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<mods:title id="5E949C42ACAA122A1289BEA97FA8D186">Cladistic analysis of Subfamily Bruchomyiinae (Diptera: Psychodidae)</mods:title>
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<paragraph id="9A0C36CAFFDEFFD3FF23F8ABE19BF8C1" blockId="17.[151,422,1879,1905]" box="[151,422,1879,1905]" pageId="17" pageNumber="168">
<heading id="C14481A6FFDEFFD3FF23F8ABE19BF8C1" bold="true" box="[151,422,1879,1905]" fontSize="11" level="1" pageId="17" pageNumber="168" reason="1">
<emphasis id="A8C7EAD8FFDEFFD3FF23F8ABE19BF8C1" bold="true" box="[151,422,1879,1905]" pageId="17" pageNumber="168">
Fossil
<taxonomicName id="5DB34D49FFDEFFD3FF52F8ABE19BF8C1" ID-CoL="87CDW" ID-ENA="43819" box="[230,422,1879,1905]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="17" pageNumber="168" phylum="Arthropoda" rank="subFamily" subFamily="Bruchomyiinae">Bruchomyiinae</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="9A0C36CAFFDEFFD0FF23F862E12FFEAD" blockId="17.[151,1436,1950,2011]" lastBlockId="18.[151,1437,151,285]" lastPageId="18" lastPageNumber="169" pageId="17" pageNumber="168">
Fossil
<taxonomicName id="5DB34D49FFDEFFD3FF56F862E1B2F807" box="[226,399,1950,1975]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="17" pageNumber="168" phylum="Arthropoda" rank="subFamily" subFamily="Bruchomyiinae">Bruchomyiinae</taxonomicName>
are documented from various deposits of Caribbean (1220 million years b.p.), Baltic (40 60 million years b.p.), and Burmese (100 million years b.p.) amber. To date the group is missing in geologically older strata,
<emphasis id="A8C7EAD8FFDDFFD0FE96FF64E176FF1F" box="[290,331,152,175]" italics="true" pageId="18" pageNumber="169">e.g.</emphasis>
Lebanese amber. Psychodids in the mentioned deposits were identified as closely related to extant Phlebotominae, Trichomyinae,
<taxonomicName id="5DB34D49FFDDFFD0FE43FF40E2BDFF65" box="[503,640,188,213]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="subFamily" subFamily="Sycoracinae">Sycoracinae</taxonomicName>
and probably
<taxonomicName id="5DB34D49FFDDFFD0FC96FF40E38EFF65" box="[802,947,188,213]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="subFamily" subFamily="Psychodinae">Psychodinae</taxonomicName>
. The absence of
<taxonomicName id="5DB34D49FFDDFFD0FBDBFF40E521FF65" box="[1135,1308,188,213]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="subFamily" subFamily="Bruchomyiinae">Bruchomyiinae</taxonomicName>
in deposits older than 100 million years b.p. may be incidental, but it remains remarkable as long as specimens are not confirmed.
</paragraph>
<caption id="CECC6642FFDDFFD0FF23FB3FE105FAA6" httpUri="https://zenodo.org/record/260843/files/figure.png" pageId="18" pageNumber="169" targetBox="[317,1270,353,1173]" targetPageId="18">
<paragraph id="9A0C36CAFFDDFFD0FF23FB3FE105FAA6" blockId="18.[151,1436,1219,1302]" pageId="18" pageNumber="169">
<emphasis id="A8C7EAD8FFDDFFD0FF23FB3FE12EFB69" bold="true" box="[151,275,1219,1241]" pageId="18" pageNumber="169">FIGURE 8.</emphasis>
Tree from Bayesian analysis of 52 species and 29 characters of extant
<taxonomicName id="5DB34D49FFDDFFD0FC5FFB3FE4B7FB69" box="[1003,1162,1219,1241]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="subFamily" subFamily="Bruchomyiinae">Bruchomyiinae</taxonomicName>
. Posterior probabilities are given. Abbreviations: B—
<taxonomicName id="5DB34D49FFDDFFD0FE1FFB1EE21AFB48" box="[427,551,1250,1272]" class="Insecta" family="Psychodidae" genus="Bruchomyia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="genus">
<emphasis id="A8C7EAD8FFDDFFD0FE1FFB1EE21AFB48" box="[427,551,1250,1272]" italics="true" pageId="18" pageNumber="169">Bruchomyia</emphasis>
</taxonomicName>
, Bo—
