treatments-xml/data/5B/56/E8/5B56E872FFB34A78FF328A2EFE9BE88B.xml

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<document id="062388B4E777CBB04C12B0B6ACA87005" ID-CLB-Dataset="26920" ID-DOI="10.11646/zootaxa.4599.1.1" ID-GBIF-Dataset="ea727a9b-324b-4c6b-a625-bc7cdd204473" ID-ISSN="1175-5326" ID-Zenodo-Dep="2668058" ID-ZooBank="7CDD24FF-5BA3-4D06-898D-A3E220515D4C" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1557127209543" checkinUser="plazi" docAuthor="Longino, John T." docDate="2019" docId="5B56E872FFB34A78FF328A2EFE9BE88B" docLanguage="en" docName="zootaxa.4599.1.1.pdf.imf" docOrigin="Zootaxa 4599 (1)" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="14" docTitle="Pheidole passivaeferox Longino 2019, new species" docType="treatment" docVersion="23" lastPageNumber="52" masterDocId="A76F900AFF804A4BFFA58B6FFFF4EF5B" masterDocTitle="Pheidole (Hymenoptera, Formicidae) of Middle American Wet Forest" masterLastPageNumber="126" masterPageNumber="1" pageNumber="52" updateTime="1732580243751" updateUser="ExternalLinkService">
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<mods:title id="AC2DB0C9925822FC52E06A36E049D608">Pheidole (Hymenoptera, Formicidae) of Middle American Wet Forest</mods:title>
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<mods:namePart id="BDE06F73A505B7796957B3F09EA5B672">Longino, John T.</mods:namePart>
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<mods:title id="A7D0BDBAB09A66ECC3118B70A397B412">Zootaxa</mods:title>
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<mods:date id="778A65ABF4ADE9092272251FC03D21BC">2019</mods:date>
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<mods:number id="B40DB46E6C3273DF17013625BE9B926C">2019-05-06</mods:number>
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<mods:number id="593E971DD3789AB78CB4DC0A1F759BD5">4599</mods:number>
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<treatment id="5B56E872FFB34A78FF328A2EFE9BE88B" ID-DOI="http://doi.org/10.5281/zenodo.5586226" ID-GBIF-Taxon="157062632" ID-Zenodo-Dep="5586226" LSID="urn:lsid:plazi:treatment:5B56E872FFB34A78FF328A2EFE9BE88B" httpUri="http://treatment.plazi.org/id/5B56E872FFB34A78FF328A2EFE9BE88B" lastPageNumber="52" pageId="51" pageNumber="52" scope_family="Formicidae" scope_order="Hymenoptera">
<subSubSection id="9BE50AEFFFB34A78FF328A2EFEF0EE26" pageId="51" pageNumber="52" type="nomenclature">
<paragraph id="D3405964FFB34A78FF328A2EFDB7EE00" blockId="51.[151,579,321,381]" box="[151,579,321,347]" pageId="51" pageNumber="52">
<heading id="8808EE08FFB34A78FF328A2EFDB7EE00" bold="true" box="[151,579,321,347]" fontSize="11" level="1" pageId="51" pageNumber="52" reason="1">
<emphasis id="E18B8576FFB34A78FF328A2EFDB7EE00" bold="true" box="[151,579,321,347]" pageId="51" pageNumber="52">
<taxonomicName id="14FF22E7FFB34A78FF328A2EFE58EE00" ID-CoL="CMLMX" authority="Longino, 2019" authorityName="Longino" authorityYear="2019" box="[151,428,321,347]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species" species="passivaeferox" status="sp. nov.">
<emphasis id="E18B8576FFB34A78FF328A2EFE58EE00" bold="true" box="[151,428,321,347]" italics="true" pageId="51" pageNumber="52">Pheidole passivaeferox</emphasis>
</taxonomicName>
<taxonomicNameLabel id="FAB8380DFFB34A78FE118A2EFDB7EE00" box="[436,579,321,347]" pageId="51" pageNumber="52" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="D3405964FFB34A78FF328A0BFEF0EE26" blockId="51.[151,579,321,381]" box="[151,260,356,381]" pageId="51" pageNumber="52">
(
