bibRefCitation id="564EB68A320EFFC2FCECFE6A782854E5" author="Roure F &amp; Alzaga-Ruiz H &amp; Callot JP" box="[885,1065,426,450]" pageId="13" pagination="169 - 89" refId="ref19935" refString="Roure F, Alzaga-Ruiz H, Callot JP et al. Long lasting interactions between tectonic loading, unroofing, post-rift thermal subsidence and sedimentary transfers along the western margin of the Gulf of Mexico: some insights from integrated quantitative studies. Tectonophysics 2009; 475: 169 - 89. https: // doi. org / 10.1016 / j. tecto. 2009.04.012" type="journal article" year="2009">
Roure
<emphasis id="00AB1769320EFFC2FC24FE6B7FEF54E5" box="[957,1006,426,450]" italics="true" pageId="13">et al.</emphasis>
2009
</bibRefCitation>
) could have stranded this population inland (
<tableCitation id="7F5DFEC0320EFFC2FCE2FE097FC754C6" box="[891,966,457,481]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="13">Table 2</tableCitation>
).
</paragraph>
<paragraph id="3260CB7B320EFFC2FCDCFE297FE65132" blockId="13.[809,1462,144,1985]" pageId="13">
As already mentioned, our phylogeny indicates that
<taxonomicName id="F5DFB0F8320EFFC2FAC9FE2979B25726" authorityName="Goode &amp; Bean" authorityYear="1879" box="[1360,1459,489,513]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FAC9FE2979B25726" box="[1360,1459,489,513]" italics="true" pageId="13">Jordanella</emphasis>
</taxonomicName>
is polyphyletic (
<figureCitation id="AAE4D7FE320EFFC2FC55FDC878025707" box="[972,1027,520,544]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
). Separation of
<taxonomicName id="F5DFB0F8320EFFC2FB39FDC978FA5707" authorityName="Goode &amp; Bean" authorityYear="1879" box="[1184,1275,520,544]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="floridae">
<emphasis id="00AB1769320EFFC2FB39FDC978FA5707" box="[1184,1275,520,544]" italics="true" pageId="13">J.floridae</emphasis>
</taxonomicName>
from
<emphasis id="00AB1769320EFFC2FAADFDC87F9F5718" italics="true" pageId="13">
<taxonomicName id="F5DFB0F8320EFFC2FAADFDC879A55707" authorityName="Miller and Walters" authorityYear="1972" box="[1332,1444,520,544]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Megupsilon</taxonomicName>
–
<taxonomicName id="F5DFB0F8320EFFC2FCB3FDE77F9F5718" box="[810,926,551,575]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">Cyprinodon</taxonomicName>
</emphasis>
23.3–10.8 Mya (
<figureCitation id="AAE4D7FE320EFFC2FBCDFDE7788F5718" box="[1108,1166,551,575]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
, node 10;
<tableCitation id="7F5DFEC0320EFFC2FA98FDE7794D5718" box="[1281,1356,551,575]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="13">Table 2</tableCitation>
) suggests that an ancestral cyprinodontid dispersed eastwards from the northwestern Gulf of
<collectingRegion id="F01B0599320EFFC2FB94FDA6785B5759" box="[1037,1114,614,638]" country="Mexico" name="Mexico" pageId="13">México</collectingRegion>
during the Early Miocene, when the nascent
<collectingRegion id="F01B0599320EFFC2FC33FD457FF457BA" box="[938,1013,645,669]" country="United States of America" name="Florida" pageId="13">Florida</collectingRegion>
Peninsula was an island (i.e. Ocala High). Our 16.8 Mya point estimate for time of divergence closely follows the onset of the MMCO, a period of high temperatures, reduced global glaciation, and elevated sea levels spanning 17.0–13.8 Mya (
<bibRefCitation id="564EB68A320EFFC2FC4AFCC37886563C" author="Miller KG &amp; Browning JV &amp; Schmelz WJ" box="[979,1159,771,795]" pageId="13" refId="ref18255" refString="Miller KG, Browning JV, Schmelz WJ et al. Cenozoic sea-level and cryospheric evolution from deep-sea geochemical and continental margin records. Science Advances 2020 a; 6: eaaz 1346. https: // doi. org / 10.1126 / sciadv. aaz 1346" type="journal volume" year="2020">
Miller
<emphasis id="00AB1769320EFFC2FB8FFCC3783F563C" box="[1046,1086,771,795]" italics="true" pageId="13">et al</emphasis>
. 2020a
</bibRefCitation>
). This timing suggests that an ancestral cyprinodontid was distributed along the northeastern Gulf Coast, ranging between the Ocala High and the mainland while the Gulf Trough, which separated these lands, was narrowed (
<figureCitation id="AAE4D7FE320EFFC2FCE2FC407FB556BF" box="[891,948,896,920]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="13">Fig. 8</figureCitation>
). Sea-level rise during the MMCO greatly broadened and deepened the Gulf Trough, shrinking the Ocala High to a small island (
<bibRefCitation id="564EB68A320EFFC2FC42FC7F786E56F0" author="Popenoe P" box="[987,1135,959,983]" pageId="13" pagination="352 - 80" refId="ref19219" refString="Popenoe P. Paleoceonography and Paleogeography of the Miocene of the Southeastern United States. In: Burnett WC, Riggs SR (eds.), Phosphate Deposits of the World. Cambridge: Cambridge University Press, 1990, 352 - 80." type="book chapter" year="1990">Popenoe 1990</bibRefCitation>
). We hypothesize that
<taxonomicName id="F5DFB0F8320EFFC2FAC1FC7F79B256F0" authorityName="Goode &amp; Bean" authorityYear="1879" box="[1368,1459,959,983]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="floridae">
<emphasis id="00AB1769320EFFC2FAC1FC7F79B256F0" box="[1368,1459,959,983]" italics="true" pageId="13">J. floridae</emphasis>
</taxonomicName>
descends from a peripheral cyprinodontid population persisting around this island.
</paragraph>
<paragraph id="3260CB7B320EFFC2FCDCFBDD787F5068" blockId="13.[809,1462,144,1985]" pageId="13">
Notably, the genus
<taxonomicName id="F5DFB0F8320EFFC2FB84FBDD78815112" authorityName="Goode &amp; Bean" authorityYear="1879" box="[1053,1152,1053,1077]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FB84FBDD78815112" box="[1053,1152,1053,1077]" italics="true" pageId="13">Jordanella</emphasis>
</taxonomicName>
has never been resolved as monophyletic with molecular data (
<bibRefCitation id="564EB68A320EFFC2FB3CFBFC795B5173" author="Miller RR &amp; Minckley WL &amp; Norris SM" box="[1189,1370,1084,1108]" pageId="13" refId="ref18488" refString="Miller RR, Minckley WL, Norris SM. Freshwater Fishes of Mexico. Chicago: The University of Chicago Press, 2005." type="book" year="2005">
Miller
<emphasis id="00AB1769320EFFC2FB75FBFD79165173" box="[1260,1303,1084,1108]" italics="true" pageId="13">et al</emphasis>
. 2005
</bibRefCitation>
,
<bibRefCitation id="564EB68A320EFFC2FAF1FBFC7FE55154" author="Echelle AA &amp; Echelle AF" pageId="13" pagination="609 - 73" refId="ref15900" refString="Echelle AA, Echelle AF. Cyprinodontidae: Pupfishes. In: Warren ML Jr, Burr BM (eds), Freshwater fishes of North America, Vol. 2, Characidae to Poeciliidae. Baltimore: Johns Hopkins University Press, 2020, 609 - 73." type="book chapter" year="2020">Echelle and Echelle 2020</bibRefCitation>
). This, along with the biogeographical evidence provided here, could justify resurrection of the name
<taxonomicName id="F5DFB0F8320EFFC2FCB3FB5A7FF05195" authorityName="Hubbs" authorityYear="1936" box="[810,1009,1178,1202]" family="Cyprinodontidae" genus="Garmanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="pulchra">
<emphasis id="00AB1769320EFFC2FCB3FB5A7FF05195" box="[810,1009,1178,1202]" italics="true" pageId="13">Garmanella pulchra</emphasis>
</taxonomicName>
for the
<collectingRegion id="F01B0599320EFFC2FBD3FB5A789A5195" box="[1098,1179,1178,1202]" country="Mexico" name="Yucatan" pageId="13">Yucatán</collectingRegion>
species. In this case, morphological similarities between species presently recognized as
<taxonomicName id="F5DFB0F8320EFFC2FCB5FB197F8E51D6" authorityName="Goode &amp; Bean" authorityYear="1879" box="[812,911,1241,1265]" family="Cyprinodontidae" genus="Jordanella" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FCB5FB197F8E51D6" box="[812,911,1241,1265]" italics="true" pageId="13">Jordanella</emphasis>
</taxonomicName>
might reflect ancestral traits that persisted in ancient peripheral-isolate taxa sequestered in ancestral cyprinodontid niches on either side of the Gulf of
<collectingRegion id="F01B0599320EFFC2FB3FFAD778F55008" box="[1190,1268,1303,1327]" country="Mexico" name="Mexico" pageId="13">México</collectingRegion>
. The taxonomy of these species needs further study.
</paragraph>
<paragraph id="3260CB7B320EFFC2FCDCFA9679A7536E" blockId="13.[809,1462,144,1985]" pageId="13">
Late Miocene divergence of
<taxonomicName id="F5DFB0F8320EFFC2FBE9FA9678E15049" authorityName="Miller and Walters" authorityYear="1972" box="[1136,1248,1366,1390]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FBE9FA9678E15049" box="[1136,1248,1366,1390]" italics="true" pageId="13">Megupsilon</emphasis>
</taxonomicName>
(14.9–7.4 Mya;
<figureCitation id="AAE4D7FE320EFFC2FA16FA967F3650AA" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Fig. 3</figureCitation>
, node 11) might represent Late Miocene emergence of ancestral Río San Fernando (Río Bravo in the work of
<bibRefCitation id="564EB68A320EFFC2FAC0FA557FF350EB" author="Snedden JW &amp; Galloway WE" pageId="13" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
) as a corridor for inland invasion (
<tableCitation id="7F5DFEC0320EFFC2FAC6FA7479AD50EB" box="[1375,1452,1460,1484]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="13">Table 2</tableCitation>
;
<figureCitation id="AAE4D7FE320EFFC2FCB3FA137F6250CC" box="[810,867,1491,1515]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="13">Fig. 8</figureCitation>
). This species (extinct in the wild) inhabited a spring-fed habitat (
<bibRefCitation id="564EB68A320EFFC2FC1BFA33782E532D" author="Miller RR &amp; Minckley WL &amp; Norris SM" box="[898,1071,1522,1547]" pageId="13" refId="ref18488" refString="Miller RR, Minckley WL, Norris SM. Freshwater Fishes of Mexico. Chicago: The University of Chicago Press, 2005." type="book" year="2005">
Miller
<emphasis id="00AB1769320EFFC2FC5EFA337FF7532C" box="[967,1014,1523,1547]" italics="true" pageId="13">et al.</emphasis>
2005
</bibRefCitation>
) and might have speciated as a spring endemic. It is possible that episodes of aridity, tectonism, or volcanism isolated the spring system, but this needs further study.
</paragraph>
<paragraph id="3260CB7B320EFFC0FCDCF9917DF3579A" blockId="13.[809,1462,144,1985]" lastBlockId="15.[112,763,144,701]" lastPageId="15" pageId="13">
The rate of
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<emphasis id="00AB1769320EFFC2FC24F9917830534E" box="[957,1073,1617,1641]" italics="true" pageId="13">Cyprinodon</emphasis>
</taxonomicName>
speciation increased upon separation from
<taxonomicName id="F5DFB0F8320EFFC2FCFDF9B07FD553AF" authorityName="Miller and Walters" authorityYear="1972" box="[868,980,1648,1672]" family="Cyprinodontidae" genus="Megupsilon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FCFDF9B07FD553AF" box="[868,980,1648,1672]" italics="true" pageId="13">Megupsilon</emphasis>
</taxonomicName>
(node 11;
<figureCitation id="AAE4D7FE320EFFC2FBD2F9B0788653AF" box="[1099,1159,1648,1672]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="13">Figs 3</figureCitation>
,
<figureCitation id="AAE4D7FE320EFFC2FB0DF9B078A053AF" box="[1172,1185,1648,1672]" captionStart="Figure 4" captionStartId="8.[127,192,1459,1483]" captionTargetBox="[132,1470,147,1429]" captionTargetId="figure-233@8.[129,1473,144,1432]" captionTargetPageId="8" captionText="Figure 4. A, BAMM phylorate plot showing speciation rates along each branch within the North American killifish families Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae. Cool colours represent slow rates; warm colours represent fast rates.B–E, speciation rate over time is shown graphically for Fundulidae (B), Cyprinodontidae (C), Profundulidae (D), and Goodeidae (E). λ = mean diversification rate." pageId="13">4</figureCitation>
). Several factors are likely to have contributed to this trend. Initially, inland invasions into five Late Miocene rivers subdivided
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<emphasis id="00AB1769320EFFC2FB0DF96F790953E0" box="[1172,1288,1711,1735]" italics="true" pageId="13">Cyprinodon</emphasis>
</taxonomicName>
into as many upland lineages, four of which dispersed far across the desert region (
<bibRefCitation id="564EB68A320EFFC2FCACF92E78755222" author="Hoagstrom CW &amp; Osborne MJ" box="[821,1140,1773,1798]" pageId="13" pagination="20 - 76" refId="ref17354" refString="Hoagstrom CW, Osborne MJ. Biogeography of Cyprinodon across the Great Plains-Chihuahuan Desert Region and Adjacent Areas. Proceedings of the Desert Fishes Council Special Publication 2021; 2021: 20 - 76." type="journal article" year="2021">Hoagstrom and Osborne 2021</bibRefCitation>
). A relative of this western inland radiation also dispersed to Yucatán, founding an endemic lineage there (
<taxonomicName id="F5DFB0F8320EFFC2FC5EF8EC78995263" authorityName="Hubbs" authorityYear="1936" box="[967,1176,1836,1860]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="species" species="artifrons">
<emphasis id="00AB1769320EFFC2FC5EF8EC78995263" box="[967,1176,1836,1860]" italics="true" pageId="13">Cyprinodon artifrons</emphasis>
</taxonomicName>
species group;
<figureCitation id="AAE4D7FE320EFFC2FADDF8EC79845263" box="[1348,1413,1836,1860]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="13">Figs 2</figureCitation>
,
<figureCitation id="AAE4D7FE320EFFC2FA0CF8EC79A35263" box="[1429,1442,1836,1860]" captionStart="Figure 5" captionStartId="10.[129,194,1836,1860]" captionTargetBox="[212,1390,147,1805]" captionTargetId="figure-7@10.[209,1393,144,1808]" captionTargetPageId="10" captionText="Figure 5. Reconstruction of ancestral areas for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifishes divided as Nearctic = A and Neotropical = B, with the boundary between realms being the Trans-Mexican Volcanic Belt (Rico et al.2022). B, ancestral areas computed using the BAYAREALIKE+J model in BIOGEOBEARS." pageId="13">5</figureCitation>
). Once inland invasions were underway, barrier displacement via drainage rearrangements, climate fluctuations, tectonism, and volcanism caused widespread allopatric diversification. Meanwhile, a maritime lineage of
<taxonomicName id="F5DFB0F8320EFFC2FB0DF869790952E6" box="[1172,1288,1961,1985]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320EFFC2FB0DF869790952E6" box="[1172,1288,1961,1985]" italics="true" pageId="13">Cyprinodon</emphasis>
</taxonomicName>
remained along the Gulf Coast (
<bibRefCitation id="564EB68A320CFFC0FE8FFF507DCC558F" author="Echelle AA &amp; Carson EW &amp; Echelle AF" box="[278,461,144,168]" pageId="15" pagination="320 - 39" refId="ref15851" refString="Echelle AA, Carson EW, Echelle AF et al. Historical biogeography of the New-World pupfish genus Cyprinodon (Teleostei: Cyprinodontidae). Copeia 2005; 2005: 320 - 39. https: // doi. org / 10.1643 / cg- 03 - 093 r 3" type="journal article" year="2005">
Echelle
<emphasis id="00AB1769320CFFC0FEFFFF517D94558F" box="[358,405,144,168]" italics="true" pageId="15">et al.</emphasis>
2005
</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FE41FF507E0D558F" author="Echelle AA &amp; Fuselier L &amp; Van Den Bussche RA" box="[472,524,144,168]" pageId="15" pagination="855 - 64" refId="ref15949" refString="Echelle AA, Fuselier L, Van Den Bussche RA et al. Molecular systematics of Hispaniolan pupfishes (Cyprinodontidae: Cyprinodon): implications for the biogeography of insular Caribbean fishes. Molecular Phylogenetics and Evolution 2006; 39: 855 - 64. https: // doi. org / 10.1016 / j. ympev. 2005.12.010" type="journal article" year="2006">2006</bibRefCitation>
). This geography is unclear in our reconstruction of ancestral habitats (
<figureCitation id="AAE4D7FE320CFFC0FDC5FF6F7E9555E0" box="[604,660,175,199]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="15">Fig. 2</figureCitation>
), because the same widespread ancestor produced sequential upland invasions from the coast, as already described. Furthermore, during the Pleistocene, maritime
<taxonomicName id="F5DFB0F8320CFFC0FE18FECD7DF45402" box="[385,501,269,293]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FE18FECD7DF45402" box="[385,501,269,293]" italics="true" pageId="15">Cyprinodon</emphasis>
</taxonomicName>
made new invasions into the desert region (
<bibRefCitation id="564EB68A320CFFC0FEB7FEED7EF45462" author="Lozano-Vilano M de &amp; Contreras-Balderas S" box="[302,757,301,325]" pageId="15" pagination="382 - 7" refId="ref17715" refString="Lozano-Vilano M de L, Contreras-Balderas S. Cyprinodon bobmilleri: new species of pupfish from Nuevo Leon, Mexico (Pisces: Cyprinodontidae). Copeia 1999; 1999: 382 - 7. https: // doi. org / 10.2307 / 1447484" type="journal article" year="1999">Lozano-Vilano and Contreras-Balderas 1999</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FFE8FE8C7DA75443" author="Hoagstrom CW &amp; Osborne MJ" box="[113,422,332,356]" pageId="15" pagination="20 - 76" refId="ref17354" refString="Hoagstrom CW, Osborne MJ. Biogeography of Cyprinodon across the Great Plains-Chihuahuan Desert Region and Adjacent Areas. Proceedings of the Desert Fishes Council Special Publication 2021; 2021: 20 - 76." type="journal article" year="2021">Hoagstrom and Osborne 2021</bibRefCitation>
) and immigrated to Caribbean islands and South America (
<bibRefCitation id="564EB68A320CFFC0FEE4FEAC7E2A54A4" author="Haney RA &amp; Silliman BR &amp; Fry AJ" box="[381,555,363,388]" pageId="15" pagination="743 - 54" refId="ref16969" refString="Haney RA, Silliman BR, Fry AJ et al. The Pleistocene history of the sheepshead minnow (Cyprinodon variegatus): non-equilibrium evolutionary dynamics within a diversifying species complex. Molecular Phylogenetics and Evolution 2007; 43: 743 - 54. https: // doi. org / 10.1016 / j. ympev. 2006.08.022" type="journal article" year="2007">
Haney
<emphasis id="00AB1769320CFFC0FE5FFEAC7DF254A4" box="[454,499,363,387]" italics="true" pageId="15">et al.</emphasis>
2007
</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FDADFEAB7E6954A4" author="Haney RA &amp; Turner BJ &amp; Rand DM" box="[564,616,363,387]" pageId="15" pagination="1108 - 14" refId="ref17027" refString="Haney RA, Turner BJ, Rand DM. A cryptic lineage within the pupfish Cyprinodon dearborni suggests multiple colonizations of South America. Journal of Fish Biology 2009; 75: 1108 - 14. https: // doi. org / 10.1111 / j. 1095 - 8649.2009.02378. x" type="journal article" year="2009">2009</bibRefCitation>
). The Yucatán lineage produced a species flock (
<bibRefCitation id="564EB68A320CFFC0FE5DFE4B7E505484" author="Strecker U" box="[452,593,395,419]" pageId="15" pagination="865 - 72" refId="ref20448" refString="Strecker U. Genetic differentiation and reproductive isolation in a Cyprinodon fish species flock from Laguna Chichancanab, Mexico. Molecular Phylogenetics and Evolution 2006; 39: 865 - 72. https: // doi. org / 10.1016 / j. ympev. 2006.01.008" type="journal article" year="2006">Strecker 2006</bibRefCitation>
;
<figureCitation id="AAE4D7FE320CFFC0FDC4FE4B7E945484" box="[605,661,395,419]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="15">Fig. 3</figureCitation>
). Beyond this, our analysis might underestimate the
<taxonomicName id="F5DFB0F8320CFFC0FD81FE6A7E8D54E5" box="[536,652,426,450]" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="genus">
<emphasis id="00AB1769320CFFC0FD81FE6A7E8D54E5" box="[536,652,426,450]" italics="true" pageId="15">Cyprinodon</emphasis>
</taxonomicName>
speciation rate, because a species flock from
<collectingRegion id="F01B0599320CFFC0FE25FE097E3E54C6" box="[444,575,457,481]" country="Bahamas" name="San Salvador" pageId="15">San Salvador</collectingRegion>
,
<collectingCountry id="4AC88BEB320CFFC0FDDEFE097EA254C6" box="[583,675,457,481]" name="Bahamas" pageId="15">Bahamas</collectingCountry>
(
<bibRefCitation id="564EB68A320CFFC0FD2BFE097D4C5726" author="Martin CH &amp; Wainwright PC" pageId="15" pagination="208 - 11" refId="ref17901" refString="Martin CH, Wainwright PC. Multiple fitness peaks on the adaptive landscape drive adaptive radiation in the wild. Science 2013; 339: 208 - 11. https: // doi. org / 10.1126 / science. 1227710" type="journal article" year="2013">Martin and Wainwright 2013</bibRefCitation>
) and the subspecies
<taxonomicName id="F5DFB0F8320CFFC0FD87FE297D245707" authority="Carr, 1936" authorityName="Carr" authorityYear="1936" family="Cyprinodontidae" genus="Cyprinodon" kingdom="Animalia" order="Cyprinodontiformes" pageId="15" phylum="Chordata" rank="subSpecies" species="variegatus" subSpecies="hubbsi">
<emphasis id="00AB1769320CFFC0FD87FE297CB05707" italics="true" pageId="15">Cyprinodon variegatus hubbsi</emphasis>
Carr, 1936
</taxonomicName>
(a potential distinct species;
<bibRefCitation id="564EB68A320CFFC0FDCAFDC87CA45718" author="Brix KV &amp; Grosell M" pageId="15" pagination="854 - 66" refId="ref14951" refString="Brix KV, Grosell M. Evaluation of pre- and post-zygotic mating barriers, hybrid fitness and phylogenetic relationship between Cyprinodon variegatus variegatus and Cyprinodon variegatus hubbsi (Cyprinodontiformes, Teleostei). Journal of Evolutionary Biology 2013; 26: 854 - 66. https: // doi. org / 10.1111 / jeb. 12099" type="journal article" year="2013">Brix and Grosell 2013</bibRefCitation>
,
<bibRefCitation id="564EB68A320CFFC0FF2DFDE87D595718" author="Jung EH &amp; Brix KV &amp; Brauner CJ" box="[180,344,551,576]" pageId="15" pagination="28 - 33" refId="ref17427" refString="Jung EH, Brix KV, Brauner CJ. The effect of temperature acclimation on thermal tolerance, hypoxia tolerance and aerobic scope in two subspecies of sheepshead minnow; Cyprinodon variegatus variegatus and Cyprinodon variegatus hubbsi. Comparative Biochemistry and Physiology. Part A, Molecular &amp; Integrative Physiology 2019; 232: 28 - 33. https: // doi. org / 10.1016 / j. cbpa. 2019.03.004" type="journal article" year="2019">
Jung
<emphasis id="00AB1769320CFFC0FF72FDE87D1C5718" box="[235,285,551,575]" italics="true" pageId="15">et al.</emphasis>
2019
</bibRefCitation>
) are absent from our phylogeny, which also leaves out seven
<taxonomicName id="F5DFB0F8320CFFC0FECBFD877E195778" box="[338,536,583,607]" pageId="15">
<emphasis id="00AB1769320CFFC0FECBFD877DC75778" box="[338,454,583,607]" italics="true" pageId="15">Cyprinodon</emphasis>
species
</taxonomicName>
for which we had no genetic data. Finally, six of the recognized species included in the analysis are likely to be polyphyletic (
<bibRefCitation id="564EB68A320CFFC0FE77FD457EEE57BA" author="Echelle AA &amp; Echelle AF" box="[494,751,645,669]" pageId="15" pagination="609 - 73" refId="ref15900" refString="Echelle AA, Echelle AF. Cyprinodontidae: Pupfishes. In: Warren ML Jr, Burr BM (eds), Freshwater fishes of North America, Vol. 2, Characidae to Poeciliidae. Baltimore: Johns Hopkins University Press, 2020, 609 - 73." type="book chapter" year="2020">Echelle and Echelle 2020</bibRefCitation>
) but are represented as one taxon here.
</paragraph>
</subSubSection>
</treatment>
</document>

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Subgenus
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First branching within subgenus
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<emphasis id="00AB17693213FFDFFE5FFEED7E215462" box="[454,544,301,325]" italics="true" pageId="16">Fundulus</emphasis>
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might reflect both ecological and geographical speciation.
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<emphasis id="00AB17693213FFDFFE75FE8C7E905443" box="[492,657,332,356]" italics="true" pageId="16">Fundulus majalis</emphasis>
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<emphasis id="00AB17693213FFDFFF59FEAB7CCC54A4" box="[192,205,363,387]" italics="true" pageId="16">–</emphasis>
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<emphasis id="00AB17693213FFDFFF55FEAB7D8B54A4" box="[204,394,363,387]" italics="true" pageId="16">Fundulus persimilis</emphasis>
<bibRefCitation id="564EB68A3213FFDFFE0AFEAB7E1254A4" author="Miller RR" box="[403,531,363,387]" pageId="16" pagination="1 - 64" refId="ref18453" refString="Miller RR. A systematic review of the Middle American fishes of the genus Profundulus. Miscellaneous Publications of the Museum of Zoology, University of Michigan 1955; 92: 1 - 64." type="journal article" year="1955">Miller, 1955</bibRefCitation>
</taxonomicName>
<emphasis id="00AB17693213FFDFFD8AFEAB7E2154A4" box="[531,544,363,387]" italics="true" pageId="16">–</emphasis>
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<emphasis id="00AB17693213FFDFFDB9FEAB7EC154A4" box="[544,704,363,387]" italics="true" pageId="16">Fundulus similis</emphasis>
(Baird &amp; Girard, 1853)
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inhabit unvegetated coastal habitats where they dive into soft sediments for cover rather than retreating to vegetation like typical
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<emphasis id="00AB17693213FFDFFEA5FE097D9754C6" box="[316,406,457,481]" italics="true" pageId="16">Fundulus</emphasis>
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(
<bibRefCitation id="564EB68A3213FFDFFE3FFE097EAD54C6" author="Martin RA &amp; Finucane JH" box="[422,684,457,481]" pageId="16" pagination="101 - 11" refId="ref17978" refString="Martin RA, Finucane JH. Reproduction and ecology of the longnose killifish. QuarterlyJournaloftheFloridaAcademyofSciences 1968; 31: 101 - 11." type="journal article" year="1968">Martin &amp; Finucane, 1968</bibRefCitation>
;
<bibRefCitation id="564EB68A3213FFDFFD23FE0A7CB45726" author="Harvey CJ" pageId="16" pagination="307 - 10" refId="ref17079" refString="Harvey CJ. Use of sandy beach habitat by Fundulus majalis, a surf-zone fish. Marine Ecology Progress Series 1998; 164: 307 - 10. https: // doi. org / 10.3354 / meps 164307" type="journal article" year="1998">Harvey, 1998</bibRefCitation>
;
<bibRefCitation id="564EB68A3213FFDFFF58FE297D6B5727" author="Miller RR &amp; Minckley WL &amp; Norris SM" box="[193,362,488,513]" pageId="16" refId="ref18488" refString="Miller RR, Minckley WL, Norris SM. Freshwater Fishes of Mexico. Chicago: The University of Chicago Press, 2005." type="book" year="2005">
Miller
<emphasis id="00AB17693213FFDFFE9DFE297D335726" box="[260,306,489,513]" italics="true" pageId="16">et al.</emphasis>
2005
</bibRefCitation>
). They are adapted for continuous swimming in the surf zone (
<bibRefCitation id="564EB68A3213FFDFFEE7FDC87E855707" author="Yetsko K &amp; Sancho G" box="[382,644,520,544]" pageId="16" pagination="827 - 33" refId="ref21196" refString="Yetsko K, Sancho G. The effects of salinity on swimming performance of two estuarine fishes, Fundulus heteroclitus and Fundulus majalis. Journal of Fish Biology 2015; 86: 827 - 33. https: // doi. org / 10.1111 / jfb. 12590" type="journal article" year="2015">Yetsko and Sancho 2015</bibRefCitation>
) and spawn in unvegetated habitats (
<bibRefCitation id="564EB68A3213FFDFFE1AFDE77E495718" author="Greeley MS &amp; Marion KR &amp; MacGregor R" box="[387,584,551,575]" pageId="16" pagination="125 - 31" refId="ref16843" refString="Greeley MS, Marion KR, MacGregor R. Semilunar spawning cycles of Fundulus similis (Cyprinodontidae). Environmental Biology of Fishes 1986; 17: 125 - 31. https: // doi. org / 10.1007 / bf 00001742" type="journal article" year="1986">
Greeley
<emphasis id="00AB17693213FFDFFE42FDE87E0C5718" box="[475,525,551,575]" italics="true" pageId="16">et al.</emphasis>
1986
</bibRefCitation>
).
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<emphasis id="00AB17693213FFDFFDF8FDE77F085718" box="[609,777,551,575]" italics="true" pageId="16">Fundulus majalis</emphasis>
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segregates from congeners in unvegetated, high-salinity waters (
<bibRefCitation id="564EB68A3213FFDFFF15FDA67D295759" author="Weisberg SB" box="[140,296,614,638]" pageId="16" pagination="249 - 57" refId="ref21072" refString="Weisberg SB. Competition and coexistence among four estuarine species of Fundulus. American Zoologist 1986; 26: 249 - 57. https: // doi. org / 10.1093 / icb / 26.1.249" type="journal article" year="1986">Weisberg 1986</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFEAEFDA67E415759" author="Wagner CM &amp; Austin HM" box="[311,576,614,638]" pageId="16" pagination="197 - 212" refId="ref20885" refString="Wagner CM, Austin HM. Correspondence between environmental gradients and summer littoral fish assemblages in low salinity reaches of the Chesapeake Bay, USA. Marine Ecology Progress Series 1999; 177: 197 - 212. https: // doi. org / 10.3354 / meps 177197" type="journal article" year="1999">Wagner and Austin 1999</bibRefCitation>
). We propose that this ecologically divergent lineage descends from an ancestor adapted for open, wave-swept coasts. Divergence 21.0–15.2 Mya (
<figureCitation id="AAE4D7FE3213FFDFFF12FD047CC257FB" box="[139,195,708,732]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="16">Fig. 3</figureCitation>
, node 16) was concurrent with sea-level rise that culminated in the MMCO (
<tableCitation id="7F5DFEC03213FFDFFECBFD237D9D57DC" box="[338,412,739,763]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="16">Table 2</tableCitation>
) and could have facilitated range expansion (
<bibRefCitation id="564EB68A3213FFDFFF79FCC37D8B563C" author="Dolby GA &amp; Hechinger R &amp; Ellingson RA" box="[224,394,771,795]" pageId="16" refId="ref15608" refString="Dolby GA, Hechinger R, Ellingson RA et al. Sea-level driven glacial-age refugia and post-glacial mixing on subtropical coasts, a palaeohabitat and genetic study. Proceedings Biological Sciences 2016; 283: 20161571. https: // doi. org / 10.1098 / rspb. 2016.1571" type="journal volume" year="2016">
Dolby
<emphasis id="00AB17693213FFDFFEBDFCC37D4D563C" box="[292,332,771,795]" italics="true" pageId="16">et al</emphasis>
. 2016
</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFE0DFCC37DC9563C" author="Dolby GA &amp; Ellingson RA &amp; Findley LT" box="[404,456,771,795]" pageId="16" pagination="994 - 1011" refId="ref15556" refString="Dolby GA, Ellingson RA, Findley LT et al. How sea level change mediates genetic divergence in coastal species across regions with varying tectonic and sediment processes. Molecular Ecology 2018; 27: 994 - 1011. https: // doi. org / 10.1111 / mec. 14487" type="journal article" year="2018">2018</bibRefCitation>
). Given that coastlines adjacent to Early Miocene deltas were wave swept and sand dominated (
<bibRefCitation id="564EB68A3213FFDFFF12FC817DC2567E" author="Snedden JW &amp; Galloway WE" box="[139,451,833,857]" pageId="16" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
), this could account for ecological specialization in this species group. In the earliest Middle Miocene (15.6 Mya), a new
<collectingCountry id="4AC88BEB3213FFDFFE3EFC407E1656BF" box="[423,535,896,920]" name="Guadeloupe" pageId="16">Guadalupe</collectingCountry>
River delta emerged to the west (
<figureCitation id="AAE4D7FE3213FFDFFF73FC5F7D255690" box="[234,292,927,951]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="16">Fig. 8</figureCitation>
). Wave-dominated conditions there (
<bibRefCitation id="564EB68A3213FFDFFD36FC5F7D4556F0" author="Snedden JW &amp; Galloway WE" pageId="16" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
) suggest that the MRCA of the
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<emphasis id="00AB17693213FFDFFD1EFC7F7EDB56F0" box="[647,730,959,983]" italics="true" pageId="16">F. similis</emphasis>
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species group evolved there.