<taxonomicName id="5DB34D49FFDDFFD0FDD9FB1EE329FB48" box="[621,788,1250,1272]" class="Insecta" family="Psychodidae" genus="Boreofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="151" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="A8C7EAD8FFDDFFD0FDD9FB1EE329FB48" box="[621,788,1250,1272]" italics="true" pageId="18" pageNumber="169">Boreofairchildia</emphasis>
</taxonomicName>
, E—
<taxonomicName id="5DB34D49FFDDFFD0FCFAFB1FE3F5FB48" box="[846,968,1251,1272]" class="Insecta" family="Psychodidae" genus="Eutonnoiria" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="genus">
<emphasis id="A8C7EAD8FFDDFFD0FCFAFB1FE3F5FB48" box="[846,968,1251,1272]" italics="true" pageId="18" pageNumber="169">Eutonnoiria</emphasis>
</taxonomicName>
, L—
<taxonomicName id="5DB34D49FFDDFFD0FBB5FB1FE4A0FB48" box="[1025,1181,1251,1272]" class="Insecta" family="Psychodidae" genus="Laurenceomyia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="151" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="A8C7EAD8FFDDFFD0FBB5FB1FE4A0FB48" box="[1025,1181,1251,1272]" italics="true" pageId="18" pageNumber="169">Laurenceomyia</emphasis>
</taxonomicName>
, N—
<taxonomicName id="5DB34D49FFDDFFD0FB6EFB1EE565FB48" box="[1242,1368,1250,1272]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="genus">
<emphasis id="A8C7EAD8FFDDFFD0FB6EFB1EE565FB48" box="[1242,1368,1250,1272]" italics="true" pageId="18" pageNumber="169">Nemopalpus</emphasis>
</taxonomicName>
, Nt—
<taxonomicName id="5DB34D49FFDDFFD0FF23FAFCE10FFAA6" box="[151,306,1280,1302]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="151" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="A8C7EAD8FFDDFFD0FF23FAFCE10FFAA6" box="[151,306,1280,1302]" italics="true" pageId="18" pageNumber="169">Notofairchildia</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="9A0C36CAFFDDFFD0FF73FABFE257F885" blockId="18.[151,1437,1347,2031]" pageId="18" pageNumber="169">
Baltic amber fossils represent separate phylogenetic lines. The most abundant species in the deposits,
<taxonomicName id="5DB34D49FFDDFFD0FA36FAB8E1F6FA30" authority="Meunier, 1905" authorityName="Meunier" authorityYear="1905" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="tertiariae">
<emphasis id="A8C7EAD8FFDDFFD0FA36FAB8E13CFACF" italics="true" pageId="18" pageNumber="169">N. tertiariae</emphasis>
(Meunier, 1905)
</taxonomicName>
, has been repeatedly photographed and illustrated, so there is no dispute about its identity; however, the interpretation of its genitalia by Hennig (1972, p.4849) is incorrect. A crested area of weak sclerotization of the elongate gonostylus lead Hennig to the assumption the distal area was a segment of its own and gonocoxites were the hypandrium. Photos and figures of the second species,
<taxonomicName id="5DB34D49FFDDFFD0FB81FA28E5A1FA5C" authority="Edwards, 1921" authorityName="Edwards" authorityYear="1921" box="[1077,1436,1491,1516]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="molophilinus">
<emphasis id="A8C7EAD8FFDDFFD0FB81FA28E4D7FA5B" box="[1077,1258,1491,1515]" italics="true" pageId="18" pageNumber="169">N. molophilinus</emphasis>
Edwards, 1921
</taxonomicName>
provided by Wagner (2006) show evident differences from
<taxonomicName id="5DB34D49FFDDFFD0FCE2FA04E3D8F9BF" box="[854,997,1528,1551]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="tertiariae">
<emphasis id="A8C7EAD8FFDDFFD0FCE2FA04E3D8F9BF" box="[854,997,1528,1551]" italics="true" pageId="18" pageNumber="169">N. tertiariae</emphasis>
</taxonomicName>
,
<emphasis id="A8C7EAD8FFDDFFD0FC4CFA04E42EF9BF" box="[1016,1043,1528,1551]" italics="true" pageId="18" pageNumber="169">i.e</emphasis>
. short and stout gonocoxites and gonostyli, epandrium with two lateral, distally divided projections.