<figureCitation id="4BC445E1FFB34A78FF3B8A0BFF08EE26" box="[158,252,356,381]" captionStart="PLATE 25" captionStartId="91.[151,238,1954,1977]" captionTargetId="figure@91.[202,1384,194,1933]" captionTargetPageId="91" captionText="PLATE 25." httpUri="https://zenodo.org/record/2668279/files/figure.png" pageId="51" pageNumber="52">Plate 25</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF328AC3FC6FEE9E" box="[151,923,428,453]" pageId="51" pageNumber="52" type="description">
<paragraph id="D3405964FFB34A78FF328AC3FC6FEE9E" blockId="51.[151,1437,428,2000]" box="[151,923,428,453]" pageId="51" pageNumber="52">
<taxonomicName id="14FF22E7FFB34A78FF328AC3FF0FEE9E" authorityName="Biolleyi Forel" authorityYear="1908" box="[151,251,428,453]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="genus">
<emphasis id="E18B8576FFB34A78FF328AC3FF0FEE9E" box="[151,251,428,453]" italics="true" pageId="51" pageNumber="52">Pheidole</emphasis>
</taxonomicName>
JTL-231: morphospecies code previously used on AntWeb.
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF628ABFFD5BED23" pageId="51" pageNumber="52" type="materials_examined">
<paragraph id="D3405964FFB34A78FF628ABFFDACED0F" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<materialsCitation id="63975339FFB34A78FF628ABFFD77ED6B" ID-GBIF-Occurrence="2237851525" collectingDate="1986-07-21" collectionCode="MUCR" collectorName="J. Longino" country="Costa Rica" elevation="50" location="La Selva Biological Station" pageId="51" pageNumber="52" specimenCount="1" stateProvince="Heredia" typeStatus="holotype">
<typeStatus id="0C44E7C6FFB34A78FF628ABFFEABEEB3" box="[199,351,464,488]" pageId="51" pageNumber="52" type="holotype">HOLOTYPE</typeStatus>
: 1 major worker,
<collectingCountry id="ABE819F4FFB34A78FD808ABFFD56EEB3" box="[549,674,463,488]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
,
<collectingRegion id="113B9786FFB34A78FD0B8AA0FCFFEEB3" box="[686,779,463,488]" country="Costa Rica" name="Heredia" pageId="51" pageNumber="52">Heredia</collectingRegion>
:
<location id="D6200FBFFFB34A78FCB38ABFFBBBEEB3" LSID="urn:lsid:plazi:treatment:5B56E872FFB34A78FF328A2EFE9BE88B:D6200FBFFFB34A78FCB38ABFFBBBEEB3" box="[790,1103,463,488]" country="Costa Rica" name="La Selva Biological Station" pageId="51" pageNumber="52" stateProvince="Heredia">La Selva Biological Station</location>
, 10.43333 -84.01667 ±
<quantity id="1407F481FFB34A78FAF98ABFFA6CEEB3" box="[1372,1432,463,488]" metricMagnitude="3" metricUnit="m" metricValue="2.0" pageId="51" pageNumber="52" unit="km" value="2.0">2 km</quantity>
,
<quantity id="1407F481FFB34A78FF328A9BFF2DED57" box="[151,217,500,525]" metricMagnitude="1" metricUnit="m" metricValue="5.0" pageId="51" pageNumber="52" unit="m" value="50.0">
<elevation id="58D2BE57FFB34A78FF328A9BFF2DED57" box="[151,217,500,525]" metricMagnitude="1" metricUnit="m" metricValue="5.0" pageId="51" pageNumber="52" unit="m" value="50.0">50 m</elevation>
</quantity>
,
<date id="A7417FA4FFB34A78FF488A9BFE83ED56" box="[237,375,500,525]" pageId="51" pageNumber="52" value="1986-07-21">
<collectingDate id="B705864CFFB34A78FF488A9BFE83ED56" box="[237,375,500,525]" pageId="51" pageNumber="52" value="1986-07-21">21-Jul-1986</collectingDate>
</date>
, lowland rainforest, nest in
<taxonomicName id="14FF22E7FFB34A78FD768A9AFC41ED56" box="[723,949,500,525]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="species" species="coenocladum">
<emphasis id="E18B8576FFB34A78FD768A9AFC41ED56" box="[723,949,500,525]" italics="true" pageId="51" pageNumber="52">Piper coenocladum</emphasis>
</taxonomicName>
(
<collectorName id="7E0A3CB2FFB34A78FC688A9BFBBBED56" box="[973,1103,500,525]" pageId="51" pageNumber="52">J. Longino</collectorName>
, JTL1429) [
<collectionCode id="B5EEC1A1FFB34A78FB408A9BFAC9ED57" box="[1253,1341,500,524]" country="Costa Rica" name="Museo de Insectos" pageId="51" pageNumber="52">MUCR</collectionCode>
, unique specimen identifier INBIOCRI002279624]
</materialsCitation>
.