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<subSubSection id="7AC598F03213FFDFFF46FBD57DAD5108" box="[223,428,1045,1071]" pageId="16" type="nomenclature">
<paragraph id="3260CB7B3213FFDFFF46FBD57DAD5108" blockId="16.[223,683,1045,1071]" box="[223,428,1045,1071]" pageId="16">
Subgenus
<taxonomicName id="F5DFB0F83213FFDFFEA5FBD67DAD5108" authorityName="Agassiz" authorityYear="1854" box="[316,428,1046,1071]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="subGenus" subGenus="Zygonetes">Zygonectes</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="7AC598F03213FFDFFE34FBD67F0B53C1" pageId="16" type="discussion">
<paragraph id="3260CB7B3213FFDFFE34FBD67EAA5108" blockId="16.[223,683,1045,1071]" box="[429,683,1045,1071]" pageId="16">
–
<taxonomicName id="F5DFB0F83213FFDFFE22FBD67E2E5108" authorityName="Cope" authorityYear="1865" box="[443,559,1045,1071]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="sciadicus">F. sciadicus</taxonomicName>
–
<taxonomicName id="F5DFB0F83213FFDFFDA7FBD57EAA5108" authorityName="Garman" authorityYear="1895" box="[574,683,1045,1071]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="genus">Plancterus</taxonomicName>
</paragraph>
<paragraph id="3260CB7B3213FFDFFF18FBFC7F0B53C1" blockId="16.[128,780,1084,1766]" pageId="16">
Our tree uniquely groups
<taxonomicName id="F5DFB0F83213FFDFFE3BFBFD7E145173" authorityName="Cope" authorityYear="1865" box="[418,533,1084,1108]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="sciadicus">
<emphasis id="00AB17693213FFDFFE3BFBFD7E145173" box="[418,533,1084,1108]" italics="true" pageId="16">F. sciadicus</emphasis>
</taxonomicName>
as sister to
<taxonomicName id="F5DFB0F83213FFDFFD3DFBFC7F0B5173" authorityName="Garman" authorityYear="1895" box="[676,778,1084,1108]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="genus">
<emphasis id="00AB17693213FFDFFD3DFBFC7F0B5173" box="[676,778,1084,1108]" italics="true" pageId="16">Plancterus</emphasis>
</taxonomicName>
(
<taxonomicName id="F5DFB0F83213FFDFFF15FB9B7DC85154" authority="Girard, 1859" authorityName="Girard" authorityYear="1859" box="[140,457,1115,1139]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="kansae">
<emphasis id="00AB17693213FFDFFF15FB9B7D345154" box="[140,309,1115,1139]" italics="true" pageId="16">Fundulus kansae</emphasis>
Girard, 1859
</taxonomicName>
<emphasis id="00AB17693213FFDFFE50FB9C7DD65154" box="[457,471,1116,1139]" italics="true" pageId="16">–</emphasis>
<taxonomicName id="F5DFB0F83213FFDFFE7AFB9B7D1151B4" authority="Jordan &amp; Gilbert, 1883" authorityName="Jordan &amp; Gilbert" authorityYear="1883" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="zebrinus">
<emphasis id="00AB17693213FFDFFE7AFB9B7E9A5154" box="[483,667,1115,1139]" italics="true" pageId="16">Fundulus zebrinus</emphasis>
Jordan &amp; Gilbert, 1883
</taxonomicName>
). Divergence of this group from
<taxonomicName id="F5DFB0F83213FFDFFDF8FBBB7ECB51B5" authorityName="Agassiz" authorityYear="1854" box="[609,714,1147,1170]" class="Actinopterygii" family="Fundulidae" genus="Zygonectes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="genus">
<emphasis id="00AB17693213FFDFFDF8FBBB7ECB51B5" box="[609,714,1147,1170]" italics="true" pageId="16">Zygonectes</emphasis>
</taxonomicName>
20.4– 13.0 Mya (
<figureCitation id="AAE4D7FE3213FFDFFF76FB5A7D275195" box="[239,294,1178,1202]" captionStart="Figure 3" captionStartId="5.[113,178,1835,1859]" captionTargetBox="[192,1381,152,1803]" captionTargetId="figure-7@5.[189,1384,142,1806]" captionTargetPageId="5" captionText="Figure 3. Phylogenetic relationships and divergence times among taxa within North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae recovered by Bayesian inference with five concatenated loci (three mitochondrial and two nuclear) and 130 species.Red dots depict nodes with posterior probabilities &lt;95%. Green arrows show fossil calibration points and brown arrows show secondary calibration points (Table 1). Numbered nodes are associated with environmental and geologic events in our biogeographical synthesis (Table 2)." pageId="16">Fig. 3</figureCitation>
, node 17;
<tableCitation id="7F5DFEC03213FFDFFE09FB5A7DDB5195" box="[400,474,1178,1202]" captionStart="Table 2" captionStartId="6.[129,184,143,167]" captionTargetPageId="6" captionText="Table 2. Divergence estimates with confidence intervals (CIs) for numbered chronogram nodes (Fig.3) that are evaluated in this study. Environmental events potentially associated with phylogenetic divergences (reciprocal illumination) are provided.Further description is provided in the main text, with references." pageId="16">Table 2</tableCitation>
) and a transition from coastal to upland habitat (
<figureCitation id="AAE4D7FE3213FFDFFEA4FB797D7451F6" box="[317,373,1209,1233]" captionStart="Figure 2" captionStartId="4.[129,194,1870,1894]" captionTargetBox="[132,1470,1224,1839]" captionTargetId="figure-539@4.[129,1473,1221,1842]" captionTargetPageId="4" captionText="Figure 2. Reconstruction of ancestral habitats for the North American killifish families Cubanichthyidae, Cyprinodontidae, Fundulidae, Goodeidae, and Profundulidae.A, distribution of North American killifish families with respect to the boundary (red line) between the coastal plain and continental interior (uplands). B, pie charts at each node represent the probability of each character state: coast including coastal plain, upland, or both." pageId="16">Fig. 2</figureCitation>
) suggest that the MRCA of
<emphasis id="00AB17693213FFDFFD09FB7A7CE651D6" italics="true" pageId="16">
<taxonomicName id="F5DFB0F83213FFDFFD09FB7A7EFD51F6" authorityName="Cope" authorityYear="1865" box="[656,764,1209,1233]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="sciadicus">F. sciadicus</taxonomicName>
–
<taxonomicName id="F5DFB0F83213FFDFFF18FB197CE651D6" authorityName="Garman" authorityYear="1895" box="[129,231,1241,1265]" family="Fundulidae" genus="Plancterus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="genus">Plancterus</taxonomicName>
</emphasis>
immigrated up the Mississippi River (
<figureCitation id="AAE4D7FE3213FFDFFDE1FB197EB251D6" box="[632,691,1241,1265]" captionStart="Figure 8" captionStartId="14.[129,194,1478,1502]" captionTargetBox="[129,1473,144,1451]" captionTargetId="figure-7@14.[129,1473,144,1451]" captionTargetPageId="14" captionText="Figure 8. Early–Middle Miocene diversification of North American killifishes in the Gulf of México region.Phylogenetic nodes (Fig.3) are indicated as N5–N7, N9–N11, and N14–N17, showing hypothetical separations between lineages (for details, see main text, Table 2, and Figs 6, 7). The Early Miocene Red River followed the Sabine route to an independent delta, potentially the estuary of origin for subgenus Fundulus, with subgenus Zygonectes arising in the Mississippi River estuary to the east (N15). The Red River joined the Mississippi River in the Middle Miocene.A Guadalupe River drainage emerged to the west in the Early–Middle Miocene, potentially the estuary of origin for the Fundulus similis species group, which diverged from subgenus Fundulus (N16). Widespread, sustained tectonism across southwestern North America isolated western tributaries from the Gulf of México, becoming endorheic drainages, some of which evidently harboured killifishes. Owing to a long history of tectonic activity, the details of these ancient endorheic drainages are unknown.Break-up of the Oligocene Río Grande drainage (Fig. 7) isolated Wileyichthys inland from Lucania (N14). Break-up of the Oligocene Madrean River drainage (Fig. 7) isolated Empetrichthyinae from Goodeinae (N6). Further break-up of the Madrean River separated Goodeinae from Characodontini-Illyodontini (N7). In the southern Gulf, capture of the upper Río Grijalva from an ancestral Río Coatzacoalcos isolated Profundulus from Tlaloc (N5). In the western Gulf, inland immigration of cyprinodontids resulted in peripheral isolates Cualac in the Río Pánuco (N9) and Megupsilon in the Late Miocene Río San Fernando (N11). The base map is a reconstruction for 20 Mya (Early Miocene, used with permission ©2023 Colorado Plateau Geosystems Inc.). The narrowed Gulf Trough follows Popenoe (1990) for Early Miocene.Later sea-level rise flooded the trough and isolated the Ocala High as an island. Two putative dispersal events are illustrated: N10, cyprinodontid eastward coastal immigration, founding Jordanella floridae on the Ocala High; and N17, Fundulus inland immigration up the Mississippi River, founding Fundulus sciadicus–Plancterus on the Northern High Plains." pageId="16">Fig. 8</figureCitation>
), which extended to the nascent Great Plains (
<bibRefCitation id="564EB68A3213FFDFFD81FB387CB45008" author="Snedden JW &amp; Galloway WE" pageId="16" refId="ref20371" refString="Snedden JW, Galloway WE. The Gulf of Mexico Sedimentary Basin: Depositional Evolution and Petroleum Applications. Cambridge: Cambridge University Press, 2019." type="book" year="2019">Snedden and Galloway 2019</bibRefCitation>
).
<taxonomicName id="F5DFB0F83213FFDFFF53FAD77D7F5008" authorityName="Cope" authorityYear="1865" box="[202,382,1303,1327]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="sciadicus">
<emphasis id="00AB17693213FFDFFF53FAD77D7F5008" box="[202,382,1303,1327]" italics="true" pageId="16">Fundulus sciadicus</emphasis>
</taxonomicName>
and
<taxonomicName id="F5DFB0F83213FFDFFE34FAD77E605008" box="[429,609,1303,1327]" pageId="16">
<emphasis id="00AB17693213FFDFFE34FAD77E125008" box="[429,531,1303,1327]" italics="true" pageId="16">Plancterus</emphasis>
species
</taxonomicName>
are grassland associates (
<bibRefCitation id="564EB68A3213FFDFFF78FAF77DBE5068" author="Cross FB &amp; Moss RE" box="[225,447,1335,1359]" pageId="16" pagination="155 - 65" refId="ref15297" refString="Cross FB, Moss RE. Historic changes in fish communities and aquatic habitats in plains streams of Kansas. In: Matthews WJ, Heins DC (eds.), Community and Evolutionary Ecology of North American Stream Fishes. Norman, Oklahoma: University of Oklahoma Press, 1987, 155 - 65." type="book chapter" year="1987">Cross and Moss 1987</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFE52FAF77ED05068" author="Fausch KD &amp; Bestgen KR" box="[459,721,1335,1359]" pageId="16" pagination="131 - 66" refId="ref16279" refString="Fausch KD, Bestgen KR. Ecology of fishes indigenous to the Central and Southwestern great plains BT. In: Knopf FL, Samson FB (eds), Ecology and Conservation of Great Plains Vertebrates. New York: Springer New York, 1997, 131 - 66." type="book chapter" year="1997">Fausch and Bestgen 1997</bibRefCitation>
). We hypothesize that the MRCA diverged from eastern relatives via adaptation to grassland habitats on the northern Great Plains. Accordingly, the 16.6 Mya point estimate for divergence of this group corresponds to a time when the Great Plains region was already ~68% open habitat (
<bibRefCitation id="564EB68A3213FFDFFE3DFA137E6A50CC" author="Edwards EJ &amp; Osborne CP &amp; Stromberg CAE" box="[420,619,1491,1515]" pageId="16" pagination="587 - 91" refId="ref16047" refString="Edwards EJ, Osborne CP, Stromberg CAE et al.; C 4 Grasses Consortium. The origins of C 4 grasslands: integrating evolutionary and ecosystem science. Science 2010; 328: 587 - 91. https: // doi. org / 10.1126 / science. 1177216" type="journal article" year="2010">
Edwards
<emphasis id="00AB17693213FFDFFD9BFA147E3350CC" box="[514,562,1491,1515]" italics="true" pageId="16">et al.</emphasis>
2010
</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFDEEFA137CD5532C" author="Andermann T &amp; Stromberg CAE &amp; Antonelli A" pageId="16" refId="ref14369" refString="Andermann T, Stromberg CAE, Antonelli A et al. The origin and evolution of open habitats in North America inferred by Bayesian deep learning models. Nature Communications 2022; 13: 4833. https: // doi. org / 10.1038 / s 41467 - 022 - 32300 - 5" type="journal volume" year="2022">
Andermann
<emphasis id="00AB17693213FFDFFD61FA147C98532C" italics="true" pageId="16">et al.</emphasis>
2022
</bibRefCitation>
). Also, the Ogallala Formation, which forms the substrate of the Great Plains, was forming at this time (
<bibRefCitation id="564EB68A3213FFDFFDE6F9D27F05530D" author="Chapin CE" box="[639,772,1554,1578]" pageId="16" pagination="976 - 91" refId="ref15100" refString="Chapin CE. Interplay of oceanographic and paleoclimate events with tectonism during middle to late Miocene sedimentation across the Southwestern USA. Geosphere 2008; 4: 976 - 91. https: // doi. org / 10.1130 / ges 00171.1" type="journal article" year="2008">Chapin 2008</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFF18F9F17D51536E" author="Galloway WE &amp; Whiteaker TL &amp; Ganey-Curry P" box="[129,336,1585,1609]" pageId="16" pagination="938 - 73" refId="ref16432" refString="Galloway WE, Whiteaker TL, Ganey-Curry P. History of Cenozoic North American drainage basin evolution, sediment yield, and accumulation in the Gulf of Mexico basin. Geosphere 2011; 7: 938 - 73. https: // doi. org / 10.1130 / ges 00647.1" type="journal article" year="2011">
Galloway
<emphasis id="00AB17693213FFDFFF7EF9F27D17536E" box="[231,278,1585,1609]" italics="true" pageId="16">et al.</emphasis>
2011
</bibRefCitation>
). The 5.0–2.6 Mya fossil †
<taxonomicName id="F5DFB0F83213FFDFFDFAF9F17D8A534E" authority="Hibbard &amp; Dunkle, 1942" authorityName="Hibbard &amp; Dunkle" authorityYear="1942" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="detillae">
<emphasis id="00AB17693213FFDFFDFAF9F17F08536E" box="[611,777,1585,1609]" italics="true" pageId="16">Fundulus detillae</emphasis>
Hibbard &amp; Dunkle, 1942
</taxonomicName>
, recovered from the Ogallala formation in Kansas (
<bibRefCitation id="564EB68A3213FFDFFEBDF9B07E2853AF" author="Ghedotti MJ &amp; Davis MP" box="[292,553,1648,1672]" pageId="16" pagination="577 - 86" refId="ref16637" refString="Ghedotti MJ, Davis MP. The taxonomic placement of three fossil Fundulus species and the timing of divergence within the North American topminnows (Teleostei: Fundulidae). Zootaxa 2017; 4250: 577 - 86. https: // doi. org / 10.11646 / zootaxa. 4250.6.5" type="journal article" year="2017">Ghedotti and Davis 2017</bibRefCitation>
,
<bibRefCitation id="564EB68A3213FFDFFDACF9B07CB45380" author="Cashner MF &amp; Echelle AA" pageId="16" pagination="493 - 502" refId="ref15008" refString="Cashner MF, Echelle AA. Profundulidae: Middle American killifishes. North America, Vol. 2, Characidae to Poeciliidae. Baltimore: Johns Hopkins University Press, 2020, 493 - 502." type="book chapter" year="2020">Cashner and Echelle 2020</bibRefCitation>
), is consistent with the hypothesis that this lineage originated on the northern plains. Altogether, this evidence suggests that the
<taxonomicName id="F5DFB0F83213FFDFFF4DF90F7D4153C1" authorityName="Cope" authorityYear="1865" box="[212,320,1742,1766]" family="Fundulidae" genus="Fundulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="16" phylum="Chordata" rank="species" species="sciadicus">
<emphasis id="00AB17693213FFDFFF4DF90F7D4153C1" box="[212,320,1742,1766]" italics="true" pageId="16">F. sciadicus</emphasis>
</taxonomicName>
group might be a case of ecological isolation.
</paragraph>
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</emphasis>
</treatment>
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125
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<mods:namePart id="239A93AAFAB983DAAEE3C2ABF07040FB">Pham, Hong Thai</mods:namePart>
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<mods:affiliation id="1D1C82B14E64CA7CF9D36E1E9B0DCF40">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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<mods:affiliation id="CBC3018C02DDB5AEE1E11874B61A0973">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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<paragraph id="432E36ED71466239FBADFBE2FC74FBE1" blockId="28.[363,1225,1005,1100]" box="[688,899,1073,1100]" pageId="28" pageNumber="29">
<figureCitation id="DBAA2A6871466239FBADFBE2FD0EFBE6" box="[688,761,1073,1099]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="28" pageNumber="29">Figs 1</figureCitation>
<figureCitation id="DBAA2A6871466239FBADFBE2FD0EFBE6" box="[688,761,1073,1099]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="28" pageNumber="29">Figs 1</figureCitation>
,
<figureCitation id="DBAA2A6871466239FA1AFBE2FCD9FBE6" box="[775,814,1073,1099]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="28" pageNumber="29">2D</figureCitation>
<figureCitation id="DBAA2A6871466239FA1AFBE2FCD9FBE6" box="[775,814,1073,1099]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="28" pageNumber="29">2D</figureCitation>
,
<figureCitation id="DBAA2A6871466239FA27FBE2FC74FBE1" box="[826,899,1073,1100]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="30.[189,232,1928,1954]" captionStartId-1="31.[189,232,1916,1942]" captionStartId-2="33.[189,232,1892,1918]" captionStartId-3="34.[189,232,1963,1989]" captionTargetBox-0="[250,1332,270,1872]" captionTargetBox-1="[189,1398,280,1853]" captionTargetBox-2="[258,1331,266,1845]" captionTargetBox-3="[237,1351,265,1919]" captionTargetId-0="figure-16@30.[243,1344,265,1885]" captionTargetId-1="figure-12@31.[189,1398,270,1862]" captionTargetId-2="figure-11@33.[257,1331,265,1845]" captionTargetId-3="figure-16@34.[237,1351,265,1919]" captionTargetPageId-0="30" captionTargetPageId-1="31" captionTargetPageId-2="33" captionTargetPageId-3="34" captionText-0="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." pageId="28" pageNumber="29">15–18</figureCitation>
<figureCitation id="DBAA2A6871466239FA27FBE2FC74FBE1" box="[826,899,1073,1100]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="30.[189,232,1928,1954]" captionStartId-1="31.[189,232,1916,1942]" captionStartId-2="33.[189,232,1892,1918]" captionStartId-3="34.[189,232,1963,1989]" captionTargetBox-0="[250,1332,270,1872]" captionTargetBox-1="[189,1398,280,1853]" captionTargetBox-2="[258,1331,266,1845]" captionTargetBox-3="[237,1351,265,1919]" captionTargetId-0="figure-16@30.[243,1344,265,1885]" captionTargetId-1="figure-12@31.[189,1398,270,1862]" captionTargetId-2="figure-11@33.[257,1331,265,1845]" captionTargetId-3="figure-16@34.[237,1351,265,1919]" captionTargetPageId-0="30" captionTargetPageId-1="31" captionTargetPageId-2="33" captionTargetPageId-3="34" captionText-0="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." figureDoi-0="http://doi.org/10.5281/zenodo.14508768" figureDoi-1="http://doi.org/10.5281/zenodo.14508770" figureDoi-2="http://doi.org/10.5281/zenodo.14508772" figureDoi-3="http://doi.org/10.5281/zenodo.14508774" httpUri-0="https://zenodo.org/record/14508768/files/figure.png" httpUri-1="https://zenodo.org/record/14508770/files/figure.png" httpUri-2="https://zenodo.org/record/14508772/files/figure.png" httpUri-3="https://zenodo.org/record/14508774/files/figure.png" pageId="28" pageNumber="29">15–18</figureCitation>
</paragraph>
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<subSubSection id="0B8B656671466239F9A0FBA2FC7FF84B" pageId="28" pageNumber="29" type="diagnosis">
@ -376,16 +377,16 @@ leg.;
<paragraph id="432E36ED71476238F9A0FB92FCBEF920" blockId="29.[188,1401,1088,1678]" pageId="29" pageNumber="30">
<smallCapsWord id="45C8A03171476238F9A0FB92FF0BFBF4" baselines="1108,1108" box="[189,252,1089,1115]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="29" pageNumber="30">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A6871476238F813FB93FE69FBF7" box="[270,414,1088,1115]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="29" pageNumber="30">Fig. 16A–D</figureCitation>
<figureCitation id="DBAA2A6871476238F813FB93FE69FBF7" box="[270,414,1088,1115]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508770" httpUri="https://zenodo.org/record/14508770/files/figure.png" pageId="29" pageNumber="30">Fig. 16A–D</figureCitation>
). Narrower than thorax and moderately elongate, with less than half of vertex length surpassing eyes. Vertex variegated brown with median line and basal portion of lateral carinae paler, sometimes also a narrow line behind anterior margin, strongly concave (longitudinally grooved), about 0.88 times as long in midline as broad basally, with sides weakly, evenly converging towards somewhat angular apex; median carina absent; all margins carinate, lateral ones strongly elevated; posterior margin angularly concave. Frons variegated brown, with dense, yellowish, minute pitting, darker along dorsal margin; series of small yellowish tubercles along lateral margins; often a yellowish marking in middle of frons; median carina well marked, extending from dorsal margin to slightly before clypeal suture; dorsal margin rather strongly concave and moderately narrow, 0.34 times as wide as length of frons in midline. Posterior side of head yellowish brown. Genae yellowish with brown to black-brown markings, at anterodorsal angle, anterior to base of antennae, and along clypeal suture. Clypeus more or less coloured as frons with pale median stripe, flat in middle portion, moderately elongate, subtriangular with fronto-clypeal suture rounded; anteclypeus yellowish, brown apically, with median blunt carina. Labium yellowish, elongate and narrow, slightly surpassing mesocoxae, with apical segment elongate, nearly as long as penultimate. Eyes reniform (not emarginate) protruding laterally; ocelli absent. Antennae rather short with scape yellowish, ring-shaped and pedicel dark brown on anterior face and yellowish on posterior one, barrel-shaped, slightly longer than broad.
</paragraph>
<paragraph id="432E36ED71476205F9A0F966FBE5FEC5" blockId="29.[189,1399,1717,2024]" lastBlockId="32.[189,1399,265,360]" lastPageId="32" lastPageNumber="33" pageId="29" pageNumber="30">
<smallCapsWord id="45C8A03171476238F9A0F966FEEEF960" baselines="1736,1736" box="[189,281,1717,1743]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="29" pageNumber="30">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A6871476238F82CF966FE3DF962" box="[305,458,1717,1743]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="29" pageNumber="30">Fig. 16A–D</figureCitation>
<figureCitation id="DBAA2A6871476238F82CF966FE3DF962" box="[305,458,1717,1743]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508770" httpUri="https://zenodo.org/record/14508770/files/figure.png" pageId="29" pageNumber="30">Fig. 16A–D</figureCitation>
). Pronotum with disc (delimited by peridiscal carina) variegated yellowish brown, usually with rather wide median brown band, smooth, with central depression with two brown impressed points; some more or less distinct yellowish tubercles on disc, along posterior margin of lateral fields and at posterodorsal angles of paranotal lobes; wide dark brown area on dorsal portion of lateral fields, continued on most surface of paranotal lobes; paranotal lobes usually with yellowish marking in middle of ventral portion; 0.60 times as long as mesonotum in midline; anterior margin strongly protruding in round angle anteriorly between eyes, with carinae directed obliquely posteriorly, not reaching hind margin of pronotum; posterior margin weakly rounded, slightly elevated; median carina obsolete; paranotal lobes (lateral view) broad with hind margin rounded, angularly rounded posteroventrally. Mesonotum variegated yellowish brown, with paler scutellum; subtriangular with nearly complete transverse anterior carina and with obsolete median carina; peridiscal carina weakly marked; disc smooth, slightly depressed in midline. Tegulae dark brown.
</paragraph>
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<paragraph id="432E36ED7144623BF9A0F85BFE26F845" blockId="30.[189,1398,1928,2024]" pageId="30" pageNumber="31">
<emphasis id="71E5EAFF7144623BF9A0F85BFEECF80F" bold="true" box="[189,283,1928,1954]" pageId="30" pageNumber="31">Fig. 15.</emphasis>
<taxonomicName id="84914D6E7144623BF83EF85BFE20F80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[291,471,1928,1954]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="30" pageNumber="31" phylum="Arthropoda" rank="species" species="quangi" status="sp. nov.">
@ -405,7 +406,7 @@ leg.;
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
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<paragraph id="432E36ED7145623AF9A0F8AFFA9CF870" blockId="31.[188,1398,1916,2014]" pageId="31" pageNumber="32">
<emphasis id="71E5EAFF7145623AF9A0F8AFFEEEF83B" bold="true" box="[189,281,1916,1942]" pageId="31" pageNumber="32">Fig. 16.</emphasis>
<taxonomicName id="84914D6E7145623AF802F8AEFE26F83B" authorityName="Constant &amp; Pham" authorityYear="2024" box="[287,465,1917,1943]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="31" pageNumber="32" phylum="Arthropoda" rank="species" species="quangi" status="sp. nov.">
@ -431,11 +432,11 @@ leg.;
<paragraph id="432E36ED717A6205F9A0FE43FDAEFCA9" blockId="32.[187,1400,399,772]" pageId="32" pageNumber="33">
<smallCapsWord id="45C8A031717A6205F9A0FE43FED5FE05" baselines="419,419" box="[189,290,400,426]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="32" pageNumber="33">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A68717A6205F82AFE5CFE39FE04" box="[311,462,399,426]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="32" pageNumber="33">Figs 15A–C</figureCitation>
<figureCitation id="DBAA2A68717A6205F82AFE5CFE39FE04" box="[311,462,399,426]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508768" httpUri="https://zenodo.org/record/14508768/files/figure.png" pageId="32" pageNumber="33">Figs 15A–C</figureCitation>
,
<figureCitation id="DBAA2A68717A6205F8C3FE5CFE08FE04" box="[478,511,399,425]" captionStart="Fig" captionStartId="34.[189,232,1963,1989]" captionTargetBox="[237,1351,265,1919]" captionTargetId="figure-16@34.[237,1351,265,1919]" captionTargetPageId="34" captionText="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." pageId="32" pageNumber="33">18</figureCitation>
<figureCitation id="DBAA2A68717A6205F8C3FE5CFE08FE04" box="[478,511,399,425]" captionStart="Fig" captionStartId="34.[189,232,1963,1989]" captionTargetBox="[237,1351,265,1919]" captionTargetId="figure-16@34.[237,1351,265,1919]" captionTargetPageId="34" captionText="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." figureDoi="http://doi.org/10.5281/zenodo.14508774" httpUri="https://zenodo.org/record/14508774/files/figure.png" pageId="32" pageNumber="33">18</figureCitation>
). Variegated yellowish brown, sometimes washed with green, with more or less extensive, scattered, irregular blackish brown markings; sometimes with more or less distinct subtriangular paler area before midlength; tegmina subcoriaceous with longitudinal veins elevated and with dense reticulum of paler veinlets; shape elongate and convex with sides broadly rounded, subparallel in middle portion, about 2.5 times as long as wide; rather narrowly rounded apically. Postclaval margin straight, slightly notched at apex of clavus. Clavus closed, reaching about 3/5 of tegmen. Venation (
<figureCitation id="DBAA2A68717A6205F9DBFD8CFEABFDD4" box="[198,348,607,634]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="32" pageNumber="33">Fig. 15A, C</figureCitation>
<figureCitation id="DBAA2A68717A6205F9DBFD8CFEABFDD4" box="[198,348,607,634]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508768" httpUri="https://zenodo.org/record/14508768/files/figure.png" pageId="32" pageNumber="33">Fig. 15A, C</figureCitation>
): ScP+R moderately developed, forking into subparallel ScP+RA and RP; MP forking rather close to base into diverging MP
<subScript id="DF1534A8717A6205FB6AFD46FD77FD0E" attach="left" box="[631,640,661,675]" fontSize="6" pageId="32" pageNumber="33">1</subScript>
and MP
@ -456,19 +457,19 @@ running subprallel to MP
<smallCapsWord id="45C8A031717A6205F9A0FCF8FF00FCE9" baselines="830,831" box="[189,247,811,837]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="32" pageNumber="33">HIND</smallCapsWord>
<smallCapsWord id="45C8A031717A6205F81CFCE3FEB2FCEE" baselines="830" box="[257,325,816,835]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="32" pageNumber="33">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A68717A6205F844FCF8FE32FCE8" box="[345,453,811,837]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="32" pageNumber="33">Fig. 16E</figureCitation>
<figureCitation id="DBAA2A68717A6205F844FCF8FE32FCE8" box="[345,453,811,837]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508770" httpUri="https://zenodo.org/record/14508770/files/figure.png" pageId="32" pageNumber="33">Fig. 16E</figureCitation>
). Dark brown with paler areas along costal margin following vein ScP+R, and along basal portion of Pcu; veins mostly coloured darker than background; wing broader than tegmen and deeply bilobed at CuP; costal margin rather weakly sinuate; CuP-Pcu-A1 lobe nearly 1.5 times as wide as ScP-R-MP-CuA lobe, the two lobes almost the same length; both lobes somewhat angularly rounded apically; postclaval margin broadly rounded; A2 lobe brown, reduced and narrow. Venation: main veins present, dense network of cross-veins; ScP+R parallel to costal margin; MP and CuA running more or less parallel; Pcu rather strongly curved before basal half of wing towards CuP; A1 curved, more or less parallel to postclaval margin; vein A2 distinct.
</paragraph>
<paragraph id="432E36ED717A6205F9A0FB8CFE27FA7E" blockId="32.[186,1400,1118,1491]" pageId="32" pageNumber="33">
<smallCapsWord id="45C8A031717A6205F9A0FB8CFF01FBDA" baselines="1138,1138" box="[189,246,1119,1145]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="32" pageNumber="33">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A68717A6205F815FB8DFE61FBD4" box="[264,406,1118,1145]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="32" pageNumber="33">Fig. 15A–E</figureCitation>
<figureCitation id="DBAA2A68717A6205F815FB8DFE61FBD4" box="[264,406,1118,1145]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508768" httpUri="https://zenodo.org/record/14508768/files/figure.png" pageId="32" pageNumber="33">Fig. 15A–E</figureCitation>
). Coxae yellowish with anterior margin brown; trochanters variegated yellowish and brown. Pro- and mesofemora yellowish with brown wide median, and anteapical rings; ventral face darker than dorsal; elongate, rather slender and slightly flattened dorsoventrally. Pro- and mesotibiae yellowish with brown more or less distinct subbasal ring; apex brown; tibiae weakly flattened dorsoventrally, narrower and longer than corresponding femora. Pro- and mesotarsi elongate and brown. Metafemora yellowish with ventral apical marking and elongate marking along anterior face, brown. Metatibiae yellowish, with one lateral spine near base (sometimes weak to obsolete), 2 lateral spines in distal half, and 8 apical spines, all spines brown, apically black. First metatarsomere elongate and slender, with a strong spine at each side and a row of 6 smaller spines in between ventrally along posterior margin; second metatarsomere short with one spine at each side, all spines black-brown apically. Metatibiotarsal formula: (2–3) 8 / 8 / 2.
</paragraph>
<paragraph id="432E36ED717A6205F9A0FA29FC80F9B9" blockId="32.[189,887,1530,1557]" box="[189,887,1530,1557]" pageId="32" pageNumber="33">
<smallCapsWord id="45C8A031717A6205F9A0FA29FEDBF9BF" baselines="1549,1549" box="[189,300,1530,1556]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Abdomen" pageId="32" pageNumber="33">ABDOMEN</smallCapsWord>
(
<figureCitation id="DBAA2A68717A6205F823FA29FE5FF9B8" box="[318,424,1530,1557]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="32" pageNumber="33">Fig. 15B</figureCitation>
<figureCitation id="DBAA2A68717A6205F823FA29FE5FF9B8" box="[318,424,1530,1557]" captionStart="Fig" captionStartId="30.[189,232,1928,1954]" captionTargetBox="[250,1332,270,1872]" captionTargetId="figure-16@30.[243,1344,265,1885]" captionTargetPageId="30" captionText="Fig. 15. Flavina quangi sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508768" httpUri="https://zenodo.org/record/14508768/files/figure.png" pageId="32" pageNumber="33">Fig. 15B</figureCitation>
). Brown with basal segment yellowish.