<taxonomicName id="5DB34D49FFDDFFD0FC28F9E0E4CBF984" authority="Wagner, 2012" authorityName="Wagner" authorityYear="2012" box="[924,1270,1564,1588]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="inexpectatus">
<emphasis id="A8C7EAD8FFDDFFD0FC28F9E0E476F983" box="[924,1099,1564,1587]" italics="true" pageId="18" pageNumber="169">N. inexpectatus</emphasis>
Wagner, 2012
</taxonomicName>
, similar in the basic structure of the genitalia to
<taxonomicName id="5DB34D49FFDDFFD0FDBCF9BCE287F9E7" box="[520,698,1599,1623]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="molophilinus">
<emphasis id="A8C7EAD8FFDDFFD0FDBCF9BCE287F9E7" box="[520,698,1599,1623]" italics="true" pageId="18" pageNumber="169">N. molophilinus</emphasis>
</taxonomicName>
, was already figured by Henning (1972, p.48, fig. 41); gonocoxite stout but gonostyli short and s-shaped, epandrium without lateral projection but with a pair of small trifid appendages. Genitalia of
<taxonomicName id="5DB34D49FFDDFFD0FE07F97BE376F910" authority="Wagner, 2006" authorityName="Wagner" authorityYear="2006" box="[435,843,1671,1696]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="hoffeinsi">
<emphasis id="A8C7EAD8FFDDFFD0FE07F97BE298F92F" box="[435,677,1671,1695]" italics="true" pageId="18" pageNumber="169">Nemopalpus hoffeinsi</emphasis>
Wagner, 2006
</taxonomicName>
, are basically different; gonocoxites and hypandrium are fused to a ventero-central complex, gonostyli inconspicuous. Epandrium well-developed, with conspicuous posterolateral prolongations. Wing venation, particularly length relation of R2+3 and R2, differs from most Old World
<taxonomicName id="5DB34D49FFDDFFD0FF50F90AE14DF8BE" box="[228,368,1782,1806]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="genus">
<emphasis id="A8C7EAD8FFDDFFD0FF50F90AE14DF8BE" box="[228,368,1782,1806]" italics="true" pageId="18" pageNumber="169">Nemopalpus</emphasis>
</taxonomicName>
; in
<taxonomicName id="5DB34D49FFDDFFD0FE2BF90BE26EF8BE" box="[415,595,1782,1806]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="molophilinus">
<emphasis id="A8C7EAD8FFDDFFD0FE2BF90BE26EF8BE" box="[415,595,1782,1806]" italics="true" pageId="18" pageNumber="169">N. molophilinus</emphasis>
</taxonomicName>
R2+3 is slightly longer than R2, in
<taxonomicName id="5DB34D49FFDDFFD0FC63F90BE45EF8BE" box="[983,1123,1783,1806]" class="Insecta" family="Psychodidae" genus="Notofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="tertiariae">
<emphasis id="A8C7EAD8FFDDFFD0FC63F90BE45EF8BE" box="[983,1123,1783,1806]" italics="true" pageId="18" pageNumber="169">N. tertiariae</emphasis>
</taxonomicName>
about 2 times longer. In
<taxonomicName id="5DB34D49FFDDFFD0FA36F90BE0CAF885" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="hoffeinsi">
<emphasis id="A8C7EAD8FFDDFFD0FA36F90BE0CAF885" italics="true" pageId="18" pageNumber="169">N. hoffeinsi</emphasis>
</taxonomicName>
, R2 is definitely longer than R2+3.