<materialsCitation id="63975339FFB34A78FD348977FDA1ED0F" ID-GBIF-Occurrence="2237851593" collectingDate="1986-07-21" collectionCode="CAS, DZUP, JTLC, MCZC, MUCR, USNM" collectorName="J. Longino" country="Costa Rica" elevation="50" location="La Selva Biological Station" pageId="51" pageNumber="52" specimenCount="1" stateProvince="Heredia" typeStatus="paratype">
<typeStatus id="0C44E7C6FFB34A78FD348977FCC4ED6B" box="[657,816,536,560]" pageId="51" pageNumber="52" type="paratype">PARATYPES</typeStatus>
: major, minor workers: same data as holotype [
<collectionCode id="B5EEC1A1FFB34A78FAF98977FA63ED6B" box="[1372,1431,536,560]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15690" lsid="urn:lsid:biocol.org:col:34700" name="California Academy of Sciences" pageId="51" pageNumber="52" type="Museum">CAS</collectionCode>
,
<collectionCode id="B5EEC1A1FFB34A78FF328952FF14ED0F" box="[151,224,573,596]" country="Brazil" httpUri="http://grbio.org/cool/5xp9-edpx" name="Universidade Federal do Parana, Colecao de Entomologia Pe. Jesus Santiago Moure" pageId="51" pageNumber="52" type="University or college">DZUP</collectionCode>
,
<collectionCode id="B5EEC1A1FFB34A78FF4E8953FEDAED0F" box="[235,302,572,596]" pageId="51" pageNumber="52">JTLC</collectionCode>
,
<collectionCode id="B5EEC1A1FFB34A78FE9C8953FE7FED0F" box="[313,395,572,596]" pageId="51" pageNumber="52">MCZC</collectionCode>
,
<collectionCode id="B5EEC1A1FFB34A78FE338953FE18ED0F" box="[406,492,572,596]" country="Costa Rica" name="Museo de Insectos" pageId="51" pageNumber="52">MUCR</collectionCode>
,
<collectionCode id="B5EEC1A1FFB34A78FE538953FDB9ED0F" box="[502,589,572,596]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="51" pageNumber="52" type="Museum">USNM</collectionCode>
]
</materialsCitation>
.
</paragraph>
<paragraph id="D3405964FFB34A78FF628930FD5BED23" blockId="51.[151,1437,428,2000]" box="[199,687,607,632]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628930FE51ED23" bold="true" box="[199,421,607,632]" pageId="51" pageNumber="52">Geographic range.</emphasis>
<collectingCountry id="ABE819F4FFB34A78FE098930FDD6ED23" box="[428,546,607,632]" name="Nicaragua" pageId="51" pageNumber="52">Nicaragua</collectingCountry>
,
<collectingCountry id="ABE819F4FFB34A78FD88890FFD5FED23" box="[557,683,607,632]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF6289EBFA91ECBB" pageId="51" pageNumber="52" type="diagnosis">
<paragraph id="D3405964FFB34A78FF6289EBFA91ECBB" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF6289EBFE66EDC7" bold="true" box="[199,402,643,669]" pageId="51" pageNumber="52">
Diagnosis.