</paragraph>
<paragraph id="432E36ED717A6206F9A0F9E8FB5FFE03" blockId="32.[187,1400,1593,2002]" lastBlockId="35.[189,1399,265,430]" lastPageId="35" lastPageNumber="36" pageId="32" pageNumber="33">
@ -477,17 +478,17 @@ running subprallel to MP
. Pygofer (
<emphasis id="71E5EAFF717A6205FB11F9EFFDDCF9F8" box="[524,555,1596,1621]" italics="true" pageId="32" pageNumber="33">Py</emphasis>
–
<figureCitation id="DBAA2A68717A6205FB50F9E8FD39F9F8" box="[589,718,1595,1621]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="32" pageNumber="33">Fig. 9A–D</figureCitation>
<figureCitation id="DBAA2A68717A6205FB50F9E8FD39F9F8" box="[589,718,1595,1621]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="32" pageNumber="33">Fig. 9A–D</figureCitation>
) massive, about 1.8 times as high as long (at ventral ⅓) in lateral view, with anterior margin weakly sinuate and posterior margin, along dorsal ⅔ more or less oblique and weakly concave then abruptly rounded at posterodorsal angle, in ventral ⅓, rather strongly excavate; posterior margin dorsally with deep, open-U-shaped excavation. Gonostyli (
<emphasis id="71E5EAFF717A6205FD82F970FB43F910" box="[1183,1204,1699,1725]" italics="true" pageId="32" pageNumber="33">G</emphasis>
–
<figureCitation id="DBAA2A68717A6205FDD0F970FABFF910" box="[1229,1352,1699,1725]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="32" pageNumber="33">Fig. 9A–C</figureCitation>
<figureCitation id="DBAA2A68717A6205FDD0F970FABFF910" box="[1229,1352,1699,1725]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="32" pageNumber="33">Fig. 9A–C</figureCitation>
) (in lateral view) longer than high (without dorsal capitulum), projecting posteriorly in distinct elongate lobe rounded apically and with ventral margin broadly rounded; capitulum (
<emphasis id="71E5EAFF717A6205FD1FF93AFBE9F8AF" box="[1026,1054,1769,1794]" italics="true" pageId="32" pageNumber="33">ca</emphasis>
–
<figureCitation id="DBAA2A68717A6205FD26F93BFB4FF8AF" box="[1083,1208,1768,1794]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="32" pageNumber="33">Fig. 9A–C</figureCitation>
<figureCitation id="DBAA2A68717A6205FD26F93BFB4FF8AF" box="[1083,1208,1768,1794]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="32" pageNumber="33">Fig. 9A–C</figureCitation>
) massive, wide, subtriangular (about as high as wide), slightly twisted, dorsally hooked cephalodorsad, and with wide, rather short neck with posterior margin rounded, and with basal lamina projection directed lateroventrad in caudal view. Aedeagus (
<figureCitation id="DBAA2A68717A6205FB14F883FD76F8C7" box="[521,641,1872,1898]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="32" pageNumber="33">Fig. 9E–J</figureCitation>
<figureCitation id="DBAA2A68717A6205FB14F883FD76F8C7" box="[521,641,1872,1898]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="32" pageNumber="33">Fig. 9E–J</figureCitation>
) symmetrical, evenly curved dorsad (in lateral view); pair of lateroventral processes (
<emphasis id="71E5EAFF717A6205F8C6F8A0FE08F820" box="[475,511,1907,1933]" italics="true" pageId="32" pageNumber="33">lvp</emphasis>
) elongate, rather wide in basal portion and directed mesocephalad (weakly sinuate), then tapering, curved laterocephalad and pointed apically; ventral lobe of periandrium (
@ -503,10 +504,10 @@ running subprallel to MP
) well developed, widely open. Anal tube (
<emphasis id="71E5EAFF71796206FC23FEFFFAA8FEEB" box="[1342,1375,300,326]" italics="true" pageId="35" pageNumber="36">An</emphasis>
–
<figureCitation id="DBAA2A6871796206F9A2FE9DFECBFEC5" box="[191,316,334,360]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="35" pageNumber="36">Fig. 9A–D</figureCitation>
<figureCitation id="DBAA2A6871796206F9A2FE9DFECBFEC5" box="[191,316,334,360]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="35" pageNumber="36">Fig. 9A–D</figureCitation>
) dorsoventrally flattened, in dorsal view widening from base towards apex in proximal half, then abruptly narrowing with apical margin truncate (slightly concave); lateral lobes well developed, angularly rounded apically and directed posteroventrad; anal opening before basal ½.
</paragraph>
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<emphasis id="71E5EAFF717B6204F9A0F8B7FEEDF8D2" bold="true" box="[189,282,1892,1919]" pageId="33" pageNumber="34">Fig. 17.</emphasis>
<taxonomicName id="84914D6E717B6204F83CF8B6FE23F8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[289,468,1893,1919]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="33" pageNumber="34" phylum="Arthropoda" rank="species" species="quangi" status="sp. nov.">
@ -542,7 +543,7 @@ running subprallel to MP
. Anteroventral view. Abbreviations: see Material and methods.
</paragraph>
</caption>
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<emphasis id="71E5EAFF71786207F9A0F878FEEEF868" bold="true" box="[189,281,1963,1989]" pageId="34" pageNumber="35">Fig. 18.</emphasis>
<taxonomicName id="84914D6E71786207F802F878FE26F868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[287,465,1963,1989]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="34" pageNumber="35" phylum="Arthropoda" rank="species" species="quangi" status="sp. nov.">
@ -568,7 +569,7 @@ running subprallel to MP
. Hind margin of sternum VII (
<emphasis id="71E5EAFF71796206FA06FE06FC96FE43" box="[795,865,469,495]" italics="true" pageId="35" pageNumber="36">St VII</emphasis>
–
<figureCitation id="DBAA2A6871796206FA99FE07FC1CFE43" box="[900,1003,468,494]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="35" pageNumber="36">Fig. 16F</figureCitation>
<figureCitation id="DBAA2A6871796206FA99FE07FC1CFE43" box="[900,1003,468,494]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508770" httpUri="https://zenodo.org/record/14508770/files/figure.png" pageId="35" pageNumber="36">Fig. 16F</figureCitation>
) concave, moderately inflated in middle portion.
</paragraph>
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@ -586,13 +587,13 @@ running subprallel to MP
was collected in May and September, in montane evergreen tropical forest, at about
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a.s.l., near the summit (
<figureCitation id="DBAA2A6871796206FBD4FD5AFCD0FD0E" box="[713,807,649,675]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="35" pageNumber="36">Fig. 2D</figureCitation>
<figureCitation id="DBAA2A6871796206FBD4FD5AFCD0FD0E" box="[713,807,649,675]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="35" pageNumber="36">Fig. 2D</figureCitation>
), and in September in lowland evergreen forest in Bach Ma, at altitudes around
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a.s.l. (
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<figureCitation id="DBAA2A6871796206FBE1FD78FCAAFD6B" box="[764,861,683,710]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="35" pageNumber="36">Fig. 2A</figureCitation>
). The species seemed rather common at that moment and specimens were collected on tree trunks densely covered in lichens and mosses while a few were also found sitting on leaves of lower plants and bushes (
<figureCitation id="DBAA2A6871796206FA8EFD22FC1CFCA6" box="[915,1003,753,779]" captionStart="Fig" captionStartId="34.[189,232,1963,1989]" captionTargetBox="[237,1351,265,1919]" captionTargetId="figure-16@34.[237,1351,265,1919]" captionTargetPageId="34" captionText="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." pageId="35" pageNumber="36">Fig. 18</figureCitation>
<figureCitation id="DBAA2A6871796206FA8EFD22FC1CFCA6" box="[915,1003,753,779]" captionStart="Fig" captionStartId="34.[189,232,1963,1989]" captionTargetBox="[237,1351,265,1919]" captionTargetId="figure-16@34.[237,1351,265,1919]" captionTargetPageId="34" captionText="Fig. 18. Flavina quangi sp. nov., live specimens in Bach Ma National Park, summit. A–B. 14 May 2023. C–E. 21 May 2023. F. 14 May 2023. G. 13 May 2023. H. 11 May 2023." figureDoi="http://doi.org/10.5281/zenodo.14508774" httpUri="https://zenodo.org/record/14508774/files/figure.png" pageId="35" pageNumber="36">Fig. 18</figureCitation>
). However, previous fieldwork in
<date id="372F102D71796206F9A0FCC0FED8FC83" box="[189,303,787,814]" pageId="35" pageNumber="36" value="2011-07">July 2011</date>
and
@ -609,7 +610,7 @@ at the same locality did not provide any specimens. When disturbed, they tend to
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<collectingCountry id="3B86767D71796206F9A0FC76FED2FC12" box="[189,293,933,959]" name="Vietnam" pageId="35" pageNumber="36">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, Bach Ma National Park (
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<figureCitation id="DBAA2A6871796206FA88FC76FC2DFC12" box="[917,986,933,959]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="35" pageNumber="36">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>

117
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<mods:identifier id="29AD34E40BB93C84FE214450304FFBA3" type="ZooBank">23F0D60C-3B5B-4AB1-B608-AD875EB424DF</mods:identifier>
</mods:mods>
<treatment id="CB3887FB714C6239FB86F922FB9DFC1C" LSID="urn:lsid:plazi:treatment:CB3887FB714C6239FB86F922FB9DFC1C" httpUri="http://treatment.plazi.org/id/CB3887FB714C6239FB86F922FB9DFC1C" lastPageId="28" lastPageNumber="29" pageId="22" pageNumber="23">
<subSubSection id="0B8B6566714C6233FB86F922FB37F89D" pageId="22" pageNumber="23" type="nomenclature">
@ -67,11 +68,11 @@ urn:lsid:zoobank.org:act:
</subSubSection>
<subSubSection id="0B8B6566714C6233FBADF8EBFC74F8FF" box="[688,899,1848,1874]" pageId="22" pageNumber="23" type="description">
<paragraph id="432E36ED714C6233FBADF8EBFC74F8FF" blockId="22.[372,1216,1777,1874]" box="[688,899,1848,1874]" pageId="22" pageNumber="23">
<figureCitation id="DBAA2A68714C6233FBADF8EBFD0EF8FF" box="[688,761,1848,1874]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="22" pageNumber="23">Figs 1</figureCitation>
<figureCitation id="DBAA2A68714C6233FBADF8EBFD0EF8FF" box="[688,761,1848,1874]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="22" pageNumber="23">Figs 1</figureCitation>
,
<figureCitation id="DBAA2A68714C6233FA1AF8EBFCD9F8FF" box="[775,814,1848,1874]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="22" pageNumber="23">2D</figureCitation>
<figureCitation id="DBAA2A68714C6233FA1AF8EBFCD9F8FF" box="[775,814,1848,1874]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="22" pageNumber="23">2D</figureCitation>
,
<figureCitation id="DBAA2A68714C6233FA27F8EBFC74F8FF" box="[826,899,1848,1874]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="24.[189,232,1928,1954]" captionStartId-1="26.[191,234,1928,1954]" captionStartId-2="27.[189,232,1892,1918]" captionTargetBox-0="[251,1336,272,1884]" captionTargetBox-1="[189,1398,444,1885]" captionTargetBox-2="[254,1333,265,1852]" captionTargetId-0="figure-16@24.[243,1344,265,1885]" captionTargetId-1="figure-65@26.[189,1398,436,1886]" captionTargetId-2="figure-12@27.[254,1333,265,1852]" captionTargetPageId-0="24" captionTargetPageId-1="26" captionTargetPageId-2="27" captionText-0="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="22" pageNumber="23">12–14</figureCitation>
<figureCitation id="DBAA2A68714C6233FA27F8EBFC74F8FF" box="[826,899,1848,1874]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="24.[189,232,1928,1954]" captionStartId-1="26.[191,234,1928,1954]" captionStartId-2="27.[189,232,1892,1918]" captionTargetBox-0="[251,1336,272,1884]" captionTargetBox-1="[189,1398,444,1885]" captionTargetBox-2="[254,1333,265,1852]" captionTargetId-0="figure-16@24.[243,1344,265,1885]" captionTargetId-1="figure-65@26.[189,1398,436,1886]" captionTargetId-2="figure-12@27.[254,1333,265,1852]" captionTargetPageId-0="24" captionTargetPageId-1="26" captionTargetPageId-2="27" captionText-0="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508762" figureDoi-1="http://doi.org/10.5281/zenodo.14508764" figureDoi-2="http://doi.org/10.5281/zenodo.14508766" httpUri-0="https://zenodo.org/record/14508762/files/figure.png" httpUri-1="https://zenodo.org/record/14508764/files/figure.png" httpUri-2="https://zenodo.org/record/14508766/files/figure.png" pageId="22" pageNumber="23">12–14</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="0B8B6566714C6232F9A0F8AAFC97FB9D" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23" type="diagnosis">
@ -231,10 +232,10 @@ leg.; day [time] collecting; I.G.: 31.933;
<paragraph id="432E36ED714D623CF9A0F985FD2BFECB" blockId="23.[188,1401,1621,2018]" lastBlockId="25.[189,1400,265,359]" lastPageId="25" lastPageNumber="26" pageId="23" pageNumber="24">
<smallCapsWord id="45C8A031714D6232F9A0F985FF0BF9C3" baselines="1641,1641" box="[189,252,1622,1648]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="23" pageNumber="24">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A68714D6232F813F986FE68F9DD" box="[270,415,1621,1648]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="23" pageNumber="24">Fig. 13A–D</figureCitation>
<figureCitation id="DBAA2A68714D6232F813F986FE68F9DD" box="[270,415,1621,1648]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508764" httpUri="https://zenodo.org/record/14508764/files/figure.png" pageId="23" pageNumber="24">Fig. 13A–D</figureCitation>
). Narrower than thorax and moderately elongate, with less than half of vertex length surpassing eyes. Vertex variegated brown with median line and basal portion of lateral carinae paler, strongly concave (longitudinally grooved), about 0.85 times as long in midline as broad basally, with sides weakly, evenly converging towards somewhat angular apex; median carina absent; all margins carinate, lateral ones strongly elevated; posterior margin angularly concave. Frons variegated brown, darker along dorsal margin; series of small yellowish tubercles along lateral margins; median carina well marked, extending from dorsal margin to slightly before clypeal suture; dorsal margin concave and moderately narrow, 0.35 times as wide as length of frons in midline. Posterior side of head yellowish. Genae yellowish with black-brown marking at anterodorsal angle and anterior to base of antennae. Clypeus coloured as frons, flat in middle portion, moderately elongate, subtriangular with fronto-clypeal suture rounded; anteclypeus yellowish, brown apically, with median blunt carina. Labium yellowish, elongate and narrow, nearly reaching metacoxae, with apical segment elongate, nearly as long as penultimate. Eyes reniform (not emarginate) protruding laterally; ocelli absent. Antennae rather short with scape yellowish, ring-shaped and pedicel dark brown on anterior face and yellowish on posterior one, barrel-shaped, slightly longer than broad.
</paragraph>
<caption id="17EE66657142623DF9A0F85BFE26F845" pageId="24" pageNumber="25" startId="24.[189,232,1928,1954]" targetBox="[251,1336,272,1884]" targetPageId="24">
<caption id="17EE66657142623DF9A0F85BFE26F845" ID-DOI="http://doi.org/10.5281/zenodo.14508762" ID-Zenodo-Dep="14508762" httpUri="https://zenodo.org/record/14508762/files/figure.png" pageId="24" pageNumber="25" startId="24.[189,232,1928,1954]" targetBox="[251,1336,272,1884]" targetPageId="24">
<paragraph id="432E36ED7142623DF9A0F85BFE26F845" blockId="24.[189,1398,1928,2024]" pageId="24" pageNumber="25">
<emphasis id="71E5EAFF7142623DF9A0F85BFEEAF80F" bold="true" box="[189,285,1928,1954]" pageId="24" pageNumber="25">Fig. 12.</emphasis>
<taxonomicName id="84914D6E7142623DF83AF85BFE37F80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[295,448,1928,1954]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="species" species="lami" status="sp. nov.">
@ -257,15 +258,15 @@ leg.; day [time] collecting; I.G.: 31.933;
<paragraph id="432E36ED7143623CF9A0FE5EFC5CFD79" blockId="25.[185,1401,396,724]" pageId="25" pageNumber="26">
<smallCapsWord id="45C8A0317143623CF9A0FE5EFEEEFE08" baselines="416,416" box="[189,281,397,423]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="25" pageNumber="26">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A687143623CF83AFE5FFE42FE0B" box="[295,437,396,423]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="25" pageNumber="26">Fig. 13A–D</figureCitation>
<figureCitation id="DBAA2A687143623CF83AFE5FFE42FE0B" box="[295,437,396,423]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508764" httpUri="https://zenodo.org/record/14508764/files/figure.png" pageId="25" pageNumber="26">Fig. 13A–D</figureCitation>
). Pronotum with disc (delimited by peridiscal carina) yellowish, smooth, with central depression with two brown impressed points; rather wide dark brown area on dorsal portion of lateral fields, continued along posterior margin of paranotal lobes; yellowish tubercles along posterior margin of paranotal lobes; 0.71 times as long as mesonotum in midline; anterior margin strongly protruding in round angle anteriorly between eyes, with carinae directed obliquely posteriorly, not reaching hind margin of pronotum; posterior margin nearly straight, slightly elevated; median carina obsolete; paranotal lobes (lateral view) broad with hind margin rounded, angularly rounded posteroventrally. Mesonotum variegated yellowish, with brown marking at base of scutellum; subtriangular with nearly complete transverse anterior carina and with obsolete median carina; peridiscal carina weakly marked; disc smooth, slightly depressed in midline. Tegulae dark brown.
</paragraph>
<paragraph id="432E36ED7143623CF9A0FD28FE37FBC9" blockId="25.[188,1399,762,1125]" pageId="25" pageNumber="26">
<smallCapsWord id="45C8A0317143623CF9A0FD28FED5FCBE" baselines="782,782" box="[189,290,763,789]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="25" pageNumber="26">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A687143623CF825FD29FE3BFCB9" box="[312,460,762,789]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="25" pageNumber="26">Fig. 12A–C</figureCitation>
<figureCitation id="DBAA2A687143623CF825FD29FE3BFCB9" box="[312,460,762,789]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508762" httpUri="https://zenodo.org/record/14508762/files/figure.png" pageId="25" pageNumber="26">Fig. 12A–C</figureCitation>
). Variegated pale yellowish, with more or less extensive, scattered, irregular blackish brown markings, larger ones at base of costal cell and at ⅔ of length along costal margin; tegmina subcoriaceous with longitudinal veins elevated and with dense reticulum of pale veinlets; shape elongate and convex with sides subparallel (weakly bisinuate) in middle portion, about 2.5 times as long as wide; angularly rounded apically. Postclaval margin straight, slightly notched at apex of clavus. Clavus closed, slightly surpassing half of tegmen length. Venation: (
<figureCitation id="DBAA2A687143623CFAEEFC71FB72FC11" box="[1011,1157,930,956]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="25" pageNumber="26">Fig. 12A, C</figureCitation>
<figureCitation id="DBAA2A687143623CFAEEFC71FB72FC11" box="[1011,1157,930,956]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508762" httpUri="https://zenodo.org/record/14508762/files/figure.png" pageId="25" pageNumber="26">Fig. 12A, C</figureCitation>
) ScP+R moderately developed, forking into subparallel ScP+RA and RP; MP forking rather close to base into diverging MP
<subScript id="DF1534A87143623CF9FAFC2BFF07FBAB" attach="left" box="[231,240,1016,1030]" fontSize="6" pageId="25" pageNumber="26">1</subScript>
and MP
@ -284,13 +285,13 @@ separated at about 2/5 of tegmen length, then running parallel to MP
<smallCapsWord id="45C8A0317143623CF9A0FB58FF00FB0E" baselines="1182,1182" box="[189,247,1163,1189]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="25" pageNumber="26">HIND</smallCapsWord>
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(
<figureCitation id="DBAA2A687143623CF84BFB59FE48FB09" box="[342,447,1162,1189]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="25" pageNumber="26">Fig. 13E</figureCitation>
<figureCitation id="DBAA2A687143623CF84BFB59FE48FB09" box="[342,447,1162,1189]" captionStart="Fig" captionStartId="26.[191,234,1928,1954]" captionTargetBox="[189,1398,444,1885]" captionTargetId="figure-65@26.[189,1398,436,1886]" captionTargetPageId="26" captionText="Fig. 13. Flavina lami sp. nov., holotype, ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508764" httpUri="https://zenodo.org/record/14508764/files/figure.png" pageId="25" pageNumber="26">Fig. 13E</figureCitation>
). Dark brown with paler brown area at base of Pcu, and narrow area along basal portion of ScP+R; veins mostly black, darker than background; wing broader than tegmen and deeply bilobed at CuP; costal margin rather weakly sinuate; CuP-Pcu-A1 lobe nearly twice as wide as ScP-R-MP-CuA lobe, the two lobes almost the same length; both lobes somewhat angularly rounded apically; postclaval margin broadly rounded; A2 lobe brown, reduced and narrow. Venation: main veins present; dense network of cross-veins; ScP+R and MP diverging in basal portion, then running subparallel; CuA running more or less straight; Pcu rather strongly curved around basal third of wing towards CuP; A1 weakly curved, more or less parallel to postclaval margin; vein A2 distinct.
</paragraph>
<paragraph id="432E36ED7143623CF9A0FA65FBB5F8B2" blockId="25.[187,1401,1461,1823]" pageId="25" pageNumber="26">
<smallCapsWord id="45C8A0317143623CF9A0FA65FF01FA63" baselines="1481,1481" box="[189,246,1462,1488]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="25" pageNumber="26">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A687143623CF817FA66FE6DFA62" box="[266,410,1461,1488]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="25" pageNumber="26">Fig. 12A–E</figureCitation>
<figureCitation id="DBAA2A687143623CF817FA66FE6DFA62" box="[266,410,1461,1488]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508762" httpUri="https://zenodo.org/record/14508762/files/figure.png" pageId="25" pageNumber="26">Fig. 12A–E</figureCitation>
). Pro- and mesocoxae yellowish, mesoanteriorly margined with brown; metacoxae brown; trochanters variegated yellowish and brown. Pro- and mesofemora yellow-brown with anteapical brown ring followed by yellowish ring before narrowly brown apex; ventral face darker than dorsal; elongate, rather slender and slightly flattened dorsoventrally. Pro- and mesotibiae yellowish with apex and subbasal ring brown; ventral face darker than dorsal; protibiae more contrasted; tibiae weakly flattened dorsoventrally, narrower and longer than corresponding femora. Pro- and mesotarsi elongate and brown. Metafemora yellowish with elongate brown marking along anteroventral portion. Metatibiae yellowish; lateral spines: 1 subbasal and
<quantity id="84699B087143623CFBBBF973FD05F917" box="[678,754,1696,1722]" metricMagnitude="-2" metricUnit="m" metricValue="6.35" metricValueMax="7.62" metricValueMin="5.08" pageId="25" pageNumber="26" unit="in" value="2.5" valueMax="3.0" valueMin="2.0">2–3 in</quantity>
distal half; 8 apical spines, all spines brown, apically black. First metatarsomere elongate and slender, with a strong spine at each side and a row of 6 smaller spines in between ventrally along posterior margin; second metatarsomere short with one spine at each side, all spines black-brown apically. Metatibiotarsal formula: (3–4) 8 / 8 / 2.
@ -298,7 +299,7 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
<paragraph id="432E36ED7143623CF9A0F896FCA4F8F2" blockId="25.[189,851,1861,1887]" box="[189,851,1861,1887]" pageId="25" pageNumber="26">
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(
<figureCitation id="DBAA2A687143623CF823F896FE5EF8F2" box="[318,425,1861,1887]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="25" pageNumber="26">Fig. 12B</figureCitation>
<figureCitation id="DBAA2A687143623CF823F896FE5EF8F2" box="[318,425,1861,1887]" captionStart="Fig" captionStartId="24.[189,232,1928,1954]" captionTargetBox="[251,1336,272,1884]" captionTargetId="figure-16@24.[243,1344,265,1885]" captionTargetPageId="24" captionText="Fig. 12. Flavina lami sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508762" httpUri="https://zenodo.org/record/14508762/files/figure.png" pageId="25" pageNumber="26">Fig. 12B</figureCitation>
). Dark brown, slightly paler basally.
</paragraph>
<paragraph id="432E36ED71436239F9A0F856FDF9FDDE" blockId="25.[188,1400,1925,2019]" lastBlockId="28.[187,1399,265,627]" lastPageId="28" lastPageNumber="29" pageId="25" pageNumber="26">
@ -307,17 +308,17 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
. Pygofer (
<emphasis id="71E5EAFF7143623CFB1FF855FDD6F832" box="[514,545,1926,1951]" italics="true" pageId="25" pageNumber="26">Py</emphasis>
–
<figureCitation id="DBAA2A687143623CFB21F856FD3EF832" box="[572,713,1925,1951]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="25" pageNumber="26">Fig. 14A–D</figureCitation>
<figureCitation id="DBAA2A687143623CFB21F856FD3EF832" box="[572,713,1925,1951]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="25" pageNumber="26">Fig. 14A–D</figureCitation>
) about 2.0 times as high as long (slightly above midheight) in lateral view, with anterior margin nearly straight and posterior margin forming a wide lobe rounded to posterodorsal angle; posterior margin dorsally with deep, V-shaped, rounded excavation. Gonostyli (
<emphasis id="71E5EAFF7140623FF9DAFEDAFF2BFE8E" box="[199,220,265,291]" italics="true" pageId="26" pageNumber="27">G</emphasis>
–
<figureCitation id="DBAA2A687140623FF9E1FEDAFE7DFE8E" box="[252,394,265,291]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="26" pageNumber="27">Fig. 14A–C</figureCitation>
<figureCitation id="DBAA2A687140623FF9E1FEDAFE7DFE8E" box="[252,394,265,291]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="26" pageNumber="27">Fig. 14A–C</figureCitation>
) (in lateral view) longer than high (without dorsal capitulum), projecting posteriorly in distinct elongate lobe rounded apically and with ventral margin weakly rounded; capitulum (
<emphasis id="71E5EAFF7140623FFC5FFEF8FAA9FEE9" box="[1346,1374,299,324]" italics="true" pageId="26" pageNumber="27">ca</emphasis>
–
<figureCitation id="DBAA2A687140623FF9A0FE9FFEBCFECB" box="[189,331,332,358]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="26" pageNumber="27">Fig. 14A–C</figureCitation>
<figureCitation id="DBAA2A687140623FF9A0FE9FFEBCFECB" box="[189,331,332,358]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="26" pageNumber="27">Fig. 14A–C</figureCitation>
) directed dorsad, rather massive, wide, falcate and short (about as high as wide), slightly twisted, with moderately long, wide neck, and with apical hook directed dorsocephalad; with basal lamina projection directed lateroventrad in caudal view. Aedeagus (
<figureCitation id="DBAA2A6871466239FAF3FEF9FB84FEE8" box="[1006,1139,298,325]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="28" pageNumber="29">Fig. 14E–J</figureCitation>
<figureCitation id="DBAA2A6871466239FAF3FEF9FB84FEE8" box="[1006,1139,298,325]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="28" pageNumber="29">Fig. 14E–J</figureCitation>
) symmetrical, evenly curved dorsad (in lateral view); pair of lateroventral processes (
<emphasis id="71E5EAFF71466239FACCFE9FFC02FECB" box="[977,1013,332,358]" italics="true" pageId="28" pageNumber="29">lvp</emphasis>
) elongate, rather wide in basal portion and directed mesocephalad (weakly sinuate), then tapering, curved laterocephalad and pointed apically; ventral lobe of periandrium (
@ -327,7 +328,7 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
) and dorsal lobe (
<emphasis id="71E5EAFF71466239FB3CFE62FDCFFE66" box="[545,568,433,459]" italics="true" pageId="28" pageNumber="29">dl</emphasis>
); aedeagus bilobed apically, each lobe with apical subtriangular process with dorsal margin undulate in caudal view (
<figureCitation id="DBAA2A6871466239FBCBFE01FCB2FE40" box="[726,837,466,493]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="28" pageNumber="29">Fig. 14G</figureCitation>
<figureCitation id="DBAA2A6871466239FBCBFE01FCB2FE40" box="[726,837,466,493]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="28" pageNumber="29">Fig. 14G</figureCitation>
), pointing laterocephalad. Connective (
<emphasis id="71E5EAFF71466239FC00FE00FACEFE41" box="[1309,1337,467,492]" italics="true" pageId="28" pageNumber="29">co</emphasis>
) and tectiductus (
@ -335,11 +336,11 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
) weak (maybe not fully mature specimen). Anal tube (
<emphasis id="71E5EAFF71466239FAF3FE26FBF8FDA2" box="[1006,1039,501,527]" italics="true" pageId="28" pageNumber="29">An</emphasis>
–
<figureCitation id="DBAA2A6871466239FD33FE27FB4AFDA3" box="[1070,1213,500,526]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="28" pageNumber="29">Fig. 14A–D</figureCitation>
<figureCitation id="DBAA2A6871466239FD33FE27FB4AFDA3" box="[1070,1213,500,526]" captionStart="Fig" captionStartId="27.[189,232,1892,1918]" captionTargetBox="[254,1333,265,1852]" captionTargetId="figure-12@27.[254,1333,265,1852]" captionTargetPageId="27" captionText="Fig. 14. Flavina lami sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="28" pageNumber="29">Fig. 14A–D</figureCitation>
) dorsoventrally flattened, in dorsal view evenly widening to midlength, with apical margin strongly bisinuate with middle portion roundly protruding, with lateral lobes strongly developed, angularly pointing posteroventrad; anal opening before basal ½.
</paragraph>
</subSubSection>
<caption id="17EE66657140623FF9A2F85BFECCF845" pageId="26" pageNumber="27" startId="26.[191,234,1928,1954]" targetBox="[189,1398,444,1885]" targetPageId="26">
<caption id="17EE66657140623FF9A2F85BFECCF845" ID-DOI="http://doi.org/10.5281/zenodo.14508764" ID-Zenodo-Dep="14508764" httpUri="https://zenodo.org/record/14508764/files/figure.png" pageId="26" pageNumber="27" startId="26.[191,234,1928,1954]" targetBox="[189,1398,444,1885]" targetPageId="26">
<paragraph id="432E36ED7140623FF9A2F85BFECCF845" blockId="26.[189,1398,1928,2024]" pageId="26" pageNumber="27">
<emphasis id="71E5EAFF7140623FF9A2F85BFED6F80F" bold="true" box="[191,289,1928,1954]" pageId="26" pageNumber="27">Fig. 13.</emphasis>
<taxonomicName id="84914D6E7140623FF836F85BFE30F80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[299,455,1928,1954]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="lami" status="sp. nov.">
@ -359,7 +360,7 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
. Right hind wing.
</paragraph>
</caption>
<caption id="17EE66657141623EF9A0F8B7FC08F845" pageId="27" pageNumber="28" startId="27.[189,232,1892,1918]" targetBox="[254,1333,265,1852]" targetPageId="27">
<caption id="17EE66657141623EF9A0F8B7FC08F845" ID-DOI="http://doi.org/10.5281/zenodo.14508766" ID-Zenodo-Dep="14508766" httpUri="https://zenodo.org/record/14508766/files/figure.png" pageId="27" pageNumber="28" startId="27.[189,232,1892,1918]" targetBox="[254,1333,265,1852]" targetPageId="27">
<paragraph id="432E36ED7141623EF9A0F8B7FC08F845" blockId="27.[188,1399,1892,2025]" pageId="27" pageNumber="28">
<emphasis id="71E5EAFF7141623EF9A0F8B7FEEBF8D2" bold="true" box="[189,284,1892,1919]" pageId="27" pageNumber="28">Fig. 14.</emphasis>
<taxonomicName id="84914D6E7141623EF838F8B6FE4AF8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[293,445,1893,1919]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="lami" status="sp. nov.">
@ -409,7 +410,7 @@ distal half; 8 apical spines, all spines brown, apically black. First metatarsom
was collected in July, in montane evergreen tropical forest, at about
<quantity id="84699B0871466239FC30FD15FEE1FCAF" metricMagnitude="3" metricUnit="m" metricValue="1.35" metricValueMax="1.4" metricValueMin="1.3" pageId="28" pageNumber="29" unit="m" value="1350.0" valueMax="1400.0" valueMin="1300.0">1300– 1400 m</quantity>
a.s.l. near the summit (
<figureCitation id="DBAA2A6871466239FB37FD34FD7FFCAC" box="[554,648,743,769]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="28" pageNumber="29">Fig. 1D</figureCitation>
<figureCitation id="DBAA2A6871466239FB37FD34FD7FFCAC" box="[554,648,743,769]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="28" pageNumber="29">Fig. 1D</figureCitation>
). The specimen was collected on tree trunks densely covered in lichens and mosses. Later fieldwork in
<date id="372F102D71466239FB96FCDAFCFBFC8E" box="[651,780,777,803]" pageId="28" pageNumber="29" value="2017-04">April 2017</date>
and
@ -426,7 +427,7 @@ at the same locality failed to provide additional specimens.