</paragraph>
<paragraph id="9A0C36CAFFDDFFD1FF73F8B8E45DFF67" blockId="18.[151,1437,1347,2031]" lastBlockId="19.[151,1436,151,359]" lastPageId="19" lastPageNumber="170" pageId="18" pageNumber="169">
<taxonomicName id="5DB34D49FFDDFFD0FF73F8B8E253F8EC" authority="Wagner, 2006" authorityName="Wagner" authorityYear="2006" box="[199,622,1860,1884]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="scheveni">
<emphasis id="A8C7EAD8FFDDFFD0FF73F8B8E182F8EC" box="[199,447,1860,1884]" italics="true" pageId="18" pageNumber="169">Nemopalpus scheveni</emphasis>
Wagner, 2006
</taxonomicName>
, from younger Caribbean amber, clearly belongs in
<taxonomicName id="5DB34D49FFDDFFD0FB68F8B8E5A8F8EC" box="[1244,1429,1860,1884]" class="Insecta" family="Psychodidae" genus="Boreofairchildia" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="151" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="A8C7EAD8FFDDFFD0FB68F8B8E5A8F8EC" box="[1244,1429,1860,1884]" italics="true" pageId="18" pageNumber="169">Boreofairchildia</emphasis>
</taxonomicName>
. Wing narrow, R2+3 more than 3 times longer than R2, r-m basal of medial fork, wing tip between R4 and R5, abdomen with lateral abdominal extensions with tufts of long setae; genitalia with elongate tergite 9, gonocoxite simple, gonostylus complicatedly built with 3 distal prolongations, and aedeagus flanked by pair of simple parameres.
<taxonomicName id="5DB34D49FFDDFFD0FEACF82AE2E1F85F" authority="Schluter, 1978" authorityName="Schluter" authorityYear="1978" box="[280,732,2006,2031]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="169" phylum="Arthropoda" rank="species" species="hennigianus">
<emphasis id="A8C7EAD8FFDDFFD0FEACF82AE209F85E" box="[280,564,2006,2030]" italics="true" pageId="18" pageNumber="169">Nemopalpus hennigianus</emphasis>
Schlüter, 1978
</taxonomicName>
was described based on a female. It is impossible to determine whether or not
<taxonomicName id="5DB34D49FFDCFFD1FEF7FF64E1F9FF1F" box="[323,452,151,175]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="170" phylum="Arthropoda" rank="species" species="scheveni">
<emphasis id="A8C7EAD8FFDCFFD1FEF7FF64E1F9FF1F" box="[323,452,151,175]" italics="true" pageId="19" pageNumber="170">N. scheveni</emphasis>
</taxonomicName>
is a synonymy of
<taxonomicName id="5DB34D49FFDCFFD1FD21FF64E37DFF1F" box="[661,832,151,175]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="170" phylum="Arthropoda" rank="species" species="hennigianus">
<emphasis id="A8C7EAD8FFDCFFD1FD21FF64E37DFF1F" box="[661,832,151,175]" italics="true" pageId="19" pageNumber="170">N. hennigianus</emphasis>
</taxonomicName>
; R2 is remarkably short, r-m basal of the medial fork, wing shape appears less slim as in other extant Neotropical species and in
<taxonomicName id="5DB34D49FFDCFFD1FC63FF43E465FF66" box="[983,1112,190,214]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="170" phylum="Arthropoda" rank="species" species="scheveni">
<emphasis id="A8C7EAD8FFDCFFD1FC63FF43E465FF66" box="[983,1112,190,214]" italics="true" pageId="19" pageNumber="170">N. scheveni</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="9A0C36CAFFDCFFD1FF73FF1EE4BAFED7" blockId="19.[151,1436,151,359]" pageId="19" pageNumber="170">
Recently another species was discovered from Burmese amber. It was impossible to identify wing venation of
<taxonomicName id="5DB34D49FFDCFFD1FF23FEFBE139FEAE" box="[151,260,262,286]" class="Insecta" family="Psychodidae" genus="Nemopalpus" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="170" phylum="Arthropoda" rank="species" species="velteni">
<emphasis id="A8C7EAD8FFDCFFD1FF23FEFBE139FEAE" box="[151,260,262,286]" italics="true" pageId="19" pageNumber="170">N. velteni</emphasis>
</taxonomicName>
, but the genitalia appear to be simple with tubular gonocoxites and gonostyli, subquadrate tergite 9 and soft oval cerci, aedeagus narrow, elongate and bifurcate. Findings of more species and specimens from Baltic and Caribbean Amber require an updated revision of fossil
<taxonomicName id="5DB34D49FFDCFFD1FCB4FEB2E390FED7" box="[768,941,334,359]" class="Insecta" family="Psychodidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="170" phylum="Arthropoda" rank="subFamily" subFamily="Bruchomyiinae">Bruchomyiinae</taxonomicName>
(Wagner, in prep.).
</paragraph>
</subSubSection>
</treatment>
</document>