<emphasis id="E18B8576FFB34A78FEEC89ECFE66EDC7" bold="true" box="[329,402,643,668]" italics="true" pageId="51" pageNumber="52">Minor</emphasis>
</emphasis>
: face smooth and shining, with variably developed faint patches of foveolation at margins; head rounded behind; promesonotal groove absent; pronotum mostly smooth and shining, with patches of weak foveolation on humeri and ventral margins; promesonotal dorsum grading to foveolate posteriorly; katepisternum foveolate; propodeal spines relatively thin and spiniform, about one third length of posterior face of propodeum; gaster smooth and shining; abundant, flexuous, almost woolly setae on mesosomal dorsum, tibiae, and gastral dorsum; color yellow brown.
<emphasis id="E18B8576FFB34A78FE578858FDCEEC0B" bold="true" box="[498,570,823,848]" italics="true" pageId="51" pageNumber="52">Major</emphasis>
: inner hypostomal teeth distinct, closely spaced; scape base terete; frontal carinae produced anteriorly as elevated, triangular teeth, most visible in profile; head elongate rectangular; scrobes shallowly impressed; face heavily sculptured, mostly reticulate rugose, rugae becoming more transversely oriented posteriorly; propodeal spines spiniform, about one fourth length of posterior face of propodeum; gastral dorsum smooth and shiny; abundant erect setae on face, sides of head, mesosomal dorsum, tibiae, and gastral dorsum.
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF628883FDE8EBE3" pageId="51" pageNumber="52" type="description">
<paragraph id="D3405964FFB34A78FF628883FD9BEB73" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628883FDD9EB5E" bold="true" box="[199,557,1004,1029]" pageId="51" pageNumber="52">Measurements, minor worker</emphasis>
: HW 0.51, HL 0.58, SL 0.40, EL 0.12, WL 0.73, PSL 0.04, PTW 0.12, PPW 0.16, CI 88, SI 79, PSLI 9, PPI 132 (n=7).
</paragraph>
<paragraph id="D3405964FFB34A78FF628F5BFD89EB2B" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628F5BFDD9EB16" bold="true" box="[199,557,1076,1101]" pageId="51" pageNumber="52">Measurements, major worker</emphasis>
: HW 0.80, HL 1.08, SL 0.43, EL 0.14, WL 1.04, PSL 0.09, PTW 0.18, PPW 0.26, CI 74, SI 54, PSLI 11, PPI 141 (n=5).
</paragraph>
<paragraph id="D3405964FFB34A78FF628F13FDE8EBE3" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628F13FE38EBCE" bold="true" box="[199,460,1148,1173]" pageId="51" pageNumber="52">Measurements, queen</emphasis>
: HW 0.71, HL 0.90, SL 0.46, EL 0.24, WL 1.29, PSL 0.07, PTW 0.20, PPW 0.32, CI 80, SI 65, PSLI 10, PPI 164 (n=2).