<paragraph id="432E36ED71466239F9A0FC44FB9DFC1C" blockId="28.[189,1130,919,948]" box="[189,1130,919,948]" pageId="28" pageNumber="29">
<collectingCountry id="3B86767D71466239F9A0FC44FED2FC1C" box="[189,293,919,945]" name="Vietnam" pageId="28" pageNumber="29">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, summit of Bach Ma National Park (
<figureCitation id="DBAA2A6871466239FD09FC44FBAEFC1C" box="[1044,1113,919,945]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="28" pageNumber="29">Fig. 1</figureCitation>
<figureCitation id="DBAA2A6871466239FD09FC44FBAEFC1C" box="[1044,1113,919,945]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="28" pageNumber="29">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>

117
data/CB/38/87/CB3887FB7152622BFB21FB55FB9DFA21.xml
View file

@ -1,54 +1,55 @@
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<mods:title id="9269121DD91D0CAE5BAAFA9E83EC7522">Issid planthoppers from Bach Ma and Phong Dien in Central Vietnam. I. Tribe Parahiraciini (Hemiptera: Fulgoromorpha: Issidae)</mods:title>
<document id="1338E932762AA10807B81FAA15209B3E" ID-DOI="10.5852/ejt.2024.975.2769" ID-ISSN="2118-9773" ID-Zenodo-Dep="14508730" ID-ZooBank="23F0D60C-3B5B-4AB1-B608-AD875EB424DF" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1734447201131" checkinUser="plazi" docAuthor="Constant, Jérôme &amp; Pham, Hong Thai" docDate="2024" docId="CB3887FB7152622BFB21FB55FB9DFA21" docLanguage="en" docName="EJT.2024.975.1-64.pdf" docOrigin="European Journal of Taxonomy 975" docSource="https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2769/12667" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Cyclopissus corticalis Constant &amp; Pham, 2024, gen. et sp. nov." docType="treatment" docVersion="2" lastPageNumber="15" masterDocId="3701FF83715A6225F91DFFD3FFF7FFAD" masterDocTitle="Issid planthoppers from Bach Ma and Phong Dien in Central Vietnam. I. Tribe Parahiraciini (Hemiptera: Fulgoromorpha: Issidae)" masterLastPageNumber="64" masterPageNumber="1" pageNumber="9" updateTime="1734447759699" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="CDD7738A3B039EABFAB3F67D799646F7">Issid planthoppers from Bach Ma and Phong Dien in Central Vietnam. I. Tribe Parahiraciini (Hemiptera: Fulgoromorpha: Issidae)</mods:title>
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<figureCitation id="DBAA2A687152622DFBA2FB19FCFFFB49" box="[703,776,1226,1252]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="8" pageNumber="9">Figs 1</figureCitation>
<figureCitation id="DBAA2A687152622DFBA2FB19FCFFFB49" box="[703,776,1226,1252]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="8" pageNumber="9">Figs 1</figureCitation>
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<figureCitation id="DBAA2A687152622DFA0BFB19FCCBFB49" box="[790,828,1226,1252]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="8" pageNumber="9">2D</figureCitation>
<figureCitation id="DBAA2A687152622DFA0BFB19FCCBFB49" box="[790,828,1226,1252]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="8" pageNumber="9">2D</figureCitation>
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<figureCitation id="DBAA2A687152622DFA54FB19FC83FB49" box="[841,884,1226,1252]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="10.[189,232,1928,1954]" captionStartId-1="11.[189,232,1850,1876]" captionStartId-2="12.[189,232,1892,1918]" captionStartId-3="13.[190,233,1963,1989]" captionTargetBox-0="[244,1333,271,1880]" captionTargetBox-1="[189,1398,277,1784]" captionTargetBox-2="[255,1333,265,1851]" captionTargetBox-3="[189,1398,505,1919]" captionTargetId-0="figure-16@10.[244,1344,265,1883]" captionTargetId-1="figure-12@11.[189,1398,265,1806]" captionTargetId-2="figure-16@12.[255,1333,265,1851]" captionTargetId-3="figure-75@13.[189,1398,505,1919]" captionTargetPageId-0="10" captionTargetPageId-1="11" captionTargetPageId-2="12" captionTargetPageId-3="13" captionText-0="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." captionText-3="Fig. 6. Cyclopissus corticalis gen. et sp. nov., live specimens in Bach Ma National Park, summit. A. 13 May 2023. B–C. 14 May 2023." pageId="8" pageNumber="9">3–6</figureCitation>
<figureCitation id="DBAA2A687152622DFA54FB19FC83FB49" box="[841,884,1226,1252]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="10.[189,232,1928,1954]" captionStartId-1="11.[189,232,1850,1876]" captionStartId-2="12.[189,232,1892,1918]" captionStartId-3="13.[190,233,1963,1989]" captionTargetBox-0="[244,1333,271,1880]" captionTargetBox-1="[189,1398,277,1784]" captionTargetBox-2="[255,1333,265,1851]" captionTargetBox-3="[189,1398,505,1919]" captionTargetId-0="figure-16@10.[244,1344,265,1883]" captionTargetId-1="figure-12@11.[189,1398,265,1806]" captionTargetId-2="figure-16@12.[255,1333,265,1851]" captionTargetId-3="figure-75@13.[189,1398,505,1919]" captionTargetPageId-0="10" captionTargetPageId-1="11" captionTargetPageId-2="12" captionTargetPageId-3="13" captionText-0="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." captionText-3="Fig. 6. Cyclopissus corticalis gen. et sp. nov., live specimens in Bach Ma National Park, summit. A. 13 May 2023. B–C. 14 May 2023." figureDoi-0="http://doi.org/10.5281/zenodo.14508738" figureDoi-1="http://doi.org/10.5281/zenodo.14508740" figureDoi-2="http://doi.org/10.5281/zenodo.14508742" figureDoi-3="http://doi.org/10.5281/zenodo.14508744" httpUri-0="https://zenodo.org/record/14508738/files/figure.png" httpUri-1="https://zenodo.org/record/14508740/files/figure.png" httpUri-2="https://zenodo.org/record/14508742/files/figure.png" httpUri-3="https://zenodo.org/record/14508744/files/figure.png" pageId="8" pageNumber="9">3–6</figureCitation>
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@ -141,7 +142,7 @@ The species epithet
•
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(dissected –
<figureCitation id="DBAA2A687152622DFB06F859FD70F808" box="[539,647,1930,1957]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="10.[189,232,1928,1954]" captionStartId-1="11.[189,232,1850,1876]" captionStartId-2="12.[189,232,1892,1918]" captionTargetBox-0="[244,1333,271,1880]" captionTargetBox-1="[189,1398,277,1784]" captionTargetBox-2="[255,1333,265,1851]" captionTargetId-0="figure-16@10.[244,1344,265,1883]" captionTargetId-1="figure-12@11.[189,1398,265,1806]" captionTargetId-2="figure-16@12.[255,1333,265,1851]" captionTargetPageId-0="10" captionTargetPageId-1="11" captionTargetPageId-2="12" captionText-0="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." pageId="8" pageNumber="9">Figs 3–5</figureCitation>
<figureCitation id="DBAA2A687152622DFB06F859FD70F808" box="[539,647,1930,1957]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="10.[189,232,1928,1954]" captionStartId-1="11.[189,232,1850,1876]" captionStartId-2="12.[189,232,1892,1918]" captionTargetBox-0="[244,1333,271,1880]" captionTargetBox-1="[189,1398,277,1784]" captionTargetBox-2="[255,1333,265,1851]" captionTargetId-0="figure-16@10.[244,1344,265,1883]" captionTargetId-1="figure-12@11.[189,1398,265,1806]" captionTargetId-2="figure-16@12.[255,1333,265,1851]" captionTargetPageId-0="10" captionTargetPageId-1="11" captionTargetPageId-2="12" captionText-0="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508738" figureDoi-1="http://doi.org/10.5281/zenodo.14508740" figureDoi-2="http://doi.org/10.5281/zenodo.14508742" httpUri-0="https://zenodo.org/record/14508738/files/figure.png" httpUri-1="https://zenodo.org/record/14508740/files/figure.png" httpUri-2="https://zenodo.org/record/14508742/files/figure.png" pageId="8" pageNumber="9">Figs 3–5</figureCitation>
);
<location id="464E60367152622DFB82F859FD48F809" LSID="urn:lsid:plazi:treatment:CB3887FB7152622BFB21FB55FB9DFA21:464E60367152622DFB82F859FD48F809" box="[671,703,1930,1956]" country="Vietnam" county="Bach Ma National Park" latitude="16.188332" longLatPrecision="21" longitude="107.84889" municipality="Province" name="Th" pageId="8" pageNumber="9">Th</location>
ừa Thiên-Huế
@ -252,29 +253,29 @@ leg.;
<paragraph id="432E36ED7153622CF9A0FD94FEFEFBAA" blockId="9.[187,1400,583,1031]" pageId="9" pageNumber="10">
<smallCapsWord id="45C8A0317153622CF9A0FD94FF0BFDF2" baselines="602,602" box="[189,252,583,609]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="9" pageNumber="10">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A687153622CF80CFD94FE60FDCC" box="[273,407,583,609]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="9" pageNumber="10">Fig. 4A–D</figureCitation>
<figureCitation id="DBAA2A687153622CF80CFD94FE60FDCC" box="[273,407,583,609]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="9" pageNumber="10">Fig. 4A–D</figureCitation>
). Head variegated brown, slightly narrower than thorax. Vertex slightly more than two times as wide as long in midline, medially grooved with margins weakly carinate; sides subparallel; anterior and posterior margins subparallel and curved. Posterior face of head largely blackish brown. Frons as wide at maximum width slightly above clypeus, as long in midline, with large oculiform black, shiny marking in dorsal portion with pale yellowish spot in dorsomedian portion, black marking finely circled in pale brown, turning to slightly wider pale yellowish in ventral portion; median carina in ventral portion, disappearing at midheight of black marking; pale yellowish tubercles along lateral margins under midheight of eye; laterodorsal angles black. Clypeus slightly paler basally, flat in middle portion, moderately elongate, subtriangular with fronto-clypeal suture rounded; anteclypeus with median blunt carina. Labium yellow-brown, elongate and narrow, reaching metacoxae, with apical segment elongate, nearly as long as penultimate. Eyes reniform (not emarginate), strongly projecting laterally; ocelli absent. Antennae dark brown, rather short with scape ring-shaped and pedicel cylindrical, slightly longer than broad.
</paragraph>
<paragraph id="432E36ED7153622CF9A0FBFCFC72FA28" blockId="9.[188,1399,1070,1414]" pageId="9" pageNumber="10">
<smallCapsWord id="45C8A0317153622CF9A0FBFCFEEEFBEA" baselines="1090,1090" box="[189,281,1071,1097]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="9" pageNumber="10">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A687153622CF834FBFDFE5CFBE4" box="[297,427,1070,1097]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="9" pageNumber="10">Fig. 4A–D</figureCitation>
<figureCitation id="DBAA2A687153622CF834FBFDFE5CFBE4" box="[297,427,1070,1097]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="9" pageNumber="10">Fig. 4A–D</figureCitation>
). Pronotum variegated brown, with more or less well marked median yellowish line on obsolete carina; impressed point on each side of median carina; anterior margin bordered posteriorly with several yellowish blunt tubercles and carinate in middle portion, strongly sinuate and strongly, roundly protruding anteriorly between eyes; posterior margin nearly straight, slightly carinate in middle portion; pale yellowish blunt tubercles on disc and sides; behind eyes, large blackish marking containing pale yellowish mark, with black area continued on inner half of paranotal lobes; lateral half of paranotal lobes yellowish brown with yellowish tubercles along posterior margin, with posteroventral angle of lobes rounded. Mesonotum yellowish with yellowish, obsolete median carina in middle of rather wide dark brown median line; weak peridiscal carinae bordered with dark brown fine line; lateral angles darker; apex of scutellum pale yellowish. Tegulae yellowish.
</paragraph>
<paragraph id="432E36ED7153622CF9A0FA7EFBD7F910" blockId="9.[188,1400,1453,1726]" pageId="9" pageNumber="10">
<smallCapsWord id="45C8A0317153622CF9A0FA7EFED5FA68" baselines="1472,1472" box="[189,290,1453,1479]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="9" pageNumber="10">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A687153622CF825FA7EFE4AFA6A" box="[312,445,1453,1479]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="9" pageNumber="10">Fig. 3A–C</figureCitation>
<figureCitation id="DBAA2A687153622CF825FA7EFE4AFA6A" box="[312,445,1453,1479]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508738" httpUri="https://zenodo.org/record/14508738/files/figure.png" pageId="9" pageNumber="10">Fig. 3A–C</figureCitation>
). Tegmina variegated brown with main veins raised and yellowish, and dense network of pale, nearly whitish, cross-veinlets; subcoriaceous, elongate, about 2.3 times as long as broad, convex with sides broadly rounded (middle portion nearly straight to weakly incurved in dorsal view); epipleuron not developed. Apex narrowly rounded. Postclaval margin weakly rounded on distal half and slightly notched at apex of clavus. Clavus closed, reaching about 3/5 of tegmen. Venation: ScP+R rather short, forking into subparallel ScP+RA and RP; RP forking at about midlength of tegmen; MP first fork at about half of tegmen length; CuA straight, unforked and reaching hind margin; Pcu fused with A1 at 3/5 of clavus length; Pcu+A1 fused with CuP at apex of clavus.
</paragraph>
<paragraph id="432E36ED71536228F9A0F936FB11FEEB" blockId="9.[189,1400,1765,1967]" lastBlockId="13.[188,1399,265,327]" lastPageId="13" lastPageNumber="14" pageId="9" pageNumber="10">
<smallCapsWord id="45C8A0317153622CF9A0F936FF00F950" baselines="1784,1784" box="[189,247,1765,1791]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="9" pageNumber="10">HIND</smallCapsWord>
<smallCapsWord id="45C8A0317153622CF81DF939FEB3F950" baselines="1784" box="[256,324,1770,1789]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="9" pageNumber="10">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A687153622CF84BF936FE44F952" box="[342,435,1765,1791]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="9" pageNumber="10">Fig. 4E</figureCitation>
<figureCitation id="DBAA2A687153622CF84BF936FE44F952" box="[342,435,1765,1791]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="9" pageNumber="10">Fig. 4E</figureCitation>
). Dark brown with contrasting yellow-brown markings, one basal along vein Pcu and one central, larger, including portion of Pcu. Hing wings wider than tegmina and deeply bilobed, strongly notched at CuP; costal margin sinuate; CuP-Pcu-A1 lobe about 1.7 times as wide as Sc-R-MP-CuA lobe, and slightly longer than latter; both lobes rounded apically; postclaval margin broadly rounded; A2 lobe reduced and narrow. Venation: main veins distinct, numerous cross-veins; ScP+R, MP and CuA running more or less parallel in basal portion, with ScP+R diverging beyond basal ⅓; Pcu strongly curved around basal third of wing towards CuP, and reaching the latter; A1 curved, more or less parallel to postclaval margin; CuP-Pcu-A1 lobe with numerous cross-veins. Vein A2 weak.
</paragraph>
<caption id="17EE66657150622FF9A0F85BFDC6F845" pageId="10" pageNumber="11" startId="10.[189,232,1928,1954]" targetBox="[244,1333,271,1880]" targetPageId="10">
<caption id="17EE66657150622FF9A0F85BFDC6F845" ID-DOI="http://doi.org/10.5281/zenodo.14508738" ID-Zenodo-Dep="14508738" httpUri="https://zenodo.org/record/14508738/files/figure.png" pageId="10" pageNumber="11" startId="10.[189,232,1928,1954]" targetBox="[244,1333,271,1880]" targetPageId="10">
<paragraph id="432E36ED7150622FF9A0F85BFDC6F845" blockId="10.[189,1399,1928,2025]" pageId="10" pageNumber="11">
<emphasis id="71E5EAFF7150622FF9A0F85BFEFCF80F" bold="true" box="[189,267,1928,1954]" pageId="10" pageNumber="11">Fig. 3.</emphasis>
<taxonomicName id="84914D6E7150622FF80EF85BFDE2F80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[275,533,1928,1954]" class="Insecta" family="Issidae" genus="Cyclopissus" kingdom="Animalia" order="Hemiptera" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="corticalis" status="gen. et sp. nov.">
@ -294,7 +295,7 @@ leg.;
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
<caption id="17EE66657151622EF9A0F8E9FEADF813" pageId="11" pageNumber="12" startId="11.[189,232,1850,1876]" targetBox="[189,1398,277,1784]" targetPageId="11">
<caption id="17EE66657151622EF9A0F8E9FEADF813" ID-DOI="http://doi.org/10.5281/zenodo.14508740" ID-Zenodo-Dep="14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="11" pageNumber="12" startId="11.[189,232,1850,1876]" targetBox="[189,1398,277,1784]" targetPageId="11">
<paragraph id="432E36ED7151622EF9A0F8E9FEADF813" blockId="11.[189,1399,1850,1982]" pageId="11" pageNumber="12">
<emphasis id="71E5EAFF7151622EF9A0F8E9FEFBF8F9" bold="true" box="[189,268,1850,1876]" pageId="11" pageNumber="12">Fig. 4.</emphasis>
<taxonomicName id="84914D6E7151622EF80BF8E9FDEDF8F9" authorityName="Constant &amp; Pham" authorityYear="2024" box="[278,538,1850,1876]" class="Insecta" family="Issidae" genus="Cyclopissus" kingdom="Animalia" order="Hemiptera" pageId="11" pageNumber="12" phylum="Arthropoda" rank="species" species="corticalis" status="gen. et sp. nov.">
@ -317,7 +318,7 @@ leg.;
. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods.
</paragraph>
</caption>
<caption id="17EE666571566229F9A0F8B7FC5EF845" pageId="12" pageNumber="13" startId="12.[189,232,1892,1918]" targetBox="[255,1333,265,1851]" targetPageId="12">
<caption id="17EE666571566229F9A0F8B7FC5EF845" ID-DOI="http://doi.org/10.5281/zenodo.14508742" ID-Zenodo-Dep="14508742" httpUri="https://zenodo.org/record/14508742/files/figure.png" pageId="12" pageNumber="13" startId="12.[189,232,1892,1918]" targetBox="[255,1333,265,1851]" targetPageId="12">
<paragraph id="432E36ED71566229F9A0F8B7FC5EF845" blockId="12.[187,1399,1892,2025]" pageId="12" pageNumber="13">
<emphasis id="71E5EAFF71566229F9A0F8B7FEF8F8D2" bold="true" box="[189,271,1892,1919]" pageId="12" pageNumber="13">Fig. 5.</emphasis>
<taxonomicName id="84914D6E71566229F806F8B6FDD5F8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[283,546,1893,1919]" class="Insecta" family="Issidae" genus="Cyclopissus" kingdom="Animalia" order="Hemiptera" pageId="12" pageNumber="13" phylum="Arthropoda" rank="species" species="corticalis" status="gen. et sp. nov.">
@ -354,10 +355,10 @@ leg.;
<paragraph id="432E36ED7157622BF9A0FEBDFAF1FE5B" blockId="13.[189,1400,366,427]" lastBlockId="14.[188,1401,265,503]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
<smallCapsWord id="45C8A03171576228F9A0FEBDFF01FE2B" baselines="385,385" box="[189,246,366,392]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="13" pageNumber="14">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A6871576228F814FEBDFEA5FE25" box="[265,338,366,392]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="13" pageNumber="14">Fig. 3</figureCitation>
<figureCitation id="DBAA2A6871576228F814FEBDFEA5FE25" box="[265,338,366,392]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508738" httpUri="https://zenodo.org/record/14508738/files/figure.png" pageId="13" pageNumber="14">Fig. 3</figureCitation>
). Pro- and mesocoxae pale yellowish with dark marking along anterior margin. Pro- and mesofemora elongate, relatively slender, shorter than corresponding tibiae, brown with apex paler and median pale yellowish ring, brown colour darker on ventral face. Pro- and mesotibiae more slender than corresponding femora, pale yellowish with subbasal and subapical brown rings leaving more or less wide pale ring in middle. Metafemora dark brown with posterior side pale yellowish in basal portion. Metatibiae yellowish with 2 lateral spines in distal half and 7 apical spines, apex of all spines black. Tarsi elongate; first metatarsomere yellowish, elongate and slender, with a strong spine at each side and a row of 5 smaller spines in between ventrally along posterior margin, apex of all spines black; second metatarsomere brown, short with one spine at each side. Metatibiotarsal formula: (2) 7 / 8 / 2.
</paragraph>
<caption id="17EE666571576228F9A3F878FD81F845" pageId="13" pageNumber="14" startId="13.[190,233,1963,1989]" targetBox="[189,1398,505,1919]" targetPageId="13">
<caption id="17EE666571576228F9A3F878FD81F845" ID-DOI="http://doi.org/10.5281/zenodo.14508744" ID-Zenodo-Dep="14508744" httpUri="https://zenodo.org/record/14508744/files/figure.png" pageId="13" pageNumber="14" startId="13.[190,233,1963,1989]" targetBox="[189,1398,505,1919]" targetPageId="13">
<paragraph id="432E36ED71576228F9A3F878FD81F845" blockId="13.[189,1399,1963,2025]" pageId="13" pageNumber="14">
<emphasis id="71E5EAFF71576228F9A3F878FEE6F868" bold="true" box="[190,273,1963,1989]" pageId="13" pageNumber="14">Fig. 6.</emphasis>
<taxonomicName id="84914D6E71576228F801F878FDD4F868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[284,547,1963,1989]" class="Insecta" family="Issidae" genus="Cyclopissus" kingdom="Animalia" order="Hemiptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="species" species="corticalis" status="gen. et sp. nov.">
@ -377,17 +378,17 @@ leg.;
. Pygofer (
<emphasis id="71E5EAFF7154622BF8E2FDCDFDE9FD9A" box="[511,542,542,567]" italics="true" pageId="14" pageNumber="15">Py</emphasis>
–
<figureCitation id="DBAA2A687154622BFB2AFDCDFD43FD95" box="[567,692,541,568]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." pageId="14" pageNumber="15">Fig. 5A–D</figureCitation>
<figureCitation id="DBAA2A687154622BFB2AFDCDFD43FD95" box="[567,692,541,568]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508742" httpUri="https://zenodo.org/record/14508742/files/figure.png" pageId="14" pageNumber="15">Fig. 5A–D</figureCitation>
) higher than long in lateral view, with anterior margin weakly concave and posterior margin broadly rounded in upper half; subcircular in caudal view; posterior margin deeply emarginate in dorsal view. Gonostyli (
<emphasis id="71E5EAFF7154622BFBD2FDB7FD13FDD3" box="[719,740,612,638]" italics="true" pageId="14" pageNumber="15">G</emphasis>
–
<figureCitation id="DBAA2A687154622BFA1CFDB7FC89FDD2" box="[769,894,612,639]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." pageId="14" pageNumber="15">Fig. 5A–C</figureCitation>
<figureCitation id="DBAA2A687154622BFA1CFDB7FC89FDD2" box="[769,894,612,639]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508742" httpUri="https://zenodo.org/record/14508742/files/figure.png" pageId="14" pageNumber="15">Fig. 5A–C</figureCitation>
) rather elongate, projecting posteriorly and with rounded posterior margin in lateral view, with capitulum (
<emphasis id="71E5EAFF7154622BFADFFD5BFC29FD0C" box="[962,990,648,673]" italics="true" pageId="14" pageNumber="15">ca</emphasis>
) strongly developed dorsad, with lateral laminate upcurving projection and with rather long neck with outer margin strongly concave and inner margin sinuate, in caudal view, and apical portion anteroposteriorly flattened, with apical hook and inner margin angularly rounded. Anal tube (
<emphasis id="71E5EAFF7154622BFBDAFD22FD1FFCA6" box="[711,744,753,779]" italics="true" pageId="14" pageNumber="15">An</emphasis>
–
<figureCitation id="DBAA2A687154622BFA1AFD22FC7FFCA1" box="[775,904,753,780]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." pageId="14" pageNumber="15">Fig. 5A–D</figureCitation>
<figureCitation id="DBAA2A687154622BFA1AFD22FC7FFCA1" box="[775,904,753,780]" captionStart="Fig" captionStartId="12.[189,232,1892,1918]" captionTargetBox="[255,1333,265,1851]" captionTargetId="figure-16@12.[255,1333,265,1851]" captionTargetPageId="12" captionText="Fig. 5. Cyclopissus corticalis gen. et sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–J. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508742" httpUri="https://zenodo.org/record/14508742/files/figure.png" pageId="14" pageNumber="15">Fig. 5A–D</figureCitation>
) dorsoventrally flattened, rather wide and subhexagonal in dorsal view and with posterolateral angles expanded lateroventrally, leaving middle portion of posterior margin strongly excavate in caudal view. Aedeagus curved dorsad (in lateral view), with symmetrical pair of elongate, apically pointed lateroventral processes (
<emphasis id="71E5EAFF7154622BFD2EFC89FBA0FCD9" box="[1075,1111,858,884]" italics="true" pageId="14" pageNumber="15">lvp</emphasis>
) directed caudad; dorsal lobe (
@ -406,7 +407,7 @@ leg.;
. Hind margin of sternum VII (
<emphasis id="71E5EAFF7154622BFA1CFC30FCB4FC51" box="[769,835,995,1021]" italics="true" pageId="14" pageNumber="15">St VII</emphasis>
–
<figureCitation id="DBAA2A687154622BFA42FC31FC43FC51" box="[863,948,994,1020]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="14" pageNumber="15">Fig. 4F</figureCitation>
<figureCitation id="DBAA2A687154622BFA42FC31FC43FC51" box="[863,948,994,1020]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="14" pageNumber="15">Fig. 4F</figureCitation>
) concave, moderately inflated towards lateral portion.
</paragraph>
</subSubSection>
@ -424,9 +425,9 @@ leg.;
was collected in May and September, in montane evergreen tropical forest, exclusively at about
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a.s.l. near the summit (
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<figureCitation id="DBAA2A687154622BFD0BFB4AFB83FB1E" box="[1046,1140,1177,1203]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="14" pageNumber="15">Fig. 1D</figureCitation>
). The species lives on tree trunks densely covered in lichens and mosses, on which it is very cryptic (
<figureCitation id="DBAA2A687154622BFD4AFB6FFB68FB7B" box="[1111,1183,1212,1238]" captionStart="Fig" captionStartId="13.[190,233,1963,1989]" captionTargetBox="[189,1398,505,1919]" captionTargetId="figure-75@13.[189,1398,505,1919]" captionTargetPageId="13" captionText="Fig. 6. Cyclopissus corticalis gen. et sp. nov., live specimens in Bach Ma National Park, summit. A. 13 May 2023. B–C. 14 May 2023." pageId="14" pageNumber="15">Fig. 6</figureCitation>
<figureCitation id="DBAA2A687154622BFD4AFB6FFB68FB7B" box="[1111,1183,1212,1238]" captionStart="Fig" captionStartId="13.[190,233,1963,1989]" captionTargetBox="[189,1398,505,1919]" captionTargetId="figure-75@13.[189,1398,505,1919]" captionTargetPageId="13" captionText="Fig. 6. Cyclopissus corticalis gen. et sp. nov., live specimens in Bach Ma National Park, summit. A. 13 May 2023. B–C. 14 May 2023." figureDoi="http://doi.org/10.5281/zenodo.14508744" httpUri="https://zenodo.org/record/14508744/files/figure.png" pageId="14" pageNumber="15">Fig. 6</figureCitation>
). When disturbed, they tend to walk away and hide on the opposite side of the trunk or branch, and jump away as a last solution.
</paragraph>
</subSubSection>
@ -439,7 +440,7 @@ a.s.l. near the summit (
<paragraph id="432E36ED7154622BF9A0FAA1FB9DFA21" blockId="14.[189,1130,1394,1423]" box="[189,1130,1394,1423]" pageId="14" pageNumber="15">
<collectingCountry id="3B86767D7154622BF9A0FAA1FED2FA21" box="[189,293,1394,1420]" name="Vietnam" pageId="14" pageNumber="15">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, summit of Bach Ma National Park (
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<figureCitation id="DBAA2A687154622BFD09FAA1FBAEFA21" box="[1044,1113,1394,1420]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="14" pageNumber="15">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>

145
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<treatment id="CB3887FB71556233FB6BFA4FFE79F966" LSID="urn:lsid:plazi:treatment:CB3887FB71556233FB6BFA4FFE79F966" httpUri="http://treatment.plazi.org/id/CB3887FB71556233FB6BFA4FFE79F966" lastPageId="22" lastPageNumber="23" pageId="15" pageNumber="16">
<subSubSection id="0B8B65667155622AFB6BFA4FFB31FA74" pageId="15" pageNumber="16" type="nomenclature">
@ -67,9 +68,9 @@ urn:lsid:zoobank.org:act:
</subSubSection>
<subSubSection id="0B8B65667155622AFBCCFA32FC95FA56" box="[721,866,1505,1531]" pageId="15" pageNumber="16" type="description">
<paragraph id="432E36ED7155622AFBCCFA32FC95FA56" blockId="15.[365,1222,1436,1531]" box="[721,866,1505,1531]" pageId="15" pageNumber="16">
<figureCitation id="DBAA2A687155622AFBCCFA32FCEBFA56" box="[721,796,1505,1531]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="15" pageNumber="16">Figs 2</figureCitation>
<figureCitation id="DBAA2A687155622AFBCCFA32FCEBFA56" box="[721,796,1505,1531]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="15" pageNumber="16">Figs 2</figureCitation>
,
<figureCitation id="DBAA2A687155622AFA34FA32FC95FA56" box="[809,866,1505,1531]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStartId-0="17.[189,232,1928,1954]" captionStartId-1="18.[189,232,1892,1918]" captionStartId-2="20.[189,232,1892,1918]" captionStartId-3="21.[189,232,1963,1989]" captionStartId-4="22.[189,232,1529,1555]" captionTargetBox-0="[245,1337,271,1876]" captionTargetBox-1="[242,1345,274,1824]" captionTargetBox-2="[257,1332,283,1846]" captionTargetBox-3="[189,1398,1438,1920]" captionTargetBox-4="[189,1398,265,1486]" captionTargetId-0="figure-12@17.[242,1345,265,1886]" captionTargetId-1="figure-17@18.[242,1345,265,1827]" captionTargetId-2="figure-16@20.[256,1332,265,1847]" captionTargetId-3="figure-455@21.[189,1398,1438,1920]" captionTargetId-4="figure-74@22.[189,1398,265,1486]" captionTargetPageId-0="17" captionTargetPageId-1="18" captionTargetPageId-2="20" captionTargetPageId-3="21" captionTargetPageId-4="22" captionText-0="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." captionText-4="Fig. 11. Flavina bachmana sp. nov., distribution map." pageId="15" pageNumber="16">7–11</figureCitation>
<figureCitation id="DBAA2A687155622AFA34FA32FC95FA56" box="[809,866,1505,1531]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStartId-0="17.[189,232,1928,1954]" captionStartId-1="18.[189,232,1892,1918]" captionStartId-2="20.[189,232,1892,1918]" captionStartId-3="21.[189,232,1963,1989]" captionStartId-4="22.[189,232,1529,1555]" captionTargetBox-0="[245,1337,271,1876]" captionTargetBox-1="[242,1345,274,1824]" captionTargetBox-2="[257,1332,283,1846]" captionTargetBox-3="[189,1398,1438,1920]" captionTargetBox-4="[189,1398,265,1486]" captionTargetId-0="figure-12@17.[242,1345,265,1886]" captionTargetId-1="figure-17@18.[242,1345,265,1827]" captionTargetId-2="figure-16@20.[256,1332,265,1847]" captionTargetId-3="figure-455@21.[189,1398,1438,1920]" captionTargetId-4="figure-74@22.[189,1398,265,1486]" captionTargetPageId-0="17" captionTargetPageId-1="18" captionTargetPageId-2="20" captionTargetPageId-3="21" captionTargetPageId-4="22" captionText-0="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." captionText-4="Fig. 11. Flavina bachmana sp. nov., distribution map." figureDoi-0="http://doi.org/10.5281/zenodo.14508748" figureDoi-1="http://doi.org/10.5281/zenodo.14508752" figureDoi-2="http://doi.org/10.5281/zenodo.14508756" figureDoi-3="http://doi.org/10.5281/zenodo.14508758" figureDoi-4="http://doi.org/10.5281/zenodo.14508760" httpUri-0="https://zenodo.org/record/14508748/files/figure.png" httpUri-1="https://zenodo.org/record/14508752/files/figure.png" httpUri-2="https://zenodo.org/record/14508756/files/figure.png" httpUri-3="https://zenodo.org/record/14508758/files/figure.png" httpUri-4="https://zenodo.org/record/14508760/files/figure.png" pageId="15" pageNumber="16">7–11</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="0B8B656671556235F9A0F9F2FC6AFD23" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16" type="diagnosis">
@ -111,14 +112,14 @@ and
which differ by much longer, curved lateroventral processes of the aedeagus (
<bibRefCitation id="27004B1C714A6235F8E3FE89FD13FED9" author="Ran H. - F. &amp; Liang A. - P." box="[510,740,346,372]" pageId="16" pageNumber="17" pagination="388 - 391" refId="ref24916" refString="Ran H. - F. &amp; Liang A. - P. 2006. Taxonomic study of the issid genus Flavina Stal (Hemiptera, Fulgoroidea, Issidae). Acta Zootaxonomica Sinica 31 (2): 388 - 391." type="journal article" year="2006">Ran &amp; Liang 2006</bibRefCitation>
: figs 7–8;
<figureCitation id="DBAA2A68714A6235FA7DFE89FC1AFED9" box="[864,1005,346,372]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 17E, K</figureCitation>
<figureCitation id="DBAA2A68714A6235FA7DFE89FC1AFED9" box="[864,1005,346,372]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508772" httpUri="https://zenodo.org/record/14508772/files/figure.png" pageId="16" pageNumber="17">Fig. 17E, K</figureCitation>
) as opposed to short and straight in
<taxonomicName id="84914D6E714A6235F9C1FEADFE84FE3A" authorityName="Constant &amp; Pham" authorityYear="2024" box="[220,371,381,407]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
<emphasis id="71E5EAFF714A6235F9C1FEADFE84FE3A" box="[220,371,381,407]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6AD65784714A6235F866FEADFE21FE35" box="[379,470,382,408]" pageId="16" pageNumber="17" rank="species">sp. nov.</taxonomicNameLabel>
(
<figureCitation id="DBAA2A68714A6235F8FBFEAEFD91FE35" box="[486,614,381,408]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 9E, K</figureCitation>
<figureCitation id="DBAA2A68714A6235F8FBFEAEFD91FE35" box="[486,614,381,408]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Fig. 9E, K</figureCitation>
).
<taxonomicName id="84914D6E714A6235FB66FEAEFCE8FE3A" authorityName="Ran &amp; Liang" authorityYear="2006" box="[635,799,381,407]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="acuta">
<emphasis id="71E5EAFF714A6235FB66FEAEFCE8FE3A" box="[635,799,381,407]" italics="true" pageId="16" pageNumber="17">Flavina acuta</emphasis>
@ -130,7 +131,7 @@ also differs by a truncate apical margin of the anal tube in dorsal view (
<emphasis id="71E5EAFF714A6235FACAFE72FB99FE16" box="[983,1134,417,443]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
(
<figureCitation id="DBAA2A68714A6235FD63FE72FB2CFE16" box="[1150,1243,417,443]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 9D</figureCitation>
<figureCitation id="DBAA2A68714A6235FD63FE72FB2CFE16" box="[1150,1243,417,443]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Fig. 9D</figureCitation>
), and a much more elongate and narrow capitulum of the gonostylus without strong apical hook (
<bibRefCitation id="27004B1C714A6235FD8DFE17FA84FE73" author="Ran H. - F. &amp; Liang A. - P." box="[1168,1395,452,478]" pageId="16" pageNumber="17" pagination="388 - 391" refId="ref24916" refString="Ran H. - F. &amp; Liang A. - P. 2006. Taxonomic study of the issid genus Flavina Stal (Hemiptera, Fulgoroidea, Issidae). Acta Zootaxonomica Sinica 31 (2): 388 - 391." type="journal article" year="2006">Ran &amp; Liang 2006</bibRefCitation>
: figs 4, 6), which is wider, shorter and apicallly hooked in
@ -138,35 +139,35 @@ also differs by a truncate apical margin of the anal tube in dorsal view (
<emphasis id="71E5EAFF714A6235FA67FE3BFBE4FDAC" box="[890,1043,487,513]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
(
<figureCitation id="DBAA2A68714A6235FD3BFE34FB5DFDAC" box="[1062,1194,487,513]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 9A–B</figureCitation>
<figureCitation id="DBAA2A68714A6235FD3BFE34FB5DFDAC" box="[1062,1194,487,513]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Fig. 9A–B</figureCitation>
).
<taxonomicName id="84914D6E714A6235FDDFFE34FA8FFDAC" authorityName="Constant &amp; Pham" authorityYear="2024" box="[1218,1400,487,513]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="quangi">
<emphasis id="71E5EAFF714A6235FDDFFE34FA8FFDAC" box="[1218,1400,487,513]" italics="true" pageId="16" pageNumber="17">Flavina quangi</emphasis>
</taxonomicName>
also differs by the obsolete carinae of the mesonotum (
<figureCitation id="DBAA2A68714A6235FA24FDD9FC54FD89" box="[825,931,522,548]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="16" pageNumber="17">Fig. 16A</figureCitation>
<figureCitation id="DBAA2A68714A6235FA24FDD9FC54FD89" box="[825,931,522,548]" captionStart="Fig" captionStartId="31.[189,232,1916,1942]" captionTargetBox="[189,1398,280,1853]" captionTargetId="figure-12@31.[189,1398,270,1862]" captionTargetPageId="31" captionText="Fig. 16. Flavina quangi sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508770" httpUri="https://zenodo.org/record/14508770/files/figure.png" pageId="16" pageNumber="17">Fig. 16A</figureCitation>
) which are well marked in
<taxonomicName id="84914D6E714A6235FDC2FDD8FA81FD89" authorityName="Constant &amp; Pham" authorityYear="2024" box="[1247,1398,522,548]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="bachmana">
<emphasis id="71E5EAFF714A6235FDC2FDD8FA81FD89" box="[1247,1398,522,548]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
(
<figureCitation id="DBAA2A68714A6235F9D8FDFEFED4FDE5" box="[197,291,557,584]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="16" pageNumber="17">Fig. 8A</figureCitation>
<figureCitation id="DBAA2A68714A6235F9D8FDFEFED4FDE5" box="[197,291,557,584]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="16" pageNumber="17">Fig. 8A</figureCitation>
), the anal tube with truncate apex in dorsal view (
<figureCitation id="DBAA2A68714A6235FA6CFDFEFC2BFDE5" box="[881,988,557,584]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 17D</figureCitation>
<figureCitation id="DBAA2A68714A6235FA6CFDFEFC2BFDE5" box="[881,988,557,584]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508772" httpUri="https://zenodo.org/record/14508772/files/figure.png" pageId="16" pageNumber="17">Fig. 17D</figureCitation>
) which is rounded in
<taxonomicName id="84914D6E714A6235FDC2FDFDFA80FDEA" authorityName="Constant &amp; Pham" authorityYear="2024" box="[1247,1399,557,583]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="bachmana">
<emphasis id="71E5EAFF714A6235FDC2FDFDFA80FDEA" box="[1247,1399,557,583]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
(
<figureCitation id="DBAA2A68714A6235F9DBFD82FED2FDC6" box="[198,293,593,619]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 9D</figureCitation>
<figureCitation id="DBAA2A68714A6235F9DBFD82FED2FDC6" box="[198,293,593,619]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Fig. 9D</figureCitation>
), and with lateral lobes strongly developed, angularly rounded and projecting posteroventrad (
<figureCitation id="DBAA2A68714A6235F9DAFDA7FEADFD23" box="[199,346,628,654]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 17 A–B</figureCitation>
<figureCitation id="DBAA2A68714A6235F9DAFDA7FEADFD23" box="[199,346,628,654]" captionStart="Fig" captionStartId="33.[189,232,1892,1918]" captionTargetBox="[258,1331,266,1845]" captionTargetId="figure-11@33.[257,1331,265,1845]" captionTargetPageId="33" captionText="Fig. 17. Flavina quangi sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508772" httpUri="https://zenodo.org/record/14508772/files/figure.png" pageId="16" pageNumber="17">Fig. 17 A–B</figureCitation>
) instead of rounded in
<taxonomicName id="84914D6E714A6235FB75FDA6FD08FD23" authorityName="Constant &amp; Pham" authorityYear="2024" box="[616,767,628,654]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="bachmana">
<emphasis id="71E5EAFF714A6235FB75FDA6FD08FD23" box="[616,767,628,654]" italics="true" pageId="16" pageNumber="17">F. bachmana</emphasis>
</taxonomicName>
(
<figureCitation id="DBAA2A68714A6235FA12FDA7FC66FD23" box="[783,913,628,654]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Fig. 9A–B</figureCitation>
<figureCitation id="DBAA2A68714A6235FA12FDA7FC66FD23" box="[783,913,628,654]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Fig. 9A–B</figureCitation>
).
</paragraph>
</subSubSection>
@ -195,7 +196,7 @@ also differs by the obsolete carinae of the mesonotum (
•
<specimenCount id="5597FD64714A6235F846FCABFE86FC3F" box="[347,369,888,914]" count="1" pageId="16" pageNumber="17" type="male">♂</specimenCount>
(dissected –
<figureCitation id="DBAA2A68714A6235FB1AFCA4FD98FC3C" box="[519,623,887,913]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="17.[189,232,1928,1954]" captionStartId-1="18.[189,232,1892,1918]" captionStartId-2="20.[189,232,1892,1918]" captionTargetBox-0="[245,1337,271,1876]" captionTargetBox-1="[242,1345,274,1824]" captionTargetBox-2="[257,1332,283,1846]" captionTargetId-0="figure-12@17.[242,1345,265,1886]" captionTargetId-1="figure-17@18.[242,1345,265,1827]" captionTargetId-2="figure-16@20.[256,1332,265,1847]" captionTargetPageId-0="17" captionTargetPageId-1="18" captionTargetPageId-2="20" captionText-0="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="16" pageNumber="17">Figs 7–9</figureCitation>
<figureCitation id="DBAA2A68714A6235FB1AFCA4FD98FC3C" box="[519,623,887,913]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="17.[189,232,1928,1954]" captionStartId-1="18.[189,232,1892,1918]" captionStartId-2="20.[189,232,1892,1918]" captionTargetBox-0="[245,1337,271,1876]" captionTargetBox-1="[242,1345,274,1824]" captionTargetBox-2="[257,1332,283,1846]" captionTargetId-0="figure-12@17.[242,1345,265,1886]" captionTargetId-1="figure-17@18.[242,1345,265,1827]" captionTargetId-2="figure-16@20.[256,1332,265,1847]" captionTargetPageId-0="17" captionTargetPageId-1="18" captionTargetPageId-2="20" captionText-0="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508748" figureDoi-1="http://doi.org/10.5281/zenodo.14508752" figureDoi-2="http://doi.org/10.5281/zenodo.14508756" httpUri-0="https://zenodo.org/record/14508748/files/figure.png" httpUri-1="https://zenodo.org/record/14508752/files/figure.png" httpUri-2="https://zenodo.org/record/14508756/files/figure.png" pageId="16" pageNumber="17">Figs 7–9</figureCitation>
);
<location id="464E6036714A6235FB9EFCA4FD54FC3C" LSID="urn:lsid:plazi:treatment:CB3887FB71556233FB6BFA4FFE79F966:464E6036714A6235FB9EFCA4FD54FC3C" box="[643,675,887,913]" country="Vietnam" county="Bach Ma National Park" latitude="16.195555" longLatPrecision="21" longitude="107.84556" municipality="Province" name="Th" pageId="16" pageNumber="17">Th</location>
ừa Thiên-Huế
@ -457,10 +458,10 @@ leg.;
<paragraph id="432E36ED714A6236F9A0F984FDA4FE7E" blockId="16.[188,1401,1623,2001]" lastBlockId="19.[189,1400,265,467]" lastPageId="19" lastPageNumber="20" pageId="16" pageNumber="17">
<smallCapsWord id="45C8A031714A6235F9A0F984FF0BF9C2" baselines="1642,1642" box="[189,252,1623,1649]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="16" pageNumber="17">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A68714A6235F80BF984FE56F9DC" box="[278,417,1623,1649]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="16" pageNumber="17">Fig. 8A–D</figureCitation>
<figureCitation id="DBAA2A68714A6235F80BF984FE56F9DC" box="[278,417,1623,1649]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="16" pageNumber="17">Fig. 8A–D</figureCitation>
). Narrower than thorax and rather elongate, with about half of vertex length surpassing eyes. Vertex yellowish with black markings apically, and laterally in distal portion, strongly concave (longitudinally grooved), about 0.9 times as long in midline as broad basally, with sides evenly converging towards rounded apex; rather weak median carina, sometimes nearly obsolete, not reaching apex, yellowish; all margins carinate, lateral ones strongly elevated; posterior margin angularly concave. Frons blackish brown, darker along sides and dorsal margin, with minute yellowish points in central portion and often rather large yellowish marking in middle; series of small yellowish tubercles along lateral margins; median carina well marked, extending from dorsal margin to slightly before clypeal suture; dorsal margin concave and narrow, 0.27 times as wide as length of frons in midline. Posterior side of head yellowish. Genae yellowish with black-brown marking between eye and anterodorsal angle, between anteroventral angle of eye and lateral margin of frons, and above clypeus. Clypeus coloured as frons, often with mediobasal yellowish spot, flat in middle portion, moderately elongate, subtriangular with fronto-clypeal suture moderately rounded; anteclypeus mostly yellowish, brown apically, with median blunt carina. Labium yellowish, elongate and narrow, nearly reaching metacoxae, with apical segment elongate, nearly as long as penultimate. Eyes reniform (not emarginate) protruding laterally; ocelli absent. Antennae rather short with scape yellowish, ring-shaped, and pedicel dark brown, barrel-shaped, slightly longer than broad.
</paragraph>
<caption id="17EE6665714B6234F9A0F85BFE26F845" pageId="17" pageNumber="18" startId="17.[189,232,1928,1954]" targetBox="[245,1337,271,1876]" targetPageId="17">
<caption id="17EE6665714B6234F9A0F85BFE26F845" ID-DOI="http://doi.org/10.5281/zenodo.14508748" ID-Zenodo-Dep="14508748" httpUri="https://zenodo.org/record/14508748/files/figure.png" pageId="17" pageNumber="18" startId="17.[189,232,1928,1954]" targetBox="[245,1337,271,1876]" targetPageId="17">
<paragraph id="432E36ED714B6234F9A0F85BFE26F845" blockId="17.[188,1398,1928,2024]" pageId="17" pageNumber="18">
<emphasis id="71E5EAFF714B6234F9A0F85BFEFDF80F" bold="true" box="[189,266,1928,1954]" pageId="17" pageNumber="18">Fig. 7.</emphasis>
<taxonomicName id="84914D6E714B6234F812F85BFE1CF80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[271,491,1928,1954]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
@ -480,7 +481,7 @@ leg.;
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
<caption id="17EE666571486237F9A0F8B7FEDFF845" pageId="18" pageNumber="19" startId="18.[189,232,1892,1918]" targetBox="[242,1345,274,1824]" targetPageId="18">
<caption id="17EE666571486237F9A0F8B7FEDFF845" ID-DOI="http://doi.org/10.5281/zenodo.14508752" ID-Zenodo-Dep="14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="18" pageNumber="19" startId="18.[189,232,1892,1918]" targetBox="[242,1345,274,1824]" targetPageId="18">
<paragraph id="432E36ED71486237F9A0F8B7FEDFF845" blockId="18.[189,1399,1892,2024]" pageId="18" pageNumber="19">
<emphasis id="71E5EAFF71486237F9A0F8B7FEE7F8D2" bold="true" box="[189,272,1892,1919]" pageId="18" pageNumber="19">Fig. 8.</emphasis>
<taxonomicName id="84914D6E71486237F801F8B6FE09F8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[284,510,1893,1919]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
@ -506,20 +507,20 @@ leg.;
<paragraph id="432E36ED71496236F9A0FE28FC83FCD9" blockId="19.[187,1400,506,884]" pageId="19" pageNumber="20">
<smallCapsWord id="45C8A03171496236F9A0FE28FEEEFDBE" baselines="526,526" box="[189,281,507,533]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="19" pageNumber="20">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A6871496236F82DFE29FE4EFDB8" box="[304,441,506,533]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="19" pageNumber="20">Fig. 8A–D</figureCitation>
<figureCitation id="DBAA2A6871496236F82DFE29FE4EFDB8" box="[304,441,506,533]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="19" pageNumber="20">Fig. 8A–D</figureCitation>
). Pronotum with disc (delimited by peridiscal carina) yellowish, with central depression often washed with brownish, lateral fields blackish brown, paranotal lobes coloured like frons, darker along posterior margin, yellowish tubercles on disc and along posterior margin of lateral fields and paranotal lobes; 0.63 times as long as mesonotum in midline; anterior margin strongly protruding anteriorly between eyes, with carinae directed obliquely posteriorly, not reaching hind margin of pronotum; posterior margin nearly straight, slightly elevated; median carina obsolete with impressed point on each side; paranotal lobes (lateral view) broad with hind margin rounded, somewhat angularly rounded posteroventrally. Mesonotum variegated yellowish, often with two black points on each side; subtriangular with nearly complete transverse anterior carina and with obsolete median carina but with supplementary carinae between midline and peridiscal carina and in lateral fields, respectively; disc smooth, slightly depressed in midline. Tegulae dark brown.
</paragraph>
<paragraph id="432E36ED71496236F9A0FC4FFDD7FA94" blockId="19.[188,1400,924,1338]" pageId="19" pageNumber="20">
<smallCapsWord id="45C8A03171496236F9A0FC4FFED5FC19" baselines="943,943" box="[189,290,924,950]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="19" pageNumber="20">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A6871496236F824FC4FFE35FC1B" box="[313,450,924,950]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="19" pageNumber="20">
<figureCitation id="DBAA2A6871496236F824FC4FFE35FC1B" box="[313,450,924,950]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508748" httpUri="https://zenodo.org/record/14508748/files/figure.png" pageId="19" pageNumber="20">
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7A–C
</figureCitation>
,
<figureCitation id="DBAA2A6871496236F8C9FC4FFE02FC1B" box="[468,501,924,950]" captionStart="Fig" captionStartId="21.[189,232,1963,1989]" captionTargetBox="[189,1398,1438,1920]" captionTargetId="figure-455@21.[189,1398,1438,1920]" captionTargetPageId="21" captionText="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." pageId="19" pageNumber="20">10</figureCitation>
<figureCitation id="DBAA2A6871496236F8C9FC4FFE02FC1B" box="[468,501,924,950]" captionStart="Fig" captionStartId="21.[189,232,1963,1989]" captionTargetBox="[189,1398,1438,1920]" captionTargetId="figure-455@21.[189,1398,1438,1920]" captionTargetPageId="21" captionText="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." figureDoi="http://doi.org/10.5281/zenodo.14508758" httpUri="https://zenodo.org/record/14508758/files/figure.png" pageId="19" pageNumber="20">10</figureCitation>
). Variegated yellowish brown, sometimes washed with green, with apex of clavus blackish brown and more or less extensive, scattered, irregular blackish brown markings; sometimes with posteromedian whitish marking; tegmina subcoriaceous with longitudinal veins moderately elevated and with dense reticulum of paler veinlets; shape elongate and convex with sides broadly rounded, subparallel in middle portion, about 2.6 times as long as wide; rather narrowly rounded apically. Postclaval margin straight, slightly notched at apex of clavus. Clavus closed, reaching about 3/5 of tegmen. Venation (
<figureCitation id="DBAA2A6871496236F8A1FBBCFDC9FB24" box="[444,574,1135,1161]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="19" pageNumber="20">Fig. 7A, C</figureCitation>
<figureCitation id="DBAA2A6871496236F8A1FBBCFDC9FB24" box="[444,574,1135,1161]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508748" httpUri="https://zenodo.org/record/14508748/files/figure.png" pageId="19" pageNumber="20">Fig. 7A, C</figureCitation>
): ScP+R moderately developed, forking into subparallel ScP+RA and RP; MP forking rather close to base into subparallel MP
<subScript id="DF1534A871496236FA5EFB76FCA9FB1E" attach="left" box="[835,862,1189,1203]" fontSize="6" pageId="19" pageNumber="20">1+2</subScript>
and MP
@ -540,16 +541,16 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
<smallCapsWord id="45C8A03171496236F9A0FAB3FF00FAD4" baselines="1395,1396" box="[189,247,1376,1402]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="19" pageNumber="20">HIND</smallCapsWord>
<smallCapsWord id="45C8A03171496236F81EFAB6FEB0FAD5" baselines="1395" box="[259,327,1381,1400]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="19" pageNumber="20">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A6871496236F841FAB3FE4CFAD7" box="[348,443,1376,1402]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="19" pageNumber="20">Fig. 8E</figureCitation>
<figureCitation id="DBAA2A6871496236F841FAB3FE4CFAD7" box="[348,443,1376,1402]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="19" pageNumber="20">Fig. 8E</figureCitation>
). Dark brown with paler area basally and along costal margin, following vein ScP+R; veins mostly coloured as background; wing broader than tegmen and deeply bilobed at CuP; costal margin rather weakly sinuate; CuP-Pcu-A1 lobe about 1.5 times as wide as ScP-R-MP-CuA lobe, the two lobes almost the same length; both lobes somewhat angularly rounded apically; postclaval margin broadly rounded; A2 lobe brown, reduced and narrow. Venation: main veins present; ScP+R merging with costal margin at nodal line, MP and CuA running more or less parallel, with numerous cross-veins; Pcu strongly curved around basal half of wing towards CuP; A1 curved, more or less parallel to postclaval margin; CuP-Pcu-A1 lobe with dense network of cross-veins; vein A2 distinct.
</paragraph>
<paragraph id="432E36ED71496230F9A0F947FAB9FEEB" blockId="19.[188,1399,1683,1987]" lastBlockId="21.[189,1399,265,326]" lastPageId="21" lastPageNumber="22" pageId="19" pageNumber="20">
<smallCapsWord id="45C8A03171496236F9A0F947FF01F901" baselines="1703,1703" box="[189,246,1684,1710]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="19" pageNumber="20">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A6871496236F816F940FE7AF903" box="[267,397,1683,1710]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="19" pageNumber="20">Fig. 7A–E</figureCitation>
<figureCitation id="DBAA2A6871496236F816F940FE7AF903" box="[267,397,1683,1710]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508748" httpUri="https://zenodo.org/record/14508748/files/figure.png" pageId="19" pageNumber="20">Fig. 7A–E</figureCitation>
). Coxae brown; trochanters variegated yellowish and brown. Pro- and mesofemora brown with median yellowish ring and more or less dense yellowish pitting in brown zones; ventral face darker than dorsal; elongate, rather slender and slightly flattened dorsoventrally. Pro- and mesotibiae yellowish with some brown pitting forming more or less distinct subbasal ring; apex brown; tibiae weakly flattened dorsoventrally, narrower and longer than corresponding femora. Pro- and mesotarsi elongate and brown. Metafemora dark brown with elongate yellowish marking along posterodorsal portion ending in subapical dorsal band. Metatibiae variegated yellowish with some brown pitting, with one lateral spine near base (sometimes weak to obsolete), 2–3 lateral spines and 8 apical spines, all spines brown, distally black. First metatarsomere elongate and slender, with a strong spine at each side and a row of 6 smaller spines in between ventrally along posterior margin; second metatarsomere short with one spine at each side, all spines black-brown apically. Metatibiotarsal formula: (3–4) 8 / 8 / 2.
</paragraph>
<caption id="17EE6665714E6231F9A0F8B7FB8FF845" pageId="20" pageNumber="21" startId="20.[189,232,1892,1918]" targetBox="[257,1332,283,1846]" targetPageId="20">
<caption id="17EE6665714E6231F9A0F8B7FB8FF845" ID-DOI="http://doi.org/10.5281/zenodo.14508756" ID-Zenodo-Dep="14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="20" pageNumber="21" startId="20.[189,232,1892,1918]" targetBox="[257,1332,283,1846]" targetPageId="20">
<paragraph id="432E36ED714E6231F9A0F8B7FB8FF845" blockId="20.[189,1399,1892,2025]" pageId="20" pageNumber="21">
<emphasis id="71E5EAFF714E6231F9A0F8B7FEE7F8D2" bold="true" box="[189,272,1892,1919]" pageId="20" pageNumber="21">Fig. 9.</emphasis>
<taxonomicName id="84914D6E714E6231F801F8B6FE0AF8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[284,509,1893,1919]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
@ -588,7 +589,7 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
<paragraph id="432E36ED714F6230F9A0FEBEFA80FE2B" blockId="21.[189,1399,364,391]" box="[189,1399,364,391]" pageId="21" pageNumber="22">
<smallCapsWord id="45C8A031714F6230F9A0FEBEFEDBFE28" baselines="384,384" box="[189,300,365,391]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Abdomen" pageId="21" pageNumber="22">ABDOMEN</smallCapsWord>
(
<figureCitation id="DBAA2A68714F6230F821FEBFFE61FE2A" box="[316,406,364,391]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="21" pageNumber="22">Fig. 7B</figureCitation>
<figureCitation id="DBAA2A68714F6230F821FEBFFE61FE2A" box="[316,406,364,391]" captionStart="Fig" captionStartId="17.[189,232,1928,1954]" captionTargetBox="[245,1337,271,1876]" captionTargetId="figure-12@17.[242,1345,265,1886]" captionTargetPageId="17" captionText="Fig. 7. Flavina bachmana sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508748" httpUri="https://zenodo.org/record/14508748/files/figure.png" pageId="21" pageNumber="22">Fig. 7B</figureCitation>
). Brown, each sternite with large, darker, crescent-shaped marking along hind margin.
</paragraph>
<paragraph id="432E36ED714F6230F9A0FE7DFDF8FC7D" blockId="21.[187,1400,429,976]" pageId="21" pageNumber="22">
@ -597,17 +598,17 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
. Pygofer (
<emphasis id="71E5EAFF714F6230F8E2FE7DFDE9FE6A" box="[511,542,430,455]" italics="true" pageId="21" pageNumber="22">Py</emphasis>
–
<figureCitation id="DBAA2A68714F6230FB2BFE7EFD44FE65" box="[566,691,429,456]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="21" pageNumber="22">Fig. 9A–D</figureCitation>
<figureCitation id="DBAA2A68714F6230FB2BFE7EFD44FE65" box="[566,691,429,456]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="21" pageNumber="22">Fig. 9A–D</figureCitation>
) about 2.0 times as high as long (at midheight) in lateral view, with anterior margin weakly sinuate and posterior margin more or less straight, weakly oblique along dorsal ⅔ then abruptly rounded at posterodorsal angle, slightly excavate in ventral ⅓; posterior margin dorsally with deep, U-shaped excavation. Gonostyli (
<emphasis id="71E5EAFF714F6230FA2CFDC5FCB1FD9D" box="[817,838,534,560]" italics="true" pageId="21" pageNumber="22">G</emphasis>
–
<figureCitation id="DBAA2A68714F6230FA79FDC6FC16FD9D" box="[868,993,533,560]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="21" pageNumber="22">Fig. 9A–C</figureCitation>
<figureCitation id="DBAA2A68714F6230FA79FDC6FC16FD9D" box="[868,993,533,560]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="21" pageNumber="22">Fig. 9A–C</figureCitation>
) (in lateral view) longer than high (without dorsal capitulum), projecting posteriorly in distinct elongate lobe rounded apically and with ventral margin more or less straight; capitulum (
<emphasis id="71E5EAFF714F6230FBE7FD8FFCE1FDD8" box="[762,790,604,629]" italics="true" pageId="21" pageNumber="22">ca</emphasis>
–
<figureCitation id="DBAA2A68714F6230FA28FD88FC45FDD8" box="[821,946,603,629]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="21" pageNumber="22">Fig. 9A–C</figureCitation>
<figureCitation id="DBAA2A68714F6230FA28FD88FC45FDD8" box="[821,946,603,629]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="21" pageNumber="22">Fig. 9A–C</figureCitation>
) with rather long, wide neck, directed dorsad, with distal portion moderately elongate and twisted, with apical hook directed dorsocephalad and with basal lamina projection directed lateroventrad in caudal view. Aedeagus (
<figureCitation id="DBAA2A68714F6230FD45FD73FB39FD17" box="[1112,1230,672,698]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="21" pageNumber="22">Fig. 9E–J</figureCitation>
<figureCitation id="DBAA2A68714F6230FD45FD73FB39FD17" box="[1112,1230,672,698]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="21" pageNumber="22">Fig. 9E–J</figureCitation>
) symmetrical, evenly curved dorsad (in lateral view); pair of pointed, short, slender, straight lateroventral processes (
<emphasis id="71E5EAFF714F6230F9DAFD36FF1CFD52" box="[199,235,741,767]" italics="true" pageId="21" pageNumber="22">lvp</emphasis>
) directed cephalad; ventral lobe of periandrium (
@ -623,7 +624,7 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
) well developed, widely open. Anal tube (
<emphasis id="71E5EAFF714F6230FB7AFC9DFD7FFCC5" box="[615,648,846,872]" italics="true" pageId="21" pageNumber="22">An</emphasis>
–
<figureCitation id="DBAA2A68714F6230FBBBFC9EFCD2FCC5" box="[678,805,845,872]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="21" pageNumber="22">Fig. 9A–D</figureCitation>
<figureCitation id="DBAA2A68714F6230FBBBFC9EFCD2FCC5" box="[678,805,845,872]" captionStart="Fig" captionStartId="20.[189,232,1892,1918]" captionTargetBox="[257,1332,283,1846]" captionTargetId="figure-16@20.[256,1332,265,1847]" captionTargetPageId="20" captionText="Fig. 9. Flavina bachmana sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–K. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Posteroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508756" httpUri="https://zenodo.org/record/14508756/files/figure.png" pageId="21" pageNumber="22">Fig. 9A–D</figureCitation>
) dorsoventrally flattened, in dorsal view widening from base towards apex, forming narrow, rounded lateral lobe and with large middle portion of apical margin rounded; lateral lobes distinct in lateral view, more or less rounded and moderately developed; anal opening before basal ½.
</paragraph>
<paragraph id="432E36ED714F6230F9A0FC24FE85FB9E" blockId="21.[189,1399,1014,1075]" pageId="21" pageNumber="22">
@ -632,7 +633,7 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
. Hind margin of sternum VII (
<emphasis id="71E5EAFF714F6230FA3CFC24FC9FFBBD" box="[801,872,1015,1041]" italics="true" pageId="21" pageNumber="22">St VII</emphasis>
–
<figureCitation id="DBAA2A68714F6230FA93FC25FC10FBBD" box="[910,999,1014,1040]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="21" pageNumber="22">Fig. 8F</figureCitation>
<figureCitation id="DBAA2A68714F6230FA93FC25FC10FBBD" box="[910,999,1014,1040]" captionStart="Fig" captionStartId="18.[189,232,1892,1918]" captionTargetBox="[242,1345,274,1824]" captionTargetId="figure-17@18.[242,1345,265,1827]" captionTargetPageId="18" captionText="Fig. 8. Flavina bachmana sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508752" httpUri="https://zenodo.org/record/14508752/files/figure.png" pageId="21" pageNumber="22">Fig. 8F</figureCitation>
) concave, moderately inflated in middle portion.
</paragraph>
</subSubSection>
@ -650,15 +651,15 @@ not reaching apex of tegmen, only running to about MP 1 -MP 2 fork; CuA unforked
was collected between April and July in montane evergreen tropical forest as well as in lowland evergreen forest in Bach Ma, at altitudes between 500 and
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a.s.l. (
<figureCitation id="DBAA2A68714F6230FDFAFB79FA9EFB68" box="[1255,1385,1194,1221]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="21" pageNumber="22">Fig. 2A–D</figureCitation>
<figureCitation id="DBAA2A68714F6230FDFAFB79FA9EFB68" box="[1255,1385,1194,1221]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="21" pageNumber="22">Fig. 2A–D</figureCitation>
); in Da Krong the species was found at a lower altitude,
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a.s.l. The species lives on tree trunks densely covered in lichens and mosses, on which it is very cryptic, but some specimens were also found sitting on leaves of lower plants and bushes, and collected by sweeping (
<figureCitation id="DBAA2A68714F6230FD0DFAC1FB90FA80" box="[1040,1127,1298,1325]" captionStart="Fig" captionStartId="21.[189,232,1963,1989]" captionTargetBox="[189,1398,1438,1920]" captionTargetId="figure-455@21.[189,1398,1438,1920]" captionTargetPageId="21" captionText="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." pageId="21" pageNumber="22">Fig. 10</figureCitation>
<figureCitation id="DBAA2A68714F6230FD0DFAC1FB90FA80" box="[1040,1127,1298,1325]" captionStart="Fig" captionStartId="21.[189,232,1963,1989]" captionTargetBox="[189,1398,1438,1920]" captionTargetId="figure-455@21.[189,1398,1438,1920]" captionTargetPageId="21" captionText="Fig. 10. Flavina bachmana sp. nov., live specimens in Bach Ma National Park.A. 14 May 2023, summit. B. 12 May 2023, Pheasant trail. C. 13 May 2023, summit." figureDoi="http://doi.org/10.5281/zenodo.14508758" httpUri="https://zenodo.org/record/14508758/files/figure.png" pageId="21" pageNumber="22">Fig. 10</figureCitation>
). When disturbed, they tend to walk away and hide on the opposite side of the trunk or branch, and jump away as a last solution.
</paragraph>
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<paragraph id="432E36ED714F6230F9A0F878FC90F845" blockId="21.[187,1399,1963,2025]" pageId="21" pageNumber="22">
<emphasis id="71E5EAFF714F6230F9A0F878FEEEF868" bold="true" box="[189,281,1963,1989]" pageId="21" pageNumber="22">Fig. 10.</emphasis>
<taxonomicName id="84914D6E714F6230F803F878FE0EF868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[286,505,1963,1989]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
@ -674,7 +675,7 @@ a.s.l. The species lives on tree trunks densely covered in lichens and mosses, o
. 13 May 2023, summit.
</paragraph>
</caption>
<caption id="17EE6665714C6233F9A0FA2AFCC1F9B9" box="[189,822,1529,1556]" pageId="22" pageNumber="23" startId="22.[189,232,1529,1555]" targetBox="[189,1398,265,1486]" targetPageId="22">
<caption id="17EE6665714C6233F9A0FA2AFCC1F9B9" ID-DOI="http://doi.org/10.5281/zenodo.14508760" ID-Zenodo-Dep="14508760" box="[189,822,1529,1556]" httpUri="https://zenodo.org/record/14508760/files/figure.png" pageId="22" pageNumber="23" startId="22.[189,232,1529,1555]" targetBox="[189,1398,265,1486]" targetPageId="22">
<paragraph id="432E36ED714C6233F9A0FA2AFCC1F9B9" blockId="22.[189,822,1529,1556]" box="[189,822,1529,1556]" pageId="22" pageNumber="23">
<emphasis id="71E5EAFF714C6233F9A0FA2AFEEEF9BE" bold="true" box="[189,281,1529,1555]" pageId="22" pageNumber="23">Fig. 11.</emphasis>
<taxonomicName id="84914D6E714C6233F83DFA2AFE0AF9BE" authorityName="Constant &amp; Pham" authorityYear="2024" box="[288,509,1529,1555]" class="Insecta" family="Issidae" genus="Flavina" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="species" species="bachmana" status="sp. nov.">
@ -695,7 +696,7 @@ a.s.l. The species lives on tree trunks densely covered in lichens and mosses, o
: Thừa Thiên-Huế Province, Bach Ma National Park, and
<collectingRegion id="8155F80F714C6233FAA6F95EFB55F90A" box="[955,1186,1677,1703]" country="Vietnam" name="Quang Tri" pageId="22" pageNumber="23">Quang Tri Province</collectingRegion>
, Da Krong Nature Reserve (
<figureCitation id="DBAA2A68714C6233F837F963FE77F966" box="[298,384,1712,1739]" captionStart="Fig" captionStartId="22.[189,232,1529,1555]" captionTargetBox="[189,1398,265,1486]" captionTargetId="figure-74@22.[189,1398,265,1486]" captionTargetPageId="22" captionText="Fig. 11. Flavina bachmana sp. nov., distribution map." pageId="22" pageNumber="23">Fig. 11</figureCitation>
<figureCitation id="DBAA2A68714C6233F837F963FE77F966" box="[298,384,1712,1739]" captionStart="Fig" captionStartId="22.[189,232,1529,1555]" captionTargetBox="[189,1398,265,1486]" captionTargetId="figure-74@22.[189,1398,265,1486]" captionTargetPageId="22" captionText="Fig. 11. Flavina bachmana sp. nov., distribution map." figureDoi="http://doi.org/10.5281/zenodo.14508760" httpUri="https://zenodo.org/record/14508760/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
).
</paragraph>
</subSubSection>

89
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@ -159,7 +160,7 @@ genus of the family
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<emphasis id="71E5EAFF715C6223F9A0F81DFEFCF845" bold="true" box="[189,267,1998,2024]" pageId="6" pageNumber="7">Fig. 1.</emphasis>
Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam.
@ -174,22 +175,22 @@ Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam.
<paragraph id="432E36ED715D6222F9A0FEE5FC91FD96" blockId="7.[189,1401,309,571]" pageId="7" pageNumber="8">
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(
<figureCitation id="DBAA2A68715D6222F813FEE6FE66FEFD" box="[270,401,309,336]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="7" pageNumber="8">Fig. 4A–D</figureCitation>
<figureCitation id="DBAA2A68715D6222F813FEE6FE66FEFD" box="[270,401,309,336]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="7" pageNumber="8">Fig. 4A–D</figureCitation>
). Head slightly narrower than thorax. Vertex wider than long, medially grooved, with sides subparallel; anterior and posterior margins subparallel and curved; all margins weakly carinate. Frons elongate with large oculiform black, shiny marking in dorsal portion, median carina in ventral portion and tubercles along lateral margins under midheight of eye. Clypeus flat in middle portion, moderately elongate, subtriangular with fronto-clypeal suture rounded; anteclypeus with median blunt carina. Labium elongate and narrow, reaching metacoxae, with apical segment elongate, nearly as long as penultimate. Eyes reniform (not emarginate); ocelli absent. Antennae rather short with scape ring-shaped and pedicel cylindrical, slightly longer than broad.