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF628FABFB9BEA5B" pageId="51" pageNumber="52" type="biology_ecology">
<paragraph id="D3405964FFB34A78FF628FABFB9BEA5B" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628FABFED3EB86" bold="true" box="[199,295,1220,1245]" pageId="51" pageNumber="52">Biology.</emphasis>
This species occurs in lower rainforest. It is a specialized plant ant, nesting in domatia of several species of myrmecophytic
<taxonomicName id="14FF22E7FFB34A78FE618F86FDF6EA5B" box="[452,514,1257,1280]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="genus">
<emphasis id="E18B8576FFB34A78FE618F86FDF6EA5B" box="[452,514,1257,1280]" italics="true" pageId="51" pageNumber="52">Piper</emphasis>
</taxonomicName>
(
<bibRefCitation id="B76E2495FFB34A78FDB58F88FD27EA5B" author="Risch, S. &amp; McClure, M. &amp; Vandermeer, J. &amp; Waltz, S." box="[528,723,1255,1280]" pageId="51" pageNumber="52" pagination="433 - 444" refId="ref52291" refString="Risch, S., McClure, M., Vandermeer, J. &amp; Waltz, S. (1977) Mutualism between three species of tropical Piper (Piperaceae) and their ant inhabitants. American Midland Naturalist, 98, 433 - 444. https: // doi. org / 10.2307 / 2424991" type="journal article" year="1977">
Risch
<emphasis id="E18B8576FFB34A78FDFC8F86FD65EA5B" box="[601,657,1255,1280]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
1977
</bibRefCitation>
,
<bibRefCitation id="B76E2495FFB34A78FD7B8F87FC54EA5B" author="Letourneau, D. K." box="[734,928,1256,1280]" pageId="51" pageNumber="52" pagination="122 - 126" refId="ref51627" refString="Letourneau, D. K. (1983) Passive aggression: an alternative hypothesis for the Piper-Pheidole association. Oecologia, 60, 122 - 126. https: // doi. org / 10.1007 / BF 00379331" type="journal article" year="1983">Letourneau 1983</bibRefCitation>
,
<bibRefCitation id="B76E2495FFB34A78FC0E8F87FB97EA5B" author="Tepe, E. &amp; Vincent, M. &amp; Watson, L." box="[939,1123,1255,1280]" pageId="51" pageNumber="52" pagination="156 - 178" refId="ref52546" refString="Tepe, E., Vincent, M. &amp; Watson, L. (2004) Phylogenetic patterns, evolutionary trends, and the origin of ant-plant associations in Piper section Macrostachys: Burger's hypothesis revisited. In: L. A. Dyer &amp; A. D. N. Palmer (Eds), Piper: a model genus for studies of phytochemistry, ecology, and evolution. Kluwer Academic Publishing, New York, pp. 156 - 178. https: // doi. org / 10.1007 / 978 - 0 - 387 - 30599 - 8 _ 9" type="book chapter" year="2004">
Tepe
<emphasis id="E18B8576FFB34A78FC428F86FBD4EA5B" box="[999,1056,1255,1280]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
2004
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF628E63FC74E8D3" pageId="51" pageNumber="52" type="discussion">
<paragraph id="D3405964FFB34A78FF628E63FB46EAA6" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628E63FEBCEA7E" bold="true" box="[199,328,1292,1317]" pageId="51" pageNumber="52">Comments</emphasis>
. One of the myrmecophytic systems that has evolved in the Neotropics involves the plant genus
<taxonomicName id="14FF22E7FFB34A78FF328E5EFF22EA13" box="[151,214,1329,1352]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="genus">
<emphasis id="E18B8576FFB34A78FF328E5EFF22EA13" box="[151,214,1329,1352]" italics="true" pageId="51" pageNumber="52">Piper</emphasis>
</taxonomicName>
and specialized
<taxonomicName id="14FF22E7FFB34A78FE338E40FE0EEA13" authorityName="Biolleyi Forel" authorityYear="1908" box="[406,506,1327,1352]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="genus">
<emphasis id="E18B8576FFB34A78FE338E40FE0EEA13" box="[406,506,1327,1352]" italics="true" pageId="51" pageNumber="52">Pheidole</emphasis>
</taxonomicName>
inhabitants. A set of multiple
<taxonomicName id="14FF22E7FFB34A78FCF88E5EFC68EA13" box="[861,924,1329,1352]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="genus">
<emphasis id="E18B8576FFB34A78FCF88E5EFC68EA13" box="[861,924,1329,1352]" italics="true" pageId="51" pageNumber="52">Piper</emphasis>
</taxonomicName>
species have modified clasping petioles and stem pith that favors nesting by
<taxonomicName id="14FF22E7FFB34A78FD8C8E3BFD79EA36" authorityName="Biolleyi Forel" authorityYear="1908" box="[553,653,1364,1389]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="genus">
<emphasis id="E18B8576FFB34A78FD8C8E3BFD79EA36" box="[553,653,1364,1389]" italics="true" pageId="51" pageNumber="52">Pheidole</emphasis>
</taxonomicName>
. These