</paragraph>
<paragraph id="432E36ED715D6222F9A0FDB2FD28FCE9" blockId="7.[189,1400,608,836]" pageId="7" pageNumber="8">
<smallCapsWord id="45C8A031715D6222F9A0FDB2FEEEFDD4" baselines="628,628" box="[189,281,609,635]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="7" pageNumber="8">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A68715D6222F831FDB3FE46FDD6" box="[300,433,608,635]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="7" pageNumber="8">Fig. 4A–D</figureCitation>
<figureCitation id="DBAA2A68715D6222F831FDB3FE46FDD6" box="[300,433,608,635]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="7" pageNumber="8">Fig. 4A–D</figureCitation>
). Pronotum slightly shorter than mesonotum in midline; anterior margin carinate in middle portion, strongly sinuate and strongly, roundly protruding anteriorly between eyes; posterior margin nearly straight, slightly carinate in middle portion; median carina obsolete anteriorly with impressed point on each side; blunt tubercles along anterior margin and irregularly on disc and sides; paranotal lobes (lateral view) broad, with tubercles along posterior margin, and with posteroventral angle rounded. Mesonotum subtriangular with obsolete median carina and weak peridiscal carinae. Tegulae moderately developed, medium sized.
</paragraph>
<paragraph id="432E36ED715D622DF9A0FCB9FABDFE66" blockId="7.[189,1398,874,934]" lastBlockId="8.[187,1400,265,460]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
<smallCapsWord id="45C8A031715D6222F9A0FCB9FED5FC2F" baselines="893,893" box="[189,290,874,900]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="7" pageNumber="8">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A68715D6222F829FCB9FE42FC29" box="[308,437,874,900]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="7" pageNumber="8">Fig. 3A–C</figureCitation>
<figureCitation id="DBAA2A68715D6222F829FCB9FE42FC29" box="[308,437,874,900]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508738" httpUri="https://zenodo.org/record/14508738/files/figure.png" pageId="7" pageNumber="8">Fig. 3A–C</figureCitation>
). Tegmina subcoriaceous with longitudinal veins slightly elevated and with rather dense reticulum of veinlets, elongate with sides broadly rounded, slightly more than 2 times as long as broad, convex, without epipleuron. Apex narrowly rounded. Postclaval margin weakly rounded on distal half and slightly notched at apex of clavus. Clavus closed, reaching about 3/5 of tegmen. Venation: ScP+R rather short, forking into subparallel ScP+RA and RP; RP forking at about midlength of tegmen; MP first fork at about half of tegmen length; CuA straight, unforked and reaching hind margin; additional vein in cubital cell, parallel to postclaval margin, originating from level of apex of clavus and not reaching apical margin; Pcu fused with A1 at 3/5 of clavus length; Pcu+A1 fused with CuP at apex of clavus.
</paragraph>
<caption id="17EE6665715D6222F9A3F878FBF0F845" pageId="7" pageNumber="8" startId="7.[190,233,1963,1989]" targetBox="[189,1398,1011,1918]" targetPageId="7">
<caption id="17EE6665715D6222F9A3F878FBF0F845" ID-DOI="http://doi.org/10.5281/zenodo.14508734" ID-Zenodo-Dep="14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="7" pageNumber="8" startId="7.[190,233,1963,1989]" targetBox="[189,1398,1011,1918]" targetPageId="7">
<paragraph id="432E36ED715D6222F9A3F878FBF0F845" blockId="7.[189,1399,1963,2025]" pageId="7" pageNumber="8">
<emphasis id="71E5EAFF715D6222F9A3F878FEFAF868" bold="true" box="[190,269,1963,1989]" pageId="7" pageNumber="8">Fig. 2.</emphasis>
Typical habitats in Bach Ma National Park, May 2023.
@ -207,13 +208,13 @@ Typical habitats in Bach Ma National Park, May 2023.
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<smallCapsWord id="45C8A0317152622DF9E3FE25FEB5FDA4" baselines="516" box="[254,322,502,521]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="8" pageNumber="9">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A687152622DF84EFE22FE59FDA6" box="[339,430,497,523]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." pageId="8" pageNumber="9">Fig. 4E</figureCitation>
<figureCitation id="DBAA2A687152622DF84EFE22FE59FDA6" box="[339,430,497,523]" captionStart="Fig" captionStartId="11.[189,232,1850,1876]" captionTargetBox="[189,1398,277,1784]" captionTargetId="figure-12@11.[189,1398,265,1806]" captionTargetPageId="11" captionText="Fig. 4. Cyclopissus corticalis gen. et sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508740" httpUri="https://zenodo.org/record/14508740/files/figure.png" pageId="8" pageNumber="9">Fig. 4E</figureCitation>
). Broader than tegmina and deeply bilobed, strongly notched at CuP; costal margin sinuate; CuP-Pcu-A1 lobe much wider than Sc-R-MP-CuA lobe, and slightly longer than latter; both lobes rounded apically; postclaval margin broadly rounded; anal lobe reduced and narrow, with weak A2 vein. Venation: main veins present; ScP+R, MP and CuA running more or less parallel in basal portion, with ScP+R diverging beyond basal ⅓, with numerous cross-veins; Pcu strongly curved around basal third of wing towards CuP and reaching the latter; A1 curved, more or less parallel to postclaval margin; CuP-Pcu-A1 lobe with numerous cross-veins.
</paragraph>
<paragraph id="432E36ED7152622DF9A0FD28FA90FCD4" blockId="8.[188,1399,762,890]" pageId="8" pageNumber="9">
<smallCapsWord id="45C8A0317152622DF9A0FD28FF01FCBE" baselines="782,782" box="[189,246,763,789]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="8" pageNumber="9">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A687152622DF81AFD29FEBBFCB9" box="[263,332,762,789]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="8" pageNumber="9">Fig. 3</figureCitation>
<figureCitation id="DBAA2A687152622DF81AFD29FEBBFCB9" box="[263,332,762,789]" captionStart="Fig" captionStartId="10.[189,232,1928,1954]" captionTargetBox="[244,1333,271,1880]" captionTargetId="figure-16@10.[244,1344,265,1883]" captionTargetPageId="10" captionText="Fig. 3. Cyclopissus corticalis gen. et sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508738" httpUri="https://zenodo.org/record/14508738/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
). Elongate and slender. Tibiae slightly longer than corresponding femora. Metatibiae with 2 lateral spines in distal half and 7 apical spines. Tarsi elongate; first metatarsomere elongate and slender, with a strong spine at each side and a row of 5 smaller spines in between ventrally along posterior margin; second metatarsomere short with one spine at each side. Metatibiotarsal formula: (2) 7 / 8 / 2.
</paragraph>
<paragraph id="432E36ED7152622DF9A0FC4CFD8AFBCC" blockId="8.[189,1400,927,1121]" pageId="8" pageNumber="9">

131
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<treatment id="CB3887FB716F621EFB4AFC14FB9DFC79" LSID="urn:lsid:plazi:treatment:CB3887FB716F621EFB4AFC14FB9DFC79" httpUri="http://treatment.plazi.org/id/CB3887FB716F621EFB4AFC14FB9DFC79" lastPageId="59" lastPageNumber="60" pageId="53" pageNumber="54">
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@ -67,11 +68,11 @@ urn:lsid:zoobank.org:act:
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<subSubSection id="0B8B6566716F6210FBADFBDEFC74FB85" box="[688,899,1037,1064]" pageId="53" pageNumber="54" type="description">
<paragraph id="432E36ED716F6210FBADFBDEFC74FB85" blockId="53.[372,1216,967,1064]" box="[688,899,1037,1064]" pageId="53" pageNumber="54">
<figureCitation id="DBAA2A68716F6210FBADFBDEFD0EFB8A" box="[688,761,1037,1063]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="53" pageNumber="54">Figs 1</figureCitation>
<figureCitation id="DBAA2A68716F6210FBADFBDEFD0EFB8A" box="[688,761,1037,1063]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="53" pageNumber="54">Figs 1</figureCitation>
,
<figureCitation id="DBAA2A68716F6210FA1AFBDEFCD9FB8A" box="[775,814,1037,1063]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." pageId="53" pageNumber="54">2D</figureCitation>
<figureCitation id="DBAA2A68716F6210FA1AFBDEFCD9FB8A" box="[775,814,1037,1063]" captionStart="Fig" captionStartId="7.[190,233,1963,1989]" captionTargetBox="[189,1398,1011,1918]" captionTargetId="figure-245@7.[189,1398,1011,1918]" captionTargetPageId="7" captionText="Fig. 2. Typical habitats in Bach Ma National Park, May 2023. A. Pheasant trail. B. Roadside (1200– 1300 m a.s.l.). C. Rhododendron trail. D. Summit (1300–1400 m a.s.l.)." figureDoi="http://doi.org/10.5281/zenodo.14508734" httpUri="https://zenodo.org/record/14508734/files/figure.png" pageId="53" pageNumber="54">2D</figureCitation>
,
<figureCitation id="DBAA2A68716F6210FA27FBDEFC74FB85" box="[826,899,1037,1064]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="55.[189,232,1928,1954]" captionStartId-1="56.[189,232,1892,1918]" captionStartId-2="57.[189,232,1892,1918]" captionStartId-3="58.[189,232,1963,1989]" captionTargetBox-0="[221,1374,273,1866]" captionTargetBox-1="[203,1384,453,1847]" captionTargetBox-2="[258,1331,285,1845]" captionTargetBox-3="[189,1398,1110,1917]" captionTargetId-0="figure-11@55.[211,1377,265,1885]" captionTargetId-1="figure-63@56.[203,1384,442,1850]" captionTargetId-2="figure-11@57.[257,1331,265,1845]" captionTargetId-3="figure-331@58.[189,1398,1110,1917]" captionTargetPageId-0="55" captionTargetPageId-1="56" captionTargetPageId-2="57" captionTargetPageId-3="58" captionText-0="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." captionText-2="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." pageId="53" pageNumber="54">32–35</figureCitation>
<figureCitation id="DBAA2A68716F6210FA27FBDEFC74FB85" box="[826,899,1037,1064]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="55.[189,232,1928,1954]" captionStartId-1="56.[189,232,1892,1918]" captionStartId-2="57.[189,232,1892,1918]" captionStartId-3="58.[189,232,1963,1989]" captionTargetBox-0="[221,1374,273,1866]" captionTargetBox-1="[203,1384,453,1847]" captionTargetBox-2="[258,1331,285,1845]" captionTargetBox-3="[189,1398,1110,1917]" captionTargetId-0="figure-11@55.[211,1377,265,1885]" captionTargetId-1="figure-63@56.[203,1384,442,1850]" captionTargetId-2="figure-11@57.[257,1331,265,1845]" captionTargetId-3="figure-331@58.[189,1398,1110,1917]" captionTargetPageId-0="55" captionTargetPageId-1="56" captionTargetPageId-2="57" captionTargetPageId-3="58" captionText-0="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." captionText-2="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." captionText-3="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." figureDoi-0="http://doi.org/10.5281/zenodo.14508802" figureDoi-1="http://doi.org/10.5281/zenodo.14508804" figureDoi-2="http://doi.org/10.5281/zenodo.14508806" figureDoi-3="http://doi.org/10.5281/zenodo.14508808" httpUri-0="https://zenodo.org/record/14508802/files/figure.png" httpUri-1="https://zenodo.org/record/14508804/files/figure.png" httpUri-2="https://zenodo.org/record/14508806/files/figure.png" httpUri-3="https://zenodo.org/record/14508808/files/figure.png" pageId="53" pageNumber="54">32–35</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="0B8B6566716F6210F9A0FB9DFAE7F8D8" pageId="53" pageNumber="54" type="diagnosis">
@ -89,28 +90,28 @@ can be recognized by:
</paragraph>
<paragraph id="432E36ED716F6210F9A0FB4CFC09FB14" blockId="53.[189,1399,1148,1521]" box="[189,1022,1182,1209]" pageId="53" pageNumber="54">
(1) the frons elongate, distinctly constricted between eyes (
<figureCitation id="DBAA2A68716F6210FA99FB4DFC07FB14" box="[900,1008,1182,1209]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." pageId="53" pageNumber="54">Fig. 33B</figureCitation>
<figureCitation id="DBAA2A68716F6210FA99FB4DFC07FB14" box="[900,1008,1182,1209]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="53" pageNumber="54">Fig. 33B</figureCitation>
);
</paragraph>
<paragraph id="432E36ED716F6210F9A0FB12FE17FA8C" blockId="53.[189,1399,1148,1521]" pageId="53" pageNumber="54">
(2) the anal tube in dorsal aspect widely spatulate with distal portion nearly circular, nearly 1.3 times as long in midline as wide with apical margin shortly truncate in middle, without distinct lateral flaps (
<emphasis id="71E5EAFF716F6210F81EFAD4FED3FA8C" box="[259,292,1287,1313]" italics="true" pageId="53" pageNumber="54">An</emphasis>
–
<figureCitation id="DBAA2A68716F6210F85CFAD5FE25FA8C" box="[321,466,1286,1313]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="53" pageNumber="54">Fig. 34A–D</figureCitation>
<figureCitation id="DBAA2A68716F6210F85CFAD5FE25FA8C" box="[321,466,1286,1313]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="53" pageNumber="54">Fig. 34A–D</figureCitation>
);
</paragraph>
<paragraph id="432E36ED716F6210F9A0FAFAFE83FACB" blockId="53.[189,1399,1148,1521]" pageId="53" pageNumber="54">
(3) the neck of the capitulum of the gonostyli rather short, less high than head of capitulum (
<emphasis id="71E5EAFF716F6210FC5CFAF9FAAAFAEE" box="[1345,1373,1322,1347]" italics="true" pageId="53" pageNumber="54">ca</emphasis>
–
<figureCitation id="DBAA2A68716F6210F9E4FA9FFE91FACB" box="[249,358,1356,1382]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="53" pageNumber="54">Fig. 34A</figureCitation>
<figureCitation id="DBAA2A68716F6210F9E4FA9FFE91FACB" box="[249,358,1356,1382]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="53" pageNumber="54">Fig. 34A</figureCitation>
);
</paragraph>
<paragraph id="432E36ED716F6210F9A0FABCFDD8FA5C" blockId="53.[189,1399,1148,1521]" pageId="53" pageNumber="54">
(4) the shape of the lateroventral processes of the aedeagus slightly inflated basally, then constricted, directed cephalad and moderately, evenly upcurved, remaining under the ventral lobe of the periandrium and reaching to basiventral lobes of periandrium; distal portion tapering and curved mesad (
<emphasis id="71E5EAFF716F6210F848FA05FE8EFA5D" box="[341,377,1494,1520]" italics="true" pageId="53" pageNumber="54">lvp</emphasis>
–
<figureCitation id="DBAA2A68716F6210F88AFA05FDD3FA5C" box="[407,548,1494,1521]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="53" pageNumber="54">Fig. 34F–K</figureCitation>
<figureCitation id="DBAA2A68716F6210F88AFA05FDD3FA5C" box="[407,548,1494,1521]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="53" pageNumber="54">Fig. 34F–K</figureCitation>
).
</paragraph>
<paragraph id="432E36ED716F6210F9A0F9C4FE3EF99C" blockId="53.[189,457,1559,1585]" box="[189,457,1559,1585]" pageId="53" pageNumber="54">
@ -191,7 +192,7 @@ also differs from the two latter species by its subparallel, slender lateroventr
•
<specimenCount id="5597FD64716C6213F843FE88FE83FED8" box="[350,372,347,373]" count="1" pageId="54" pageNumber="55" type="male">♂</specimenCount>
(
<figureCitation id="DBAA2A68716C6213F899FE89FDF9FED8" box="[388,526,346,373]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="55.[189,232,1928,1954]" captionStartId-1="56.[189,232,1892,1918]" captionStartId-2="57.[189,232,1892,1918]" captionTargetBox-0="[221,1374,273,1866]" captionTargetBox-1="[203,1384,453,1847]" captionTargetBox-2="[258,1331,285,1845]" captionTargetId-0="figure-11@55.[211,1377,265,1885]" captionTargetId-1="figure-63@56.[203,1384,442,1850]" captionTargetId-2="figure-11@57.[257,1331,265,1845]" captionTargetPageId-0="55" captionTargetPageId-1="56" captionTargetPageId-2="57" captionText-0="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." captionText-2="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="54" pageNumber="55">Figs 32–34</figureCitation>
<figureCitation id="DBAA2A68716C6213F899FE89FDF9FED8" box="[388,526,346,373]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="55.[189,232,1928,1954]" captionStartId-1="56.[189,232,1892,1918]" captionStartId-2="57.[189,232,1892,1918]" captionTargetBox-0="[221,1374,273,1866]" captionTargetBox-1="[203,1384,453,1847]" captionTargetBox-2="[258,1331,285,1845]" captionTargetId-0="figure-11@55.[211,1377,265,1885]" captionTargetId-1="figure-63@56.[203,1384,442,1850]" captionTargetId-2="figure-11@57.[257,1331,265,1845]" captionTargetPageId-0="55" captionTargetPageId-1="56" captionTargetPageId-2="57" captionText-0="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." captionText-2="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508802" figureDoi-1="http://doi.org/10.5281/zenodo.14508804" figureDoi-2="http://doi.org/10.5281/zenodo.14508806" httpUri-0="https://zenodo.org/record/14508802/files/figure.png" httpUri-1="https://zenodo.org/record/14508804/files/figure.png" httpUri-2="https://zenodo.org/record/14508806/files/figure.png" pageId="54" pageNumber="55">Figs 32–34</figureCitation>
); [Thừa Thiên-Huế Province],
<location id="464E6036716C6213FA64FE89FB65FED9" LSID="urn:lsid:plazi:treatment:CB3887FB716F621EFB4AFC14FB9DFC79:464E6036716C6213FA64FE89FB65FED9" box="[889,1170,346,373]" country="Vietnam" latitude="16.2" longLatPrecision="1282" longitude="107.86667" name="Bach Ma National Park" pageId="54" pageNumber="55">Bach Ma National Park</location>
, summit;
@ -227,7 +228,7 @@ leg.; day [time] collecting; I.G.: 31.933;
•
<specimenCount id="5597FD64716C6213F842FDD5FE7CFD8C" box="[351,395,518,545]" count="1" pageId="54" pageNumber="55" type="female">1 ♀</specimenCount>
(
<figureCitation id="DBAA2A68716C6213F881FDD5FE02FD8C" box="[412,501,518,545]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." pageId="54" pageNumber="55">Fig. 35</figureCitation>
<figureCitation id="DBAA2A68716C6213F881FDD5FE02FD8C" box="[412,501,518,545]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508808" httpUri="https://zenodo.org/record/14508808/files/figure.png" pageId="54" pageNumber="55">Fig. 35</figureCitation>
);
<location id="464E6036716C6213FB16FDD5FDDCFD8D" LSID="urn:lsid:plazi:treatment:CB3887FB716F621EFB4AFC14FB9DFC79:464E6036716C6213FB16FDD5FDDCFD8D" box="[523,555,518,544]" country="Vietnam" county="Bach Ma National Park" latitude="16.188332" longLatPrecision="21" longitude="107.84889" municipality="Province" name="Th" pageId="54" pageNumber="55">Th</location>
ừa Thiên-Huế
@ -278,21 +279,21 @@ leg.;
<paragraph id="432E36ED716C6213F9A0FD2CFE01FAAA" blockId="54.[187,1401,766,1287]" pageId="54" pageNumber="55">
<smallCapsWord id="45C8A031716C6213F9A0FD2CFF0BFCBA" baselines="786,786" box="[189,252,767,793]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="54" pageNumber="55">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A68716C6213F813FD2DFE68FCB4" box="[270,415,766,793]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." pageId="54" pageNumber="55">Fig. 33A–D</figureCitation>
<figureCitation id="DBAA2A68716C6213F813FD2DFE68FCB4" box="[270,415,766,793]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="54" pageNumber="55">Fig. 33A–D</figureCitation>
). Vertex variegated brown with anterior angles black, distinctly broader than long in midline and with lateral margins longer than midline, with all margins carinate; anterior margin nearly straight with median carina of frons distinct in middle, posterior margin deeply concave, angularly rounded, and lateral ones subparallel; disc shallowly excavate with obsolete median carina. Side of head yellowish brown with black marking at anterodorsal angle extending on a distance along anterior margin, darker area between anteroventral portion of eye and anterior margin, followed by pale yellowish area extending to base of antenna and anterior margin of gena; pale yellowish marking along clypeal suture; anteroventral angle moderately projecting anteriorly in a small round lobe. Frons elongate and generally smooth, variegated brown with dorsal angles blackish, distinct yellowish spot in middle on each side of median carina and more or less disctinct yellowish band along frontoclypeal suture; median carina extending from dorsal margin down nearly to rounded frontoclypeal suture, brown on disc; no distinct row of tubercles along dorsal and lateral margins; dorsal margin distinctly concave, lateral margins distinctly sinuate. Clypeus variegated dark brown, weakly elevated medially. Labium brown with last segment longer than broad, and shorter than penultimate. Scape short, ring-shaped, yellowish; pedicel bulbous, yellowish brown.
</paragraph>
<paragraph id="432E36ED716C6213F9A0FAFCFBE7F92A" blockId="54.[188,1400,1327,1671]" pageId="54" pageNumber="55">
<smallCapsWord id="45C8A031716C6213F9A0FAFCFEEEFAEA" baselines="1346,1346" box="[189,281,1327,1353]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="54" pageNumber="55">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A68716C6213F834FAFCFE2BFAE4" box="[297,476,1327,1353]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." pageId="54" pageNumber="55">Fig. 33A, C–D</figureCitation>
<figureCitation id="DBAA2A68716C6213F834FAFCFE2BFAE4" box="[297,476,1327,1353]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="54" pageNumber="55">Fig. 33A, C–D</figureCitation>
). Pronotum variegated yellowish brown; about 0.6 times as long as mesonotum in midline; anterior margin carinate, strongly, angularly protruding anteriorly between eyes, with carinae directed obliquely posteriorly, not reaching hind margin of pronotum; posterior margin almost straight, raised; no median carina but with impressed point on each side of median line; lateral portion behind eye very narrow, laminate; blunt, pale yellowish tubercles scattered on disc with row following anterior margin, and on outer margin of paranotal lobes; paranotal lobes (lateral view) broad, brown in outer portion, pale yellowish in inner portion and with large black marking behind lower portion of eye, and with posteroventral angle straight. Mesonotum variegated yellowish brown with longitudinal carinae obsolete but marked by dark brown lines; smooth, slightly depressed before scutellum; scutellum paler; nearly complete transverse carina along anterior margin. Tegulae brown.
</paragraph>
<paragraph id="432E36ED716C621FF9A0F97CFDA7FEC7" blockId="54.[188,1399,1710,2026]" lastBlockId="58.[189,1399,265,362]" lastPageId="58" lastPageNumber="59" pageId="54" pageNumber="55">
<smallCapsWord id="45C8A031716C6213F9A0F97CFED5F96A" baselines="1730,1730" box="[189,290,1711,1737]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="54" pageNumber="55">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A68716C6213F82FF97DFE34F965" box="[306,451,1710,1737]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="54" pageNumber="55">Figs 32A–C</figureCitation>
<figureCitation id="DBAA2A68716C6213F82FF97DFE34F965" box="[306,451,1710,1737]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508802" httpUri="https://zenodo.org/record/14508802/files/figure.png" pageId="54" pageNumber="55">Figs 32A–C</figureCitation>
,
<figureCitation id="DBAA2A68716C6213F8D2F97DFDDFF965" box="[463,552,1710,1737]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." pageId="54" pageNumber="55">35A–C</figureCitation>
<figureCitation id="DBAA2A68716C6213F8D2F97DFDDFF965" box="[463,552,1710,1737]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508808" httpUri="https://zenodo.org/record/14508808/files/figure.png" pageId="54" pageNumber="55">35A–C</figureCitation>
). Variegated yellowish brown (greener in live specimens); subcoriaceous with longitudinal veins yellow brown, elevated and with a dense reticulum of pale yellow veinlets; shape elongate and convex with sides broadly rounded, widest at basal ⅓, about 2.1 times as long as broad; narrowly rounded apically. Postclaval margin weakly rounded on distal half and weakly notched at apex of clavus. Clavus closed, reaching slightly over midlength of tegmen. Venation: ScP+R forking close to base after rather short common stem, ScP+RA and RP running more or less parallel to costal margin and not forked; MP forked rather close to base after common stem, resulting veins both forked again further; CuA simple, sinuate, more or less parallel to claval joint, then to sutural margin and merging with latter before apex of tegmen; Pcu and A
<subScript id="DF1534A8716C6213FB59F80FFDBAF847" attach="left" box="[580,589,2012,2026]" fontSize="6" pageId="54" pageNumber="55">1</subScript>
fused at apical ⅓ of clavus, resulting Pcu+A
@ -301,12 +302,12 @@ reaching apex of clavus.
<smallCapsWord id="45C8A0317162621DF9A0FEDAFF00FE8F" baselines="284,285" box="[189,247,265,291]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="56" pageNumber="57">HIND</smallCapsWord>
<smallCapsWord id="45C8A0317162621DF81FFEDDFEB1FE8C" baselines="284" box="[258,326,270,289]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="56" pageNumber="57">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A687162621DF847FEDAFE31FE8E" box="[346,454,265,291]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." pageId="56" pageNumber="57">Fig. 33E</figureCitation>
<figureCitation id="DBAA2A687162621DF847FEDAFE31FE8E" box="[346,454,265,291]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="56" pageNumber="57">Fig. 33E</figureCitation>
). Brown with large basicostal portion of lobe Sc-R-MP-CuA and base of lobe CuP-Pcu-A
<subScript id="DF1534A87162621DF859FEECFEBAFEE0" attach="left" box="[324,333,319,333]" fontSize="6" pageId="56" pageNumber="57">1</subScript>
yellow-brown; venation darker than corresponding background; wing broader than tegmen and deeply bilobed at CuP; costal margin moderately sinuate; CuP-Pcu-A1 lobe broadly rounded along postclaval margin and about 1.5 times as wide as Sc-R-MP-CuA lobe; both lobes approximately the same length, angularly rounded at apex; A2 lobe reduced and narrow. Venation: longitudinal veins ScP-R-MP-Cu well distinct; Pcu and A
</paragraph>
<caption id="17EE6665716D6212F9A0F85BFDC6F845" pageId="55" pageNumber="56" startId="55.[189,232,1928,1954]" targetBox="[221,1374,273,1866]" targetPageId="55">
<caption id="17EE6665716D6212F9A0F85BFDC6F845" ID-DOI="http://doi.org/10.5281/zenodo.14508802" ID-Zenodo-Dep="14508802" httpUri="https://zenodo.org/record/14508802/files/figure.png" pageId="55" pageNumber="56" startId="55.[189,232,1928,1954]" targetBox="[221,1374,273,1866]" targetPageId="55">
<paragraph id="432E36ED716D6212F9A0F85BFDC6F845" blockId="55.[189,1399,1928,2025]" pageId="55" pageNumber="56">
<emphasis id="71E5EAFF716D6212F9A0F85BFEE9F80F" bold="true" box="[189,286,1928,1954]" pageId="55" pageNumber="56">Fig. 32.</emphasis>
<taxonomicName id="84914D6E716D6212F834F85BFDBFF80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[297,584,1928,1954]" class="Insecta" family="Issidae" genus="Pusulissus" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Hemiptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="bachmaensis" status="sp. nov.">
@ -326,7 +327,7 @@ yellow-brown; venation darker than corresponding background; wing broader than t
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
<caption id="17EE66657162621DF9A0F8B7FEDFF845" pageId="56" pageNumber="57" startId="56.[189,232,1892,1918]" targetBox="[203,1384,453,1847]" targetPageId="56">
<caption id="17EE66657162621DF9A0F8B7FEDFF845" ID-DOI="http://doi.org/10.5281/zenodo.14508804" ID-Zenodo-Dep="14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="56" pageNumber="57" startId="56.[189,232,1892,1918]" targetBox="[203,1384,453,1847]" targetPageId="56">
<paragraph id="432E36ED7162621DF9A0F8B7FEDFF845" blockId="56.[189,1399,1892,2024]" pageId="56" pageNumber="57">
<emphasis id="71E5EAFF7162621DF9A0F8B7FEEDF8D2" bold="true" box="[189,282,1892,1919]" pageId="56" pageNumber="57">Fig. 33.</emphasis>
<taxonomicName id="84914D6E7162621DF83CF8B6FDCAF8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[289,573,1893,1919]" class="Insecta" family="Issidae" genus="Pusulissus" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Hemiptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="species" species="bachmaensis" status="sp. nov.">
@ -349,7 +350,7 @@ yellow-brown; venation darker than corresponding background; wing broader than t
. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods.
</paragraph>
</caption>
<caption id="17EE66657163621CF9A0F8B7FB49F845" pageId="57" pageNumber="58" startId="57.[189,232,1892,1918]" targetBox="[258,1331,285,1845]" targetPageId="57">
<caption id="17EE66657163621CF9A0F8B7FB49F845" ID-DOI="http://doi.org/10.5281/zenodo.14508806" ID-Zenodo-Dep="14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="57" pageNumber="58" startId="57.[189,232,1892,1918]" targetBox="[258,1331,285,1845]" targetPageId="57">
<paragraph id="432E36ED7163621CF9A0F8B7FB49F845" blockId="57.[187,1399,1892,2025]" pageId="57" pageNumber="58">
<emphasis id="71E5EAFF7163621CF9A0F8B7FED4F8D2" bold="true" box="[189,291,1892,1919]" pageId="57" pageNumber="58">Fig. 34.</emphasis>
<taxonomicName id="84914D6E7163621CF82EF8B6FDA0F8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[307,599,1893,1919]" class="Insecta" family="Issidae" genus="Pusulissus" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Hemiptera" pageId="57" pageNumber="58" phylum="Arthropoda" rank="species" species="bachmaensis" status="sp. nov.">
@ -390,13 +391,13 @@ yellow-brown; venation darker than corresponding background; wing broader than t
<paragraph id="432E36ED7160621FF9A0FE41FCDFFDB8" blockId="58.[187,1399,401,533]" pageId="58" pageNumber="59">
<smallCapsWord id="45C8A0317160621FF9A0FE41FF01FE07" baselines="421,421" box="[189,246,402,428]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="58" pageNumber="59">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A687160621FF818FE42FE65FE06" box="[261,402,401,428]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="58" pageNumber="59">Fig. 32A–E</figureCitation>
<figureCitation id="DBAA2A687160621FF818FE42FE65FE06" box="[261,402,401,428]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508802" httpUri="https://zenodo.org/record/14508802/files/figure.png" pageId="58" pageNumber="59">Fig. 32A–E</figureCitation>
). Moderately elongate and slender, yellowish brown femora darker than corresponding tibiae; apex of femora and base and pro- and mesotibiae yellowish; posterior margin of pro- and mesofemora with row of acute irregular minute teeth. Metatibiae with 2 lateral spines in distal half and 8 apical spines. Metatibiotarsal formula: (2) 8 / 8 / 2.
</paragraph>
<paragraph id="432E36ED7160621FF9A0FDEEFBFAFDFA" blockId="58.[189,1037,573,599]" box="[189,1037,573,599]" pageId="58" pageNumber="59">
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(
<figureCitation id="DBAA2A687160621FF823FDEEFE5EFDFA" box="[318,425,573,599]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="58" pageNumber="59">Fig. 32B</figureCitation>
<figureCitation id="DBAA2A687160621FF823FDEEFE5EFDFA" box="[318,425,573,599]" captionStart="Fig" captionStartId="55.[189,232,1928,1954]" captionTargetBox="[221,1374,273,1866]" captionTargetId="figure-11@55.[211,1377,265,1885]" captionTargetPageId="55" captionText="Fig. 32. Pusulissus bachmaensis sp. nov., dissected holotype, ♂ (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508802" httpUri="https://zenodo.org/record/14508802/files/figure.png" pageId="58" pageNumber="59">Fig. 32B</figureCitation>
). Yellowish with wide, black brown band in middle.
</paragraph>
<paragraph id="432E36ED7160621EF9A0FD53FC20FE75" blockId="58.[187,1400,639,1061]" lastBlockId="59.[189,1399,265,472]" lastPageId="59" lastPageNumber="60" pageId="58" pageNumber="59">
@ -405,15 +406,15 @@ yellow-brown; venation darker than corresponding background; wing broader than t
. Pygofer (
<emphasis id="71E5EAFF7160621FF8E2FD53FDE9FD34" box="[511,542,640,665]" italics="true" pageId="58" pageNumber="59">Py</emphasis>
–
<figureCitation id="DBAA2A687160621FFB2AFDACFD34FD34" box="[567,707,639,665]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="58" pageNumber="59">Fig. 34A–D</figureCitation>
<figureCitation id="DBAA2A687160621FFB2AFDACFD34FD34" box="[567,707,639,665]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="58" pageNumber="59">Fig. 34A–D</figureCitation>
) about 2.2 times as high as long in lateral view, with anterior margin sinuate and posterior margin more or less straight in middle portion, rounded to posterodorsal angle, and obliquely excavate in ventral ⅓; posterior margin dorsally deeply excavate (U-shaped excavation). Gonostyli (
<emphasis id="71E5EAFF7160621FF8C5FD38FE1AFCA8" box="[472,493,747,773]" italics="true" pageId="58" pageNumber="59">G</emphasis>
–
<figureCitation id="DBAA2A687160621FFB17FD38FD62FCA8" box="[522,661,747,773]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="58" pageNumber="59">Fig. 34A–C</figureCitation>
<figureCitation id="DBAA2A687160621FFB17FD38FD62FCA8" box="[522,661,747,773]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="58" pageNumber="59">Fig. 34A–C</figureCitation>
) (in lateral view) strongly elongate (without dorsal capitulum), projecting posteriorly in a distinct elongate lobe rounded apically, and concave; capitulum (
<emphasis id="71E5EAFF7160621FFC0EFCC3FAD8FC84" box="[1299,1327,784,809]" italics="true" pageId="58" pageNumber="59">ca</emphasis>
) with neck rather short and wide, less high than head of capitulum, with head massive, falcate in lateral view with apical hook directed cephalad, with strong tooth on inner margin directed mesad, and with lateral laminate projection in dorsal half with outer margin under it strongly concave in caudal view. Aedeagus (
<figureCitation id="DBAA2A687160621FF9D8FC4CFEA6FC14" box="[197,337,927,953]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="58" pageNumber="59">Fig. 34E–K</figureCitation>
<figureCitation id="DBAA2A687160621FF9D8FC4CFEA6FC14" box="[197,337,927,953]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="58" pageNumber="59">Fig. 34E–K</figureCitation>
) symmetrical, rather strongly curved dorsad (in lateral view), but with distal ¼ more or less straight; pair of elongate lateroventral processes (
<emphasis id="71E5EAFF7160621FFBE3FC10FCD5FC70" box="[766,802,963,989]" italics="true" pageId="58" pageNumber="59">lvp</emphasis>
) slightly inflated basally, then constricted, directed cephalad and moderately, evenly upcurved, remaining under ventral lobe of periandrium and reaching to basiventral lobes of periandrium; distal portion tapering and curved mesad; ventral lobe of periandrium (
@ -425,10 +426,10 @@ yellow-brown; venation darker than corresponding background; wing broader than t
) well developed. Anal tube (
<emphasis id="71E5EAFF7161621EF823FEA6FEA8FE22" box="[318,351,373,399]" italics="true" pageId="59" pageNumber="60">An</emphasis>
–
<figureCitation id="DBAA2A687161621EF864FEA6FDF2FE22" box="[377,517,373,399]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." pageId="59" pageNumber="60">Fig. 34A–D</figureCitation>
<figureCitation id="DBAA2A687161621EF864FEA6FDF2FE22" box="[377,517,373,399]" captionStart="Fig" captionStartId="57.[189,232,1892,1918]" captionTargetBox="[258,1331,285,1845]" captionTargetId="figure-11@57.[257,1331,265,1845]" captionTargetPageId="57" captionText="Fig. 34. Pusulissus bachmaensis sp. nov., holotype, ♂ (RBINS). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Caudal view. D. Dorsal view. E–L. Aedeagus. E. Posteroventral view. F. Left lateral view. G. Left laterodorsal view. H. Anterodorsal view. I. Dorsal view. J. Ventral view. K. Anteroventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508806" httpUri="https://zenodo.org/record/14508806/files/figure.png" pageId="59" pageNumber="60">Fig. 34A–D</figureCitation>
) dorsoventrally flattened, in dorsal view widely spatulate with distal portion nearly circular, nearly 1.3 times as long in midline as wide, with apical margin shortly truncate in middle, without distinct lateral flaps; anal opening at about basal ⅓.
</paragraph>
<caption id="17EE66657160621FF9A0F878FC8CF845" pageId="58" pageNumber="59" startId="58.[189,232,1963,1989]" targetBox="[189,1398,1110,1917]" targetPageId="58">
<caption id="17EE66657160621FF9A0F878FC8CF845" ID-DOI="http://doi.org/10.5281/zenodo.14508808" ID-Zenodo-Dep="14508808" httpUri="https://zenodo.org/record/14508808/files/figure.png" pageId="58" pageNumber="59" startId="58.[189,232,1963,1989]" targetBox="[189,1398,1110,1917]" targetPageId="58">
<paragraph id="432E36ED7160621FF9A0F878FC8CF845" blockId="58.[189,1398,1963,2025]" pageId="58" pageNumber="59">
<emphasis id="71E5EAFF7160621FF9A0F878FEECF868" bold="true" box="[189,283,1963,1989]" pageId="58" pageNumber="59">Fig. 35.</emphasis>
<taxonomicName id="84914D6E7160621FF83FF878FDCAF868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[290,573,1963,1989]" class="Insecta" family="Issidae" genus="Pusulissus" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Hemiptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="bachmaensis" status="sp. nov.">
@ -446,7 +447,7 @@ yellow-brown; venation darker than corresponding background; wing broader than t
. Hind margin of sternum VII (
<emphasis id="71E5EAFF7161621EFA1CFDD3FCB4FDB4" box="[769,835,512,538]" italics="true" pageId="59" pageNumber="60">St VII</emphasis>
–
<figureCitation id="DBAA2A687161621EFA42FE2CFC34FDB4" box="[863,963,511,537]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." pageId="59" pageNumber="60">Fig. 33F</figureCitation>
<figureCitation id="DBAA2A687161621EFA42FE2CFC34FDB4" box="[863,963,511,537]" captionStart="Fig" captionStartId="56.[189,232,1892,1918]" captionTargetBox="[203,1384,453,1847]" captionTargetId="figure-63@56.[203,1384,442,1850]" captionTargetPageId="56" captionText="Fig. 33. Pusulissus bachmaensis sp. nov. A–E. Holotype ♂ (RBINS). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (VNMN), terminalia, ventral view. Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508804" httpUri="https://zenodo.org/record/14508804/files/figure.png" pageId="59" pageNumber="60">Fig. 33F</figureCitation>
) concave, with middle portion nearly straight; small digitiform process directed posterad, derived from middle of sternum VII and slightly surpassing posterior margin.
</paragraph>
</subSubSection>
@ -472,9 +473,9 @@ in
, both in montane evergreen tropical forest, at about
<quantity id="84699B087161621EFA49FD0EFC01FD5A" box="[852,1014,733,759]" metricMagnitude="3" metricUnit="m" metricValue="1.35" metricValueMax="1.4" metricValueMin="1.3" pageId="59" pageNumber="60" unit="m" value="1350.0" valueMax="1400.0" valueMin="1300.0">1300–1400 m</quantity>
a.s.l., near the summit (
<figureCitation id="DBAA2A687161621EFC16FD0EFA9FFD5A" box="[1291,1384,733,759]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="59" pageNumber="60">Fig. 1D</figureCitation>
<figureCitation id="DBAA2A687161621EFC16FD0EFA9FFD5A" box="[1291,1384,733,759]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="59" pageNumber="60">Fig. 1D</figureCitation>
). The specimens were collected on tree trunks densely covered in lichens and mosses (the female was photographed in a cage –
<figureCitation id="DBAA2A687161621EF8F5FCF6FDC9FCED" box="[488,574,805,832]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." pageId="59" pageNumber="60">Fig. 35</figureCitation>
<figureCitation id="DBAA2A687161621EF8F5FCF6FDC9FCED" box="[488,574,805,832]" captionStart="Fig" captionStartId="58.[189,232,1963,1989]" captionTargetBox="[189,1398,1110,1917]" captionTargetId="figure-331@58.[189,1398,1110,1917]" captionTargetPageId="58" captionText="Fig. 35. Pusulissus bachmaensis sp. nov., paratype, ♀ (VNMN) from Bach Ma National Park, summit, 21 May 2023. A–D. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508808" httpUri="https://zenodo.org/record/14508808/files/figure.png" pageId="59" pageNumber="60">Fig. 35</figureCitation>
). When disturbed, they tend to walk away and hide on the opposite side of the trunk or branch, and jump away as a last solution.
</paragraph>
</subSubSection>
@ -487,7 +488,7 @@ a.s.l., near the summit (
<paragraph id="432E36ED7161621EF9A0FC69FB9DFC79" blockId="59.[189,1130,954,983]" box="[189,1130,954,983]" pageId="59" pageNumber="60">
<collectingCountry id="3B86767D7161621EF9A0FC69FED2FC79" box="[189,293,954,980]" name="Vietnam" pageId="59" pageNumber="60">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, summit of Bach Ma National Park (
<figureCitation id="DBAA2A687161621EFD09FC69FBAEFC79" box="[1044,1113,954,980]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="59" pageNumber="60">Fig. 1</figureCitation>
<figureCitation id="DBAA2A687161621EFD09FC69FBAEFC79" box="[1044,1113,954,980]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="59" pageNumber="60">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>

127
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@ -1,52 +1,53 @@
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<mods:affiliation id="585CCC95CDF9F3ACBD5B070C0DECF528">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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<mods:affiliation id="53A90635BE736D2FC70EE2E7F95BCF54">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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@ -67,9 +68,9 @@ urn:lsid:zoobank.org:act:
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<paragraph id="432E36ED7173620CFBD4F954FC9DF90F" blockId="41.[368,1219,1606,1698]" box="[713,874,1671,1698]" pageId="41" pageNumber="42">
<figureCitation id="DBAA2A687173620CFBD4F954FCE5F90F" box="[713,786,1671,1698]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="41" pageNumber="42">Figs 1</figureCitation>
<figureCitation id="DBAA2A687173620CFBD4F954FCE5F90F" box="[713,786,1671,1698]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="41" pageNumber="42">Figs 1</figureCitation>
,
<figureCitation id="DBAA2A687173620CFA3CF95BFC9DF90F" box="[801,874,1672,1698]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="43.[189,232,1928,1954]" captionStartId-1="44.[189,232,1892,1918]" captionStartId-2="45.[189,232,1892,1918]" captionStartId-3="46.[189,232,1963,1989]" captionTargetBox-0="[252,1336,270,1884]" captionTargetBox-1="[255,1328,272,1845]" captionTargetBox-2="[258,1331,282,1845]" captionTargetBox-3="[189,1398,1062,1920]" captionTargetId-0="figure-11@43.[243,1344,265,1885]" captionTargetId-1="figure-16@44.[255,1333,265,1851]" captionTargetId-2="figure-11@45.[257,1331,265,1845]" captionTargetId-3="figure-293@46.[189,1398,1062,1920]" captionTargetPageId-0="43" captionTargetPageId-1="44" captionTargetPageId-2="45" captionTargetPageId-3="46" captionText-0="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." captionText-3="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="41" pageNumber="42">24–27</figureCitation>
<figureCitation id="DBAA2A687173620CFA3CF95BFC9DF90F" box="[801,874,1672,1698]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="43.[189,232,1928,1954]" captionStartId-1="44.[189,232,1892,1918]" captionStartId-2="45.[189,232,1892,1918]" captionStartId-3="46.[189,232,1963,1989]" captionTargetBox-0="[252,1336,270,1884]" captionTargetBox-1="[255,1328,272,1845]" captionTargetBox-2="[258,1331,282,1845]" captionTargetBox-3="[189,1398,1062,1920]" captionTargetId-0="figure-11@43.[243,1344,265,1885]" captionTargetId-1="figure-16@44.[255,1333,265,1851]" captionTargetId-2="figure-11@45.[257,1331,265,1845]" captionTargetId-3="figure-293@46.[189,1398,1062,1920]" captionTargetPageId-0="43" captionTargetPageId-1="44" captionTargetPageId-2="45" captionTargetPageId-3="46" captionText-0="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." captionText-3="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi-0="http://doi.org/10.5281/zenodo.14508786" figureDoi-1="http://doi.org/10.5281/zenodo.14508788" figureDoi-2="http://doi.org/10.5281/zenodo.14508790" figureDoi-3="http://doi.org/10.5281/zenodo.14508792" httpUri-0="https://zenodo.org/record/14508786/files/figure.png" httpUri-1="https://zenodo.org/record/14508788/files/figure.png" httpUri-2="https://zenodo.org/record/14508790/files/figure.png" httpUri-3="https://zenodo.org/record/14508792/files/figure.png" pageId="41" pageNumber="42">24–27</figureCitation>
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@ -87,14 +88,14 @@ can be recognized by:
</paragraph>
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(1) the mostly bright green colour of the insect (
<figureCitation id="DBAA2A687173620CFA1EF8C0FC61F880" box="[771,918,1811,1837]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="41" pageNumber="42">Figs 24A–C</figureCitation>
<figureCitation id="DBAA2A687173620CFA1EF8C0FC61F880" box="[771,918,1811,1837]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508786" httpUri="https://zenodo.org/record/14508786/files/figure.png" pageId="41" pageNumber="42">Figs 24A–C</figureCitation>
,
<figureCitation id="DBAA2A687173620CFABFF8C0FC0FF880" box="[930,1016,1811,1837]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="41" pageNumber="42">27C–E</figureCitation>
<figureCitation id="DBAA2A687173620CFABFF8C0FC0FF880" box="[930,1016,1811,1837]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="41" pageNumber="42">27C–E</figureCitation>
);
</paragraph>
<paragraph id="432E36ED7173620CF9A0F8E0FC80F8E3" blockId="41.[189,1030,1778,1870]" box="[189,887,1843,1870]" pageId="41" pageNumber="42">
(2) the yellow-brown colour of the hind wings (
<figureCitation id="DBAA2A687173620CFBE2F8E0FC9DF8E3" box="[767,874,1843,1870]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." pageId="41" pageNumber="42">Fig. 25E</figureCitation>
<figureCitation id="DBAA2A687173620CFBE2F8E0FC9DF8E3" box="[767,874,1843,1870]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508788" httpUri="https://zenodo.org/record/14508788/files/figure.png" pageId="41" pageNumber="42">Fig. 25E</figureCitation>
).
</paragraph>
<paragraph id="432E36ED7173620CF9A0F8A7FE3EF823" blockId="41.[189,457,1908,1934]" box="[189,457,1908,1934]" pageId="41" pageNumber="42">
@ -159,7 +160,7 @@ series in Bach Ma National Park during our 2023 expedition.
•
<specimenCount id="5597FD647170620FF842FCD0FE82FCB0" box="[351,373,771,797]" count="1" pageId="42" pageNumber="43" type="male">♂</specimenCount>
(dissected –
<figureCitation id="DBAA2A687170620FFB0FFCD1FD6BFCB1" box="[530,668,770,796]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="43.[189,232,1928,1954]" captionStartId-1="44.[189,232,1892,1918]" captionStartId-2="45.[189,232,1892,1918]" captionTargetBox-0="[252,1336,270,1884]" captionTargetBox-1="[255,1328,272,1845]" captionTargetBox-2="[258,1331,282,1845]" captionTargetId-0="figure-11@43.[243,1344,265,1885]" captionTargetId-1="figure-16@44.[255,1333,265,1851]" captionTargetId-2="figure-11@45.[257,1331,265,1845]" captionTargetPageId-0="43" captionTargetPageId-1="44" captionTargetPageId-2="45" captionText-0="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="42" pageNumber="43">Figs 24–26</figureCitation>
<figureCitation id="DBAA2A687170620FFB0FFCD1FD6BFCB1" box="[530,668,770,796]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="43.[189,232,1928,1954]" captionStartId-1="44.[189,232,1892,1918]" captionStartId-2="45.[189,232,1892,1918]" captionTargetBox-0="[252,1336,270,1884]" captionTargetBox-1="[255,1328,272,1845]" captionTargetBox-2="[258,1331,282,1845]" captionTargetId-0="figure-11@43.[243,1344,265,1885]" captionTargetId-1="figure-16@44.[255,1333,265,1851]" captionTargetId-2="figure-11@45.[257,1331,265,1845]" captionTargetPageId-0="43" captionTargetPageId-1="44" captionTargetPageId-2="45" captionText-0="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." captionText-2="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508786" figureDoi-1="http://doi.org/10.5281/zenodo.14508788" figureDoi-2="http://doi.org/10.5281/zenodo.14508790" httpUri-0="https://zenodo.org/record/14508786/files/figure.png" httpUri-1="https://zenodo.org/record/14508788/files/figure.png" httpUri-2="https://zenodo.org/record/14508790/files/figure.png" pageId="42" pageNumber="43">Figs 24–26</figureCitation>
);
<location id="464E60367170620FFBADFCD1FD27FCB1" LSID="urn:lsid:plazi:treatment:CB3887FB7173620AFB48F995FC1CFAC4:464E60367170620FFBADFCD1FD27FCB1" box="[688,720,770,796]" country="Vietnam" county="YesHue Eco" latitude="16.218056" longLatPrecision="21" longitude="107.72417" municipality="Bach Ma National Park" name="Th" pageId="42" pageNumber="43">Th</location>
ừa Thiên-Huế
@ -239,26 +240,26 @@ leg.;
<paragraph id="432E36ED7170620FF9A0FB62FC40F986" blockId="42.[187,1401,1201,1579]" pageId="42" pageNumber="43">
<smallCapsWord id="45C8A0317170620FF9A0FB62FF0BFB64" baselines="1220,1220" box="[189,252,1201,1227]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="42" pageNumber="43">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A687170620FF813FB62FE68FB61" box="[270,415,1201,1228]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." pageId="42" pageNumber="43">Fig. 25A–D</figureCitation>
<figureCitation id="DBAA2A687170620FF813FB62FE68FB61" box="[270,415,1201,1228]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508788" httpUri="https://zenodo.org/record/14508788/files/figure.png" pageId="42" pageNumber="43">Fig. 25A–D</figureCitation>
). Strongly elongated anteriorly in cephalic process; vertex olivaceous green, usually getting slightly darker towards apex; distinctly longer in midline than broad before eyes (nearly 2.9 times), widening from base to anterior margin of eyes, tapering beyond eyes to midlength, then with sides parallel to rounded apex, median carina weakly distinct, lateral carinae distinct; posterior margin moderately incurved. Side of head brown, darker than vertex anteriorly on cephalic process, yellowish under eye to clypeal suture. Frons yellowish in basal portion then turning dark brown from middle of eyes towards apex, with yellow area extended anteriorly in middle; lateral carinae reaching apex, median carina distinct in distal half; elongate, widest slightly anterior to eyes, roundly tapering towards clypeal suture; clypeal suture rounded. Clypeus black-brown with base shortly yellow, weakly elevated medially in distal portion. Labium brown with last segment longer than broad, and shorter than penultimate. Antennae dark brown; scape short, ring-shaped; pedicel bulbous.
</paragraph>
<paragraph id="432E36ED7170620FF9A0F980FDCBF8CF" blockId="42.[189,1399,1617,1890]" pageId="42" pageNumber="43">
<smallCapsWord id="45C8A0317170620FF9A0F980FEEEF9C6" baselines="1638,1638" box="[189,281,1619,1645]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="42" pageNumber="43">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A687170620FF834F981FE4EF9C0" box="[297,441,1618,1645]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." pageId="42" pageNumber="43">Fig. 25A–D</figureCitation>
<figureCitation id="DBAA2A687170620FF834F981FE4EF9C0" box="[297,441,1618,1645]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508788" httpUri="https://zenodo.org/record/14508788/files/figure.png" pageId="42" pageNumber="43">Fig. 25A–D</figureCitation>
). Pronotum yellowish brown; about ⅔ of length of mesonotum in midline; anterior margin weakly carinate, sinuate and moderately protruding anteriorly between eyes, with peridiscal carinae weakly distinct; posterior margin nearly straight; no median carina but with impressed point on each side of median line; weak, blunt tubercles along anterior and posterior margins; paranotal lobes (lateral view) moderately broad, turning dark brown in lower portion, under level of eye and with posteroventral angle rather acutely rounded. Mesonotum yellowish brown with longitudinal carinae obsolete; two black-brown impressed points on disc and smaller black-brown point near anterolateral angle. Tegulae yellowish brown.
</paragraph>
<paragraph id="432E36ED7170620BF9A0F858FDA1FDB7" blockId="42.[189,1401,1930,1991]" lastBlockId="46.[189,1401,263,538]" lastPageId="46" lastPageNumber="47" pageId="42" pageNumber="43">
<smallCapsWord id="45C8A0317170620FF9A0F858FED5F80E" baselines="1950,1950" box="[189,290,1931,1957]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="42" pageNumber="43">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A687170620FF82BF859FE3CF809" box="[310,459,1930,1956]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="42" pageNumber="43">Figs 24A–C</figureCitation>
<figureCitation id="DBAA2A687170620FF82BF859FE3CF809" box="[310,459,1930,1956]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508786" httpUri="https://zenodo.org/record/14508786/files/figure.png" pageId="42" pageNumber="43">Figs 24A–C</figureCitation>
,
<figureCitation id="DBAA2A687170620FF8C4F859FDDBF808" box="[473,556,1930,1957]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="42" pageNumber="43">27C–E</figureCitation>
<figureCitation id="DBAA2A687170620FF8C4F859FDDBF808" box="[473,556,1930,1957]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="42" pageNumber="43">27C–E</figureCitation>
). Green (tending to fade to yellow in collection specimens), narrowly washed with brown along apical margin; subcoriaceous with longitudinal veins distinct and with dense reticulum of veinlets; shape elongate and convex with sides broadly rounded, widest at basal ⅓, about 2.7 times as long as broad; narrowly rounded apically. Postclaval margin nearly straight, very weakly rounded on distal half, and notched at apex of clavus. Clavus closed, reaching about ⅔ of tegmen. Venation: ScP+R forking close to base after short common stem, ScP+RA and RP running more or less parallel to costal margin; MP forked slightly before basal ⅓, resulting veins running more or less parallel; CuA weakly diverging from claval joint and forked before apex of latter; Pcu and A 1 fused at distal ¼ of clavus, resulting Pcu+A
<subScript id="DF1534A87174620BFB63FE3CFD70FE50" attach="left" box="[638,647,495,509]" fontSize="6" pageId="46" pageNumber="47">1</subScript>
reaching claval joint before apex of clavus; dense reticulum of veinlets, espcially beyond basal ⅓.
</paragraph>
<caption id="17EE66657171620EF9A0F85BFDC6F845" pageId="43" pageNumber="44" startId="43.[189,232,1928,1954]" targetBox="[252,1336,270,1884]" targetPageId="43">
<caption id="17EE66657171620EF9A0F85BFDC6F845" ID-DOI="http://doi.org/10.5281/zenodo.14508786" ID-Zenodo-Dep="14508786" httpUri="https://zenodo.org/record/14508786/files/figure.png" pageId="43" pageNumber="44" startId="43.[189,232,1928,1954]" targetBox="[252,1336,270,1884]" targetPageId="43">
<paragraph id="432E36ED7171620EF9A0F85BFDC6F845" blockId="43.[189,1399,1928,2025]" pageId="43" pageNumber="44">
<emphasis id="71E5EAFF7171620EF9A0F85BFEE9F80F" bold="true" box="[189,286,1928,1954]" pageId="43" pageNumber="44">Fig. 24.</emphasis>
<taxonomicName id="84914D6E7171620EF835F85BFDBDF80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[296,586,1928,1954]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="lindae" status="sp. nov.">
@ -278,7 +279,7 @@ reaching claval joint before apex of clavus; dense reticulum of veinlets, espcia
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
<caption id="17EE666571766209F9A0F8B7FEDFF845" pageId="44" pageNumber="45" startId="44.[189,232,1892,1918]" targetBox="[255,1328,272,1845]" targetPageId="44">
<caption id="17EE666571766209F9A0F8B7FEDFF845" ID-DOI="http://doi.org/10.5281/zenodo.14508788" ID-Zenodo-Dep="14508788" httpUri="https://zenodo.org/record/14508788/files/figure.png" pageId="44" pageNumber="45" startId="44.[189,232,1892,1918]" targetBox="[255,1328,272,1845]" targetPageId="44">
<paragraph id="432E36ED71766209F9A0F8B7FEDFF845" blockId="44.[188,1399,1892,2024]" pageId="44" pageNumber="45">
<emphasis id="71E5EAFF71766209F9A0F8B7FEEDF8D2" bold="true" box="[189,282,1892,1919]" pageId="44" pageNumber="45">Fig. 25.</emphasis>
<taxonomicName id="84914D6E71766209F83DF8B6FDC8F8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[288,575,1893,1919]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="lindae" status="sp. nov.">
@ -301,7 +302,7 @@ reaching claval joint before apex of clavus; dense reticulum of veinlets, espcia
. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods.
</paragraph>
</caption>
<caption id="17EE666571776208F9A0F8B7FA80F845" pageId="45" pageNumber="46" startId="45.[189,232,1892,1918]" targetBox="[258,1331,282,1845]" targetPageId="45">
<caption id="17EE666571776208F9A0F8B7FA80F845" ID-DOI="http://doi.org/10.5281/zenodo.14508790" ID-Zenodo-Dep="14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="45" pageNumber="46" startId="45.[189,232,1892,1918]" targetBox="[258,1331,282,1845]" targetPageId="45">
<paragraph id="432E36ED71776208F9A0F8B7FA80F845" blockId="45.[187,1399,1892,2025]" pageId="45" pageNumber="46">
<emphasis id="71E5EAFF71776208F9A0F8B7FEEEF8D2" bold="true" box="[189,281,1892,1919]" pageId="45" pageNumber="46">Fig. 26.</emphasis>
<taxonomicName id="84914D6E71776208F802F8B6FDCAF8D2" authorityName="Constant &amp; Pham" authorityYear="2024" box="[287,573,1893,1919]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="lindae" status="sp. nov.">
@ -343,7 +344,7 @@ reaching claval joint before apex of clavus; dense reticulum of veinlets, espcia
<smallCapsWord id="45C8A0317174620BF9A0FD92FF00FDF7" baselines="596,597" box="[189,247,577,603]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="46" pageNumber="47">HIND</smallCapsWord>
<smallCapsWord id="45C8A0317174620BF9E2FD95FEB4FDF4" baselines="596" box="[255,323,582,601]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="46" pageNumber="47">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A687174620BF848FD92FE37FDF6" box="[341,448,577,603]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="46" pageNumber="47">Fig. 26E</figureCitation>
<figureCitation id="DBAA2A687174620BF848FD92FE37FDF6" box="[341,448,577,603]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="46" pageNumber="47">Fig. 26E</figureCitation>
). Yellow-brown turning to dark brown along margin in distal portion of lobe Sc-R-MP-CuA and narrowly along margin in distal portion of lobe CuP-Pcu-A
<subScript id="DF1534A87174620BFD23FDA4FBB0FD28" attach="left" box="[1086,1095,631,645]" fontSize="6" pageId="46" pageNumber="47">1</subScript>
, A
@ -359,10 +360,10 @@ lobe moderately reduced and narrow. Venation: longitudinal veins ScP-R-MP-Cu wel
<paragraph id="432E36ED7174620BF9A0FC85FA9FFC1B" blockId="46.[189,1399,853,950]" pageId="46" pageNumber="47">
<smallCapsWord id="45C8A0317174620BF9A0FC85FF01FCC3" baselines="873,873" box="[189,246,854,880]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="46" pageNumber="47">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A687174620BF817FC86FE6CFCDD" box="[266,411,853,880]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="46" pageNumber="47">Fig. 24A–E</figureCitation>
<figureCitation id="DBAA2A687174620BF817FC86FE6CFCDD" box="[266,411,853,880]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508786" httpUri="https://zenodo.org/record/14508786/files/figure.png" pageId="46" pageNumber="47">Fig. 24A–E</figureCitation>
). Moderately elongate, slender; femora brown, darker in basal portion, wider than corresponding tibiae; tibiae paler than corresponding femora, yellow-brown; pro- and meso tarsi black; metatibiae with 2 lateral spines in distal half and 7 apical spines. Metatibiotarsal formula: (2) 7 / 9 / 2.
</paragraph>
<caption id="17EE66657174620BF9A0F878FD1DF845" pageId="46" pageNumber="47" startId="46.[189,232,1963,1989]" targetBox="[189,1398,1062,1920]" targetPageId="46">
<caption id="17EE66657174620BF9A0F878FD1DF845" ID-DOI="http://doi.org/10.5281/zenodo.14508792" ID-Zenodo-Dep="14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="46" pageNumber="47" startId="46.[189,232,1963,1989]" targetBox="[189,1398,1062,1920]" targetPageId="46">
<paragraph id="432E36ED7174620BF9A0F878FD1DF845" blockId="46.[189,1399,1963,2025]" pageId="46" pageNumber="47">
<emphasis id="71E5EAFF7174620BF9A0F878FEE8F868" bold="true" box="[189,287,1963,1989]" pageId="46" pageNumber="47">Fig. 27.</emphasis>
<taxonomicName id="84914D6E7174620BF837F878FDBAF868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[298,589,1963,1989]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="46" pageNumber="47" phylum="Arthropoda" rank="species" species="lindae" status="sp. nov.">
@ -381,7 +382,7 @@ in Bach Ma National Park, Yes Hue Eco, 17 May 2023.
<paragraph id="432E36ED7175620AF9A0FEDAFB59FE8E" blockId="47.[189,1198,265,291]" box="[189,1198,265,291]" pageId="47" pageNumber="48">
<smallCapsWord id="45C8A0317175620AF9A0FEDAFEDBFE8C" baselines="284,284" box="[189,300,265,291]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Abdomen" pageId="47" pageNumber="48">ABDOMEN</smallCapsWord>
(
<figureCitation id="DBAA2A687175620AF823FEDAFE5EFE8E" box="[318,425,265,291]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="47" pageNumber="48">Fig. 24B</figureCitation>
<figureCitation id="DBAA2A687175620AF823FEDAFE5EFE8E" box="[318,425,265,291]" captionStart="Fig" captionStartId="43.[189,232,1928,1954]" captionTargetBox="[252,1336,270,1884]" captionTargetId="figure-11@43.[243,1344,265,1885]" captionTargetPageId="43" captionText="Fig. 24. Pseudochoutagus lindae sp. nov., dissected holotype, ♂ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508786" httpUri="https://zenodo.org/record/14508786/files/figure.png" pageId="47" pageNumber="48">Fig. 24B</figureCitation>
). Yellowish brown, middle portion of basal segments dark brown.
</paragraph>
<paragraph id="432E36ED7175620AF9A0FE98FB44FC73" blockId="47.[189,1401,330,991]" pageId="47" pageNumber="48">
@ -390,15 +391,15 @@ in Bach Ma National Park, Yes Hue Eco, 17 May 2023.
. Pygofer (
<emphasis id="71E5EAFF7175620AFB0DFE98FDD8FEC9" box="[528,559,331,356]" italics="true" pageId="47" pageNumber="48">Py</emphasis>
–
<figureCitation id="DBAA2A687175620AFB4EFE99FD12FEC9" box="[595,741,330,356]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="47" pageNumber="48">Fig. 26A–D</figureCitation>
<figureCitation id="DBAA2A687175620AFB4EFE99FD12FEC9" box="[595,741,330,356]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="47" pageNumber="48">Fig. 26A–D</figureCitation>
) about 2.3 times as high as long in lateral view, with anterior and posterior margins subparallel, sinuate; posterior margin with posterodorsal angle nearly right and dorsal margin straight; in dorsal view, posterior margin very deeply and abruptly excavate. Gonostyli (
<emphasis id="71E5EAFF7175620AF85EFE67FEAFFE63" box="[323,344,436,462]" italics="true" pageId="47" pageNumber="48">G</emphasis>
–
<figureCitation id="DBAA2A687175620AF865FE67FDDEFE63" box="[376,553,436,462]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="47" pageNumber="48">Fig. 26A–B, D</figureCitation>
<figureCitation id="DBAA2A687175620AF865FE67FDDEFE63" box="[376,553,436,462]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="47" pageNumber="48">Fig. 26A–B, D</figureCitation>
) (in lateral view) longer than high (without dorsal capitulum), tapering posteriorly in a distinct lobe narrowly rounded apically, and concave; capitulum (
<emphasis id="71E5EAFF7175620AFD97FE0BFB51FE5C" box="[1162,1190,472,497]" italics="true" pageId="47" pageNumber="48">ca</emphasis>
) with wide neck, curved laterad and directed posterodorsad, with distal portion obliquely flattened, and with outer margin under lateral laminate projection, strongly concave in caudal view. Aedeagus (
<figureCitation id="DBAA2A687175620AFD66FDCDFAE4FD95" box="[1147,1299,541,568]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="47" pageNumber="48">Fig. 26 E–L</figureCitation>
<figureCitation id="DBAA2A687175620AFD66FDCDFAE4FD95" box="[1147,1299,541,568]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="47" pageNumber="48">Fig. 26 E–L</figureCitation>
) mostly symmetrical, except for lateroventral processes (
<emphasis id="71E5EAFF7175620AFA18FD92FCDEFDF6" box="[773,809,577,603]" italics="true" pageId="47" pageNumber="48">lvp</emphasis>
), rather strongly curved dorsad (in lateral view), longer and wider than ventral lobe of periandrium, and with pair of posterolateral pointed processes directed cephalad; pair of elongate lateroventral processes (
@ -414,7 +415,7 @@ in Bach Ma National Park, Yes Hue Eco, 17 May 2023.
). Anal tube (
<emphasis id="71E5EAFF7175620AFB7FFC88FD74FCD8" box="[610,643,859,885]" italics="true" pageId="47" pageNumber="48">An</emphasis>
–
<figureCitation id="DBAA2A687175620AFBBDFC89FCD9FCD9" box="[672,814,858,884]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." pageId="47" pageNumber="48">Fig. 26A–D</figureCitation>
<figureCitation id="DBAA2A687175620AFBBDFC89FCD9FCD9" box="[672,814,858,884]" captionStart="Fig" captionStartId="45.[189,232,1892,1918]" captionTargetBox="[258,1331,282,1845]" captionTargetId="figure-11@45.[257,1331,265,1845]" captionTargetPageId="45" captionText="Fig. 26. Pseudochoutagus lindae sp. nov., holotype, ♂ (VNMN). A–D. Pygofer, anal tube and gonostyli. A. Left lateral view. B. Posterolateral view. C. Dorsal view. D. Caudal view. E–L. Aedeagus. E. Left lateral view. F. Left laterodorsal view. G. Left lateroventral view. H. Dorsal view. I. Anterodorsal view. J. Ventral view.K. Posteroventral view.L. Anteroventral view.Abbreviations: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508790" httpUri="https://zenodo.org/record/14508790/files/figure.png" pageId="47" pageNumber="48">Fig. 26A–D</figureCitation>
) dorsoventrally flattened, in dorsal view spatulate with apical margin roundly emarginate; basal portion narrow, widening to strong sinuation at level of anal opening; distal portion weakly tapering to lateroposterior rounded angles; in lateral view, strongly curved near base, then gently curved posteroventrad; anal opening at about midlength.
</paragraph>
<paragraph id="432E36ED7175620AF9A0FBD5FD55FBEE" blockId="47.[189,1398,1029,1091]" pageId="47" pageNumber="48">
@ -423,7 +424,7 @@ in Bach Ma National Park, Yes Hue Eco, 17 May 2023.