<taxonomicName id="14FF22E7FFB34A78FD568E3AFCC5EA37" box="[755,817,1365,1388]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="genus">
<emphasis id="E18B8576FFB34A78FD568E3AFCC5EA37" box="[755,817,1365,1388]" italics="true" pageId="51" pageNumber="52">Piper</emphasis>
</taxonomicName>
species occur in lowland rainforests from parts of
<collectingCountry id="ABE819F4FFB34A78FF328E18FEF9EACB" box="[151,269,1399,1424]" name="Nicaragua" pageId="51" pageNumber="52">Nicaragua</collectingCountry>
, across all of
<collectingCountry id="ABE819F4FFB34A78FE0B8E17FDD8EACB" box="[430,556,1399,1424]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
, and into parts of adjacent
<collectingCountry id="ABE819F4FFB34A78FCC08E17FC35EAD4" box="[869,961,1400,1423]" name="Panama" pageId="51" pageNumber="52">Panama</collectingCountry>
. The evolution of the plants involved has received attention (
<bibRefCitation id="B76E2495FFB34A78FEC88EF3FDD3EAEF" author="Tepe, E. &amp; Vincent, M. &amp; Watson, L." box="[365,551,1436,1461]" pageId="51" pageNumber="52" pagination="156 - 178" refId="ref52546" refString="Tepe, E., Vincent, M. &amp; Watson, L. (2004) Phylogenetic patterns, evolutionary trends, and the origin of ant-plant associations in Piper section Macrostachys: Burger's hypothesis revisited. In: L. A. Dyer &amp; A. D. N. Palmer (Eds), Piper: a model genus for studies of phytochemistry, ecology, and evolution. Kluwer Academic Publishing, New York, pp. 156 - 178. https: // doi. org / 10.1007 / 978 - 0 - 387 - 30599 - 8 _ 9" type="book chapter" year="2004">
Tepe
<emphasis id="E18B8576FFB34A78FE098EF2FE10EAEE" box="[428,484,1436,1461]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
2004
</bibRefCitation>
), as have the ecological relationships of ants and plants (
<bibRefCitation id="B76E2495FFB34A78FB058EF3FA90EAEF" author="Letourneau, D. K." box="[1184,1380,1436,1460]" pageId="51" pageNumber="52" pagination="122 - 126" refId="ref51627" refString="Letourneau, D. K. (1983) Passive aggression: an alternative hypothesis for the Piper-Pheidole association. Oecologia, 60, 122 - 126. https: // doi. org / 10.1007 / BF 00379331" type="journal article" year="1983">Letourneau 1983</bibRefCitation>
), but the taxonomy of the ants has been neglected. The ant involved has been assumed to be a single species,
<taxonomicName id="14FF22E7FFB34A78FABB8EAEFA66EA83" authorityName="Forel" authorityYear="1899" box="[1310,1426,1471,1496]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="species" species="bicornis">
<emphasis id="E18B8576FFB34A78FABB8EAEFA66EA83" box="[1310,1426,1471,1496]" italics="true" pageId="51" pageNumber="52">P. bicornis</emphasis>
</taxonomicName>
. However, there are morphological discontinuities that suggest a clade of three related species.
</paragraph>
<paragraph id="D3405964FFB34A78FF628D68FA94E98F" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
The types of
<taxonomicName id="14FF22E7FFB34A78FEC68D66FE2EE97B" authorityName="Forel" authorityYear="1899" box="[355,474,1543,1568]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="species" species="bicornis">
<emphasis id="E18B8576FFB34A78FEC68D66FE2EE97B" box="[355,474,1543,1568]" italics="true" pageId="51" pageNumber="52">P. bicornis</emphasis>
</taxonomicName>
, which are from far southwestern
<collectingCountry id="ABE819F4FFB34A78FCD78D67FC3AE944" box="[882,974,1544,1567]" name="Panama" pageId="51" pageNumber="52">Panama</collectingCountry>
, and almost all the collections from the adjacent Pacific lowlands of
<collectingCountry id="ABE819F4FFB34A78FE458D43FDAFE91E" box="[480,603,1580,1605]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
have major workers with relatively broad heads (CI 80-83; n=2). All the collections from the Atlantic lowlands of
<collectingCountry id="ABE819F4FFB34A78FDCD8D3FFD16E933" box="[616,738,1615,1640]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
and
<collectingCountry id="ABE819F4FFB34A78FCBD8D20FC7FE933" box="[792,907,1615,1640]" name="Nicaragua" pageId="51" pageNumber="52">Nicaragua</collectingCountry>
have noticeably more narrow heads (CI 72-76; n=5). Both of these species have the minor workers with a smooth, shiny face. Disjunctions of insect clades across the mountains of
<collectingCountry id="ABE819F4FFB34A78FED28DF7FE0FE9EB" box="[375,507,1687,1712]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
are repeatedly being shown to be relatively old, with varying degrees of morphological divergence and at a level of genetic and morphological separation that warrants species status.