. Hind margin of sternum VII (
<emphasis id="71E5EAFF7175620AFA06FBD5FC96FBB2" box="[795,865,1030,1056]" italics="true" pageId="47" pageNumber="48">St VII</emphasis>
–
<figureCitation id="DBAA2A687175620AFA99FBD6FC1BFB8D" box="[900,1004,1029,1056]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." pageId="47" pageNumber="48">Fig. 25F</figureCitation>
<figureCitation id="DBAA2A687175620AFA99FBD6FC1BFB8D" box="[900,1004,1029,1056]" captionStart="Fig" captionStartId="44.[189,232,1892,1918]" captionTargetBox="[255,1328,272,1845]" captionTargetId="figure-16@44.[255,1333,265,1851]" captionTargetPageId="44" captionText="Fig. 25. Pseudochoutagus lindae sp. nov. A–E. Holotype ♂ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing. F. Paratype ♀ (RBINS), terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508788" httpUri="https://zenodo.org/record/14508788/files/figure.png" pageId="47" pageNumber="48">Fig. 25F</figureCitation>
) with massive, widely spatulate, apically weakly concave median process.
</paragraph>
</subSubSection>
@ -439,13 +440,13 @@ in Bach Ma National Park, Yes Hue Eco, 17 May 2023.
</taxonomicName>
<taxonomicNameLabel id="6AD657847175620AF8F7FB4AFDB0FB1E" box="[490,583,1177,1203]" pageId="47" pageNumber="48" rank="species">sp. nov.</taxonomicNameLabel>
was collected in May in a marshy microhabitat (
<figureCitation id="DBAA2A687175620AFD83FB4AFAFCFB1E" box="[1182,1291,1177,1203]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="47" pageNumber="48">Fig. 27A</figureCitation>
<figureCitation id="DBAA2A687175620AFD83FB4AFAFCFB1E" box="[1182,1291,1177,1203]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="47" pageNumber="48">Fig. 27A</figureCitation>
) along a stream by sweeping a fern (
<figureCitation id="DBAA2A687175620AFB1CFB6FFD9BFB7B" box="[513,620,1212,1238]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="47" pageNumber="48">Fig. 27B</figureCitation>
<figureCitation id="DBAA2A687175620AFB1CFB6FFD9BFB7B" box="[513,620,1212,1238]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="47" pageNumber="48">Fig. 27B</figureCitation>
), in moist evergreen tropical forest at about
<quantity id="84699B087175620AFD6EFB6FFB4AFB7B" box="[1139,1213,1212,1239]" metricMagnitude="2" metricUnit="m" metricValue="2.5" pageId="47" pageNumber="48" unit="m" value="250.0">250 m</quantity>
in altitude. The green colour of adult specimens (
<figureCitation id="DBAA2A687175620AFB59FB0CFD39FB54" box="[580,718,1247,1273]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="47" pageNumber="48">Fig. 27C–E</figureCitation>
<figureCitation id="DBAA2A687175620AFB59FB0CFD39FB54" box="[580,718,1247,1273]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="47" pageNumber="48">Fig. 27C–E</figureCitation>
) probably helps their camouflage on their host plants.
</paragraph>
</subSubSection>
@ -458,7 +459,7 @@ in altitude. The green colour of adult specimens (
<paragraph id="432E36ED7175620AF9A0FA9CFC1CFAC4" blockId="47.[189,1003,1359,1388]" box="[189,1003,1359,1388]" pageId="47" pageNumber="48">
<collectingCountry id="3B86767D7175620AF9A0FA9CFED2FAC4" box="[189,293,1359,1385]" name="Vietnam" pageId="47" pageNumber="48">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, Bach Ma National Park (
<figureCitation id="DBAA2A687175620AFA88FA9CFC2DFAC4" box="[917,986,1359,1385]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="47" pageNumber="48">Fig. 1</figureCitation>
<figureCitation id="DBAA2A687175620AFA88FA9CFC2DFAC4" box="[917,986,1359,1385]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="47" pageNumber="48">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>

117
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@ -1,52 +1,53 @@
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<mods:affiliation id="F19BA7F0A767FBC20B60CDACFF722BB5">Royal Belgian Institute of Natural Sciences, O. D. Phylogeny and Taxonomy, Entomology, Vautier Street 29, B- 1000 Brussels, Belgium.</mods:affiliation>
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<mods:namePart id="228ED308273491683FE309B3309613C5">Pham, Hong Thai</mods:namePart>
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<mods:affiliation id="4030DF0488200C3E71D89CCF8B36ADFF">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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<mods:affiliation id="B92187041DC34319C921F85D99DFA2E0">Mientrung Institute for Scientific Research, Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Street, Hanoi, Vietnam. &amp; Graduate School of Science and Technology, Vietnam Academy of Science and Technology, Hanoi, Vietnam.</mods:affiliation>
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<treatment id="CB3887FB71756211FB48FA43FC4CFA2F" LSID="urn:lsid:plazi:treatment:CB3887FB71756211FB48FA43FC4CFA2F" httpUri="http://treatment.plazi.org/id/CB3887FB71756211FB48FA43FC4CFA2F" lastPageId="52" lastPageNumber="53" pageId="47" pageNumber="48">
<subSubSection id="0B8B65667175620AFB48FA43FB33FA63" pageId="47" pageNumber="48" type="nomenclature">
@ -67,9 +68,9 @@ urn:lsid:zoobank.org:act:
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<subSubSection id="0B8B65667175620AFBD4FA04FC9DFA5C" box="[713,874,1495,1521]" pageId="47" pageNumber="48" type="description">
<paragraph id="432E36ED7175620AFBD4FA04FC9DFA5C" blockId="47.[368,1220,1424,1521]" box="[713,874,1495,1521]" pageId="47" pageNumber="48">
<figureCitation id="DBAA2A687175620AFBD4FA04FCE5FA5C" box="[713,786,1495,1521]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="47" pageNumber="48">Figs 1</figureCitation>
<figureCitation id="DBAA2A687175620AFBD4FA04FCE5FA5C" box="[713,786,1495,1521]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="47" pageNumber="48">Figs 1</figureCitation>
,
<figureCitation id="DBAA2A687175620AFA3CFA04FC9DFA5C" box="[801,874,1495,1521]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="48.[189,232,1928,1954]" captionStartId-1="49.[189,232,1928,1954]" captionStartId-2="50.[190,233,1963,1989]" captionStartId-3="51.[189,232,1963,1989]" captionTargetBox-0="[251,1333,275,1884]" captionTargetBox-1="[243,1344,269,1872]" captionTargetBox-2="[472,1115,1470,1916]" captionTargetBox-3="[189,1398,556,1920]" captionTargetId-0="figure-16@48.[243,1344,265,1885]" captionTargetId-1="figure-11@49.[243,1344,265,1885]" captionTargetId-2="figure-340@50.[472,1115,1470,1922]" captionTargetId-3="figure-100@51.[189,1398,556,1920]" captionTargetPageId-0="48" captionTargetPageId-1="49" captionTargetPageId-2="50" captionTargetPageId-3="51" captionText-0="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." captionText-3="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." pageId="47" pageNumber="48">28–31</figureCitation>
<figureCitation id="DBAA2A687175620AFA3CFA04FC9DFA5C" box="[801,874,1495,1521]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="48.[189,232,1928,1954]" captionStartId-1="49.[189,232,1928,1954]" captionStartId-2="50.[190,233,1963,1989]" captionStartId-3="51.[189,232,1963,1989]" captionTargetBox-0="[251,1333,275,1884]" captionTargetBox-1="[243,1344,269,1872]" captionTargetBox-2="[472,1115,1470,1916]" captionTargetBox-3="[189,1398,556,1920]" captionTargetId-0="figure-16@48.[243,1344,265,1885]" captionTargetId-1="figure-11@49.[243,1344,265,1885]" captionTargetId-2="figure-340@50.[472,1115,1470,1922]" captionTargetId-3="figure-100@51.[189,1398,556,1920]" captionTargetPageId-0="48" captionTargetPageId-1="49" captionTargetPageId-2="50" captionTargetPageId-3="51" captionText-0="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." captionText-3="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." figureDoi-0="http://doi.org/10.5281/zenodo.14508794" figureDoi-1="http://doi.org/10.5281/zenodo.14508796" figureDoi-2="http://doi.org/10.5281/zenodo.14508798" figureDoi-3="http://doi.org/10.5281/zenodo.14508800" httpUri-0="https://zenodo.org/record/14508794/files/figure.png" httpUri-1="https://zenodo.org/record/14508796/files/figure.png" httpUri-2="https://zenodo.org/record/14508798/files/figure.png" httpUri-3="https://zenodo.org/record/14508800/files/figure.png" pageId="47" pageNumber="48">28–31</figureCitation>
</paragraph>
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<subSubSection id="0B8B65667175620AF9A0F9CBFDB2F809" pageId="47" pageNumber="48" type="diagnosis">
@ -87,12 +88,12 @@ can be recognized by:
</paragraph>
<paragraph id="432E36ED7175620AF9A0F9B9FE57F90A" blockId="47.[189,1398,1607,1739]" pageId="47" pageNumber="48">
(1) the variegated brown general colour of the insect, with distinctly darker, black-brown frons (
<figureCitation id="DBAA2A687175620AF81FF95EFE64F90A" box="[258,403,1677,1703]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="47" pageNumber="48">Fig. 28A–C</figureCitation>
<figureCitation id="DBAA2A687175620AF81FF95EFE64F90A" box="[258,403,1677,1703]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508794" httpUri="https://zenodo.org/record/14508794/files/figure.png" pageId="47" pageNumber="48">Fig. 28A–C</figureCitation>
);
</paragraph>
<paragraph id="432E36ED7175620AF9A0F962FABDF966" blockId="47.[189,1398,1607,1739]" box="[189,1354,1712,1739]" pageId="47" pageNumber="48">
(2) the robust cephalic process, with vertex about 1.9 times as long in midline as wide (
<figureCitation id="DBAA2A687175620AFDCDF963FAC9F966" box="[1232,1342,1712,1739]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="47" pageNumber="48">Fig. 29A</figureCitation>
<figureCitation id="DBAA2A687175620AFDCDF963FAC9F966" box="[1232,1342,1712,1739]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508796" httpUri="https://zenodo.org/record/14508796/files/figure.png" pageId="47" pageNumber="48">Fig. 29A</figureCitation>
).
</paragraph>
<paragraph id="432E36ED7175620AF9A0F921FE3EF8A1" blockId="47.[189,457,1778,1804]" box="[189,457,1778,1804]" pageId="47" pageNumber="48">
@ -119,7 +120,7 @@ The closest species is
), and by the frons not distinctly darker than the sides of head and vertex.
</paragraph>
</subSubSection>
<caption id="17EE6665716A6215F9A0F85BFE26F845" pageId="48" pageNumber="49" startId="48.[189,232,1928,1954]" targetBox="[251,1333,275,1884]" targetPageId="48">
<caption id="17EE6665716A6215F9A0F85BFE26F845" ID-DOI="http://doi.org/10.5281/zenodo.14508794" ID-Zenodo-Dep="14508794" httpUri="https://zenodo.org/record/14508794/files/figure.png" pageId="48" pageNumber="49" startId="48.[189,232,1928,1954]" targetBox="[251,1333,275,1884]" targetPageId="48">
<paragraph id="432E36ED716A6215F9A0F85BFE26F845" blockId="48.[189,1398,1928,2024]" pageId="48" pageNumber="49">
<emphasis id="71E5EAFF716A6215F9A0F85BFEEBF80F" bold="true" box="[189,284,1928,1954]" pageId="48" pageNumber="49">Fig. 28.</emphasis>
<taxonomicName id="84914D6E716A6215F83AF85BFDB0F80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[295,583,1928,1954]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="48" pageNumber="49" phylum="Arthropoda" rank="species" species="trungi" status="sp. nov.">
@ -139,7 +140,7 @@ The closest species is
. Metatarsus and apex of metatibia, ventral view.
</paragraph>
</caption>
<caption id="17EE6665716B6214F9A0F85BFE51F845" pageId="49" pageNumber="50" startId="49.[189,232,1928,1954]" targetBox="[243,1344,269,1872]" targetPageId="49">
<caption id="17EE6665716B6214F9A0F85BFE51F845" ID-DOI="http://doi.org/10.5281/zenodo.14508796" ID-Zenodo-Dep="14508796" httpUri="https://zenodo.org/record/14508796/files/figure.png" pageId="49" pageNumber="50" startId="49.[189,232,1928,1954]" targetBox="[243,1344,269,1872]" targetPageId="49">
<paragraph id="432E36ED716B6214F9A0F85BFE51F845" blockId="49.[189,1399,1928,2025]" pageId="49" pageNumber="50">
<emphasis id="71E5EAFF716B6214F9A0F85BFEE8F80F" bold="true" box="[189,287,1928,1954]" pageId="49" pageNumber="50">Fig. 29.</emphasis>
<taxonomicName id="84914D6E716B6214F837F85BFDBAF80F" authorityName="Constant &amp; Pham" authorityYear="2024" box="[298,589,1928,1954]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="49" pageNumber="50" phylum="Arthropoda" rank="species" species="trungi" status="sp. nov.">
@ -184,7 +185,7 @@ The closest species is
•
<specimenCount id="5597FD6471686217F846FE3DFE86FDA5" box="[347,369,494,520]" count="1" pageId="50" pageNumber="51" type="female">♀</specimenCount>
(
<figureCitation id="DBAA2A6871686217F89DFE3EFDFFFDA5" box="[384,520,493,520]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="48.[189,232,1928,1954]" captionStartId-1="49.[189,232,1928,1954]" captionStartId-2="50.[190,233,1963,1989]" captionTargetBox-0="[251,1333,275,1884]" captionTargetBox-1="[243,1344,269,1872]" captionTargetBox-2="[472,1115,1470,1916]" captionTargetId-0="figure-16@48.[243,1344,265,1885]" captionTargetId-1="figure-11@49.[243,1344,265,1885]" captionTargetId-2="figure-340@50.[472,1115,1470,1922]" captionTargetPageId-0="48" captionTargetPageId-1="49" captionTargetPageId-2="50" captionText-0="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." pageId="50" pageNumber="51">Figs 28–30</figureCitation>
<figureCitation id="DBAA2A6871686217F89DFE3EFDFFFDA5" box="[384,520,493,520]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="48.[189,232,1928,1954]" captionStartId-1="49.[189,232,1928,1954]" captionStartId-2="50.[190,233,1963,1989]" captionTargetBox-0="[251,1333,275,1884]" captionTargetBox-1="[243,1344,269,1872]" captionTargetBox-2="[472,1115,1470,1916]" captionTargetId-0="figure-16@48.[243,1344,265,1885]" captionTargetId-1="figure-11@49.[243,1344,265,1885]" captionTargetId-2="figure-340@50.[472,1115,1470,1922]" captionTargetPageId-0="48" captionTargetPageId-1="49" captionTargetPageId-2="50" captionText-0="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." captionText-1="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." captionText-2="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." figureDoi-0="http://doi.org/10.5281/zenodo.14508794" figureDoi-1="http://doi.org/10.5281/zenodo.14508796" figureDoi-2="http://doi.org/10.5281/zenodo.14508798" httpUri-0="https://zenodo.org/record/14508794/files/figure.png" httpUri-1="https://zenodo.org/record/14508796/files/figure.png" httpUri-2="https://zenodo.org/record/14508798/files/figure.png" pageId="50" pageNumber="51">Figs 28–30</figureCitation>
);
<location id="464E603671686217FB01FE3EFDCBFDAA" LSID="urn:lsid:plazi:treatment:CB3887FB71756211FB48FA43FC4CFA2F:464E603671686217FB01FE3EFDCBFDAA" box="[540,572,493,519]" country="Vietnam" county="Phong Dien District" latitude="16.5075" longLatPrecision="21" longitude="107.26806" municipality="Province" name="Th" pageId="50" pageNumber="51">Th</location>
ừa Thiên-Huế
@ -254,16 +255,16 @@ leg.; I.G.: 34.640;
<paragraph id="432E36ED71686217F9A0FC99FBB9FBD0" blockId="50.[188,1400,842,1150]" pageId="50" pageNumber="51">
<smallCapsWord id="45C8A03171686217F9A0FC99FF0BFCCF" baselines="861,861" box="[189,252,842,868]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Head" pageId="50" pageNumber="51">HEAD</smallCapsWord>
(
<figureCitation id="DBAA2A6871686217F80BFC99FE58FCC9" box="[278,431,842,868]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="50" pageNumber="51">Fig. 29A–D</figureCitation>
<figureCitation id="DBAA2A6871686217F80BFC99FE58FCC9" box="[278,431,842,868]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508796" httpUri="https://zenodo.org/record/14508796/files/figure.png" pageId="50" pageNumber="51">Fig. 29A–D</figureCitation>
). Strongly elongated anteriorly in cephalic process; vertex variegated brown; distinctly longer in midline than broad before eyes (about 1.9 times), widening from base to anterior margin of eyes, tapering beyond eyes to narrowly rounded apex, median and lateral carinae distinct; posterior margin moderately incurved. Side of head yellow-brown but coloured as vertex on sides of cephalic process. Frons entirely blackish brown, distinctly darker than sides of head and vertex; lateral carinae reaching apex, median carina distinct in distal half; elongate, widest slightly anterior to eyes, roundly tapering towards clypeal suture; clypeal suture rounded. Clypeus black-brown, weakly elevated medially in distal portion. Labium brown with last segment longer than broad, and shorter than penultimate. Antennae dark brown; scape short, ring-shaped; pedicel bulbous.
</paragraph>
<paragraph id="432E36ED71686216F9A0FB76FEBDFEEB" blockId="50.[189,1399,1187,1391]" lastBlockId="51.[188,1398,265,326]" lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51">
<smallCapsWord id="45C8A03171686217F9A0FB76FEEEFB10" baselines="1208,1208" box="[189,281,1189,1215]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Thorax" pageId="50" pageNumber="51">THORAX</smallCapsWord>
(
<figureCitation id="DBAA2A6871686217F833FB76FE33FB12" box="[302,452,1189,1215]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="50" pageNumber="51">Fig. 29A–D</figureCitation>
<figureCitation id="DBAA2A6871686217F833FB76FE33FB12" box="[302,452,1189,1215]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508796" httpUri="https://zenodo.org/record/14508796/files/figure.png" pageId="50" pageNumber="51">Fig. 29A–D</figureCitation>
). Pronotum coloured as vertex; nearly ⅔ of length of mesonotum in midline; anterior margin weakly carinate, sinuate and moderately protruding anteriorly between eyes, with peridiscal carinae indistinct; posterior margin nearly straight; median carina obsolete; weak impressed point on each side of median line; weak, blunt tubercles along anterior and posterior margins; paranotal lobes (lateral view) moderately broad, turning dark brown in lower portion, under level of eye and with posteroventral angle rather narrowly rounded. Mesonotum coloured like vertex with longitudinal carinae very weak and blunt; two weakly impressed points on disc; tip of scutellum pale yellow. Tegulae dark brown.
</paragraph>
<caption id="17EE666571686217F9A3F878FE06F845" pageId="50" pageNumber="51" startId="50.[190,233,1963,1989]" targetBox="[472,1115,1470,1916]" targetPageId="50">
<caption id="17EE666571686217F9A3F878FE06F845" ID-DOI="http://doi.org/10.5281/zenodo.14508798" ID-Zenodo-Dep="14508798" httpUri="https://zenodo.org/record/14508798/files/figure.png" pageId="50" pageNumber="51" startId="50.[190,233,1963,1989]" targetBox="[472,1115,1470,1916]" targetPageId="50">
<paragraph id="432E36ED71686217F9A3F878FE06F845" blockId="50.[189,1399,1963,2025]" pageId="50" pageNumber="51">
<emphasis id="71E5EAFF71686217F9A3F878FEECF868" bold="true" box="[190,283,1963,1989]" pageId="50" pageNumber="51">Fig. 30.</emphasis>
<taxonomicName id="84914D6E71686217F802F878FDCAF868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[287,573,1963,1989]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="50" pageNumber="51" phylum="Arthropoda" rank="species" species="trungi" status="sp. nov.">
@ -276,17 +277,17 @@ leg.; I.G.: 34.640;
<paragraph id="432E36ED71696211F9A0FEBDFA81FE5A" blockId="51.[189,1399,366,497]" lastBlockId="52.[189,1399,263,503]" lastPageId="52" lastPageNumber="53" pageId="51" pageNumber="52">
<smallCapsWord id="45C8A03171696216F9A0FEBDFED5FE2B" baselines="385,385" box="[189,290,366,392]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Tegmina" pageId="51" pageNumber="52">TEGMINA</smallCapsWord>
(
<figureCitation id="DBAA2A6871696216F82EFEBDFE32FE25" box="[307,453,366,392]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="51" pageNumber="52">
<figureCitation id="DBAA2A6871696216F82EFEBDFE32FE25" box="[307,453,366,392]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508794" httpUri="https://zenodo.org/record/14508794/files/figure.png" pageId="51" pageNumber="52">
<smallCapsWord id="45C8A03171696216F82EFEBDFE91FE2B" baselines="385" box="[307,358,366,392]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="s" pageId="51" pageNumber="52">FigS</smallCapsWord>
28A–C
</figureCitation>
,
<figureCitation id="DBAA2A6871696216F8CFFEBDFDD0FE25" box="[466,551,366,392]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." pageId="51" pageNumber="52">31C–F</figureCitation>
<figureCitation id="DBAA2A6871696216F8CFFEBDFDD0FE25" box="[466,551,366,392]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508800" httpUri="https://zenodo.org/record/14508800/files/figure.png" pageId="51" pageNumber="52">31C–F</figureCitation>
). Dark brown, washed with olivaceous in basal portion, black transverse band just beyond midlength, at about basal ⅓, transverse band of white wax, apical ⅓ covered in white wax, basal ⅓ with scattered small points of white wax; veins concolourous except in black band, where red; subcoriaceous with longitudinal veins distinct and with dense reticulum of veinlets; shape elongate and convex with sides broadly rounded, widest at basal ⅓, about 2.5 times as long as broad; narrowly rounded apically. Postclaval margin nearly straight, very weakly rounded on distal half, and notched at apex of clavus. Clavus closed, reaching about ⅔ of tegmen. Venation: ScP+R forking close to base after short common stem, ScP+RA and RP unforked, running more or less parallel to costal margin in a large basal portion; MP forked slightly before basal ⅓, resulting veins unforked, running more or less parallel; CuA weakly diverging from claval joint; Pcu and A 1 fused at distal ¼ of clavus, resulting Pcu+A
<subScript id="DF1534A8716E6211F812FE3CFEEFFE50" attach="left" box="[271,280,495,509]" fontSize="6" pageId="52" pageNumber="53">1</subScript>
reaching claval joint before apex of clavus; dense reticulum of veinlets, especially after basal ⅓.
</paragraph>
<caption id="17EE666571696216F9A0F878FDBEF845" pageId="51" pageNumber="52" startId="51.[189,232,1963,1989]" targetBox="[189,1398,556,1920]" targetPageId="51">
<caption id="17EE666571696216F9A0F878FDBEF845" ID-DOI="http://doi.org/10.5281/zenodo.14508800" ID-Zenodo-Dep="14508800" httpUri="https://zenodo.org/record/14508800/files/figure.png" pageId="51" pageNumber="52" startId="51.[189,232,1963,1989]" targetBox="[189,1398,556,1920]" targetPageId="51">
<paragraph id="432E36ED71696216F9A0F878FDBEF845" blockId="51.[187,1399,1963,2024]" pageId="51" pageNumber="52">
<emphasis id="71E5EAFF71696216F9A0F878FEEFF868" bold="true" box="[189,280,1963,1989]" pageId="51" pageNumber="52">Fig. 31.</emphasis>
<taxonomicName id="84914D6E71696216F803F878FDCCF868" authorityName="Constant &amp; Pham" authorityYear="2024" box="[286,571,1963,1989]" class="Insecta" family="Issidae" genus="Pseudochoutagus" kingdom="Animalia" order="Hemiptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species" species="trungi" status="sp. nov.">
@ -304,7 +305,7 @@ in Phong Dien District, 17 May 2023.
<smallCapsWord id="45C8A031716E6211F9A0FDCDFF00FD9B" baselines="561,561" box="[189,247,542,568]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Hind" pageId="52" pageNumber="53">HIND</smallCapsWord>
<smallCapsWord id="45C8A031716E6211F81AFDF0FEBCFD9B" baselines="561" box="[263,331,547,566]" lowerCaseFontSize="8" mainFontSize="11" normCase="lower" normString="wings" pageId="52" pageNumber="53">WINGS</smallCapsWord>
(
<figureCitation id="DBAA2A68716E6211F879FDCDFE20FD95" box="[356,471,542,568]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." pageId="52" pageNumber="53">Fig. 29E</figureCitation>
<figureCitation id="DBAA2A68716E6211F879FDCDFE20FD95" box="[356,471,542,568]" captionStart="Fig" captionStartId="49.[189,232,1928,1954]" captionTargetBox="[243,1344,269,1872]" captionTargetId="figure-11@49.[243,1344,265,1885]" captionTargetPageId="49" captionText="Fig. 29. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Head and thorax, dorsal view. B. Frons, perpendicular view. C. Head and thorax, lateral view. D. Head and thorax, anterolateral view. E. Right hind wing." figureDoi="http://doi.org/10.5281/zenodo.14508796" httpUri="https://zenodo.org/record/14508796/files/figure.png" pageId="52" pageNumber="53">Fig. 29E</figureCitation>
). Black-brown, slightly paler along costal margin in basal half; venation concolourous; wing broader than tegmen and deeply bilobed at CuP; costal margin sinuate; CuP-Pcu-A 1 lobe broadly rounded along postclaval margin and about 1.3 times as wide as Sc-R-MP-CuA lobe, both about the same length and apically rounded; A
<subScript id="DF1534A8716E6211FA30FD49FCC1FD05" attach="left" box="[813,822,666,680]" fontSize="6" pageId="52" pageNumber="53">2</subScript>
lobe moderately reduced and narrow. Venation: longitudinal veins ScP-R-MP-Cu well distinct; Pcu and A
@ -316,13 +317,13 @@ rather weak.
<paragraph id="432E36ED716E6211F9A0FCC3FD9BFC3E" blockId="52.[188,1399,783,915]" pageId="52" pageNumber="53">
<smallCapsWord id="45C8A031716E6211F9A0FCC3FF01FC85" baselines="803,803" box="[189,246,784,810]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Legs" pageId="52" pageNumber="53">LEGS</smallCapsWord>
(
<figureCitation id="DBAA2A68716E6211F81AFCDCFE63FC84" box="[263,404,783,810]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="52" pageNumber="53">Fig. 28A–E</figureCitation>
<figureCitation id="DBAA2A68716E6211F81AFCDCFE63FC84" box="[263,404,783,810]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508794" httpUri="https://zenodo.org/record/14508794/files/figure.png" pageId="52" pageNumber="53">Fig. 28A–E</figureCitation>
). Moderately elongate, slender; brown with apex of pro- and mesotibiae, posterobasal portion of metafemora and basal half of first segment of metatarsi, yellowish; femora wider than corresponding tibiae; metatibiae with two lateral spines in distal half and seven apical spines. Metatibiotarsal formula: (2) 7 / 9 / 2.
</paragraph>
<paragraph id="432E36ED716E6211F9A0FC69FBEBFC79" blockId="52.[189,1052,954,981]" box="[189,1052,954,981]" pageId="52" pageNumber="53">
<smallCapsWord id="45C8A031716E6211F9A0FC69FEDBFC7E" baselines="973,974" box="[189,300,954,980]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Abdomen" pageId="52" pageNumber="53">ABDOMEN</smallCapsWord>
(
<figureCitation id="DBAA2A68716E6211F823FC69FE5FFC79" box="[318,424,954,981]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." pageId="52" pageNumber="53">Fig. 28B</figureCitation>
<figureCitation id="DBAA2A68716E6211F823FC69FE5FFC79" box="[318,424,954,981]" captionStart="Fig" captionStartId="48.[189,232,1928,1954]" captionTargetBox="[251,1333,275,1884]" captionTargetId="figure-16@48.[243,1344,265,1885]" captionTargetPageId="48" captionText="Fig. 28. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral view. D. Right hind leg, ventral view. E. Metatarsus and apex of metatibia, ventral view." figureDoi="http://doi.org/10.5281/zenodo.14508794" httpUri="https://zenodo.org/record/14508794/files/figure.png" pageId="52" pageNumber="53">Fig. 28B</figureCitation>
). Yellowish brown with dark brown median marking.
</paragraph>
<paragraph id="432E36ED716E6211F9A0FC2FFE7BFB94" blockId="52.[189,1398,1019,1081]" pageId="52" pageNumber="53">
@ -331,7 +332,7 @@ rather weak.
. Hind margin of sternum VII with massive, long, apically rounded median process (
<emphasis id="71E5EAFF716E6211F9DAFBCCFEFDFB95" box="[199,266,1055,1081]" italics="true" pageId="52" pageNumber="53">St VII</emphasis>
–
<figureCitation id="DBAA2A68716E6211F835FBCCFE89FB94" box="[296,382,1055,1081]" captionStart="Fig" captionStartId="50.[190,233,1963,1989]" captionTargetBox="[472,1115,1470,1916]" captionTargetId="figure-340@50.[472,1115,1470,1922]" captionTargetPageId="50" captionText="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." pageId="52" pageNumber="53">Fig. 30</figureCitation>
<figureCitation id="DBAA2A68716E6211F835FBCCFE89FB94" box="[296,382,1055,1081]" captionStart="Fig" captionStartId="50.[190,233,1963,1989]" captionTargetBox="[472,1115,1470,1916]" captionTargetId="figure-340@50.[472,1115,1470,1922]" captionTargetPageId="50" captionText="Fig. 30. Pseudochoutagus trungi sp. nov., holotype ♀ (VNMN). Terminalia, ventral view. Abbreviation: see Material and methods." figureDoi="http://doi.org/10.5281/zenodo.14508798" httpUri="https://zenodo.org/record/14508798/files/figure.png" pageId="52" pageNumber="53">Fig. 30</figureCitation>
).
</paragraph>
</subSubSection>
@ -347,13 +348,13 @@ rather weak.
</taxonomicName>
<taxonomicNameLabel id="6AD65784716E6211F8FDFB5CFDCEFB04" box="[480,569,1167,1193]" pageId="52" pageNumber="53" rank="species">sp. nov.</taxonomicNameLabel>
was collected in May in a marshy microhabitat (
<figureCitation id="DBAA2A68716E6211FD71FB5CFB22FB04" box="[1132,1237,1167,1193]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." pageId="52" pageNumber="53">Fig. 31A</figureCitation>
<figureCitation id="DBAA2A68716E6211FD71FB5CFB22FB04" box="[1132,1237,1167,1193]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508800" httpUri="https://zenodo.org/record/14508800/files/figure.png" pageId="52" pageNumber="53">Fig. 31A</figureCitation>
) by sweeping the lower vegetation (mostly ferns and palms) (
<figureCitation id="DBAA2A68716E6211FBE4FB61FC91FB61" box="[761,870,1202,1228]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." pageId="52" pageNumber="53">Fig. 31B</figureCitation>
<figureCitation id="DBAA2A68716E6211FBE4FB61FC91FB61" box="[761,870,1202,1228]" captionStart="Fig" captionStartId="51.[189,232,1963,1989]" captionTargetBox="[189,1398,556,1920]" captionTargetId="figure-100@51.[189,1398,556,1920]" captionTargetPageId="51" captionText="Fig. 31. Pseudochoutagus trungi sp. nov. in Phong Dien District, 17 May 2023.A–B. Habitat.C–F. Live specimen (photographed in cage)." figureDoi="http://doi.org/10.5281/zenodo.14508800" httpUri="https://zenodo.org/record/14508800/files/figure.png" pageId="52" pageNumber="53">Fig. 31B</figureCitation>
), in moist evergreen tropical forest at about
<quantity id="84699B08716E6211F9A2FB06FEFCFB5D" box="[191,267,1237,1264]" metricMagnitude="2" metricUnit="m" metricValue="3.5" pageId="52" pageNumber="53" unit="m" value="350.0">350 m</quantity>
in altitude. The variegated brown colour of adult specimens (
<figureCitation id="DBAA2A68716E6211FAF2FB06FB8DFB42" box="[1007,1146,1237,1263]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." pageId="52" pageNumber="53">Fig. 27C–F</figureCitation>
<figureCitation id="DBAA2A68716E6211FAF2FB06FB8DFB42" box="[1007,1146,1237,1263]" captionStart="Fig" captionStartId="46.[189,232,1963,1989]" captionTargetBox="[189,1398,1062,1920]" captionTargetId="figure-293@46.[189,1398,1062,1920]" captionTargetPageId="46" captionText="Fig. 27. Pseudochoutagus lindae sp. nov. in Bach Ma National Park, Yes Hue Eco, 17 May 2023. A. Habitat. B. Host plant. C–E. Live specimen." figureDoi="http://doi.org/10.5281/zenodo.14508792" httpUri="https://zenodo.org/record/14508792/files/figure.png" pageId="52" pageNumber="53">Fig. 27C–F</figureCitation>
) probably helps their camouflage on their host plants.
</paragraph>
</subSubSection>
@ -366,7 +367,7 @@ in altitude. The variegated brown colour of adult specimens (
<paragraph id="432E36ED716E6211F9A0FABBFC4CFA2F" blockId="52.[189,955,1384,1413]" box="[189,955,1384,1413]" pageId="52" pageNumber="53">
<collectingCountry id="3B86767D716E6211F9A0FABBFED2FA2F" box="[189,293,1384,1410]" name="Vietnam" pageId="52" pageNumber="53">Vietnam</collectingCountry>
: Thừa Thiên-Huế Province, Phong Dien district (
<figureCitation id="DBAA2A68716E6211FA78FABBFC5DFA2F" box="[869,938,1384,1410]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." pageId="52" pageNumber="53">Fig. 1</figureCitation>
<figureCitation id="DBAA2A68716E6211FA78FABBFC5DFA2F" box="[869,938,1384,1410]" captionStart="Fig" captionStartId="6.[189,232,1998,2024]" captionTargetBox="[189,1398,729,1950]" captionTargetId="figure-138@6.[189,1398,729,1950]" captionTargetPageId="6" captionText="Fig. 1. Map of the sampled localities in Thừa Thiên-Huế Province, Vietnam." figureDoi="http://doi.org/10.5281/zenodo.14508732" httpUri="https://zenodo.org/record/14508732/files/figure.png" pageId="52" pageNumber="53">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>