</paragraph>
<paragraph id="D3405964FFB34A78FF628D8FFC74E8D3" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
An unexpected discovery was a third species, described here as
<taxonomicName id="14FF22E7FFB34A78FC098D8EFB98E9A3" authorityName="Longino" authorityYear="2019" box="[940,1132,1759,1784]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="51" pageNumber="52" phylum="Arthropoda" rank="species" species="bicornisculpta">
<emphasis id="E18B8576FFB34A78FC098D8EFB98E9A3" box="[940,1132,1759,1784]" italics="true" pageId="51" pageNumber="52">P. bicornisculpta</emphasis>
</taxonomicName>
, from an ant
<taxonomicName id="14FF22E7FFB34A78FAA98D8EFABEE9A3" box="[1292,1354,1761,1784]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="genus">
<emphasis id="E18B8576FFB34A78FAA98D8EFABEE9A3" box="[1292,1354,1761,1784]" italics="true" pageId="51" pageNumber="52">Piper</emphasis>
</taxonomicName>
on the Pacific slope of
<collectingCountry id="ABE819F4FFB34A78FEF78C6BFE24E846" box="[338,464,1796,1821]" name="Costa Rica" pageId="51" pageNumber="52">Costa Rica</collectingCountry>
, very near populations of true
<taxonomicName id="14FF22E7FFB34A78FC948C6AFC53E846" authorityName="Forel" authorityYear="1899" box="[817,935,1796,1821]" class="Magnoliopsida" family="Piperaceae" genus="Piper" kingdom="Plantae" order="Piperales" pageId="51" pageNumber="52" phylum="Tracheophyta" rank="species" species="bicornis">
<emphasis id="E18B8576FFB34A78FC948C6AFC53E846" box="[817,935,1796,1821]" italics="true" pageId="51" pageNumber="52">P. bicornis</emphasis>
</taxonomicName>
. This species has the major worker smaller overall (HW 0.92 vs.&gt; 1.00), intermediate with respect to head shape (CI 78), and with a striking difference in the head shape and sculpture of the minor worker. The face of the minor is fully foveolate, overlain with faint rugulae. Additional differentiating characters are in the species description.
</paragraph>
</subSubSection>
<subSubSection id="9BE50AEFFFB34A78FF628CFBFE9BE88B" pageId="51" pageNumber="52" type="etymology">
<paragraph id="D3405964FFB34A78FF628CFBFE9BE88B" blockId="51.[151,1437,428,2000]" pageId="51" pageNumber="52">
<emphasis id="E18B8576FFB34A78FF628CFBFEBCE8F6" bold="true" box="[199,328,1940,1965]" pageId="51" pageNumber="52">Etymology</emphasis>
: In reference to the &quot;passive aggressive&quot; nature of host plant defense by this specialized plant ant (
<bibRefCitation id="B76E2495FFB34A78FF3B8CD7FE97E88B" author="Letourneau, D. K." box="[158,355,1976,2000]" pageId="51" pageNumber="52" pagination="122 - 126" refId="ref51627" refString="Letourneau, D. K. (1983) Passive aggression: an alternative hypothesis for the Piper-Pheidole association. Oecologia, 60, 122 - 126. https: // doi. org / 10.1007 / BF 00379331" type="journal article" year="1983">Letourneau 1983</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>