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<mods:title id="FF2418F80D24B5947B8AE6CB8A43E771">The first fossil spider cricket (Orthoptera: Gryllidae: Phalangopsinae): 20 million years of troglobiomorphosis or exaptation in the dark?</mods:title>
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<mods:namePart id="9FC5977B76002BCCB61A4C87FF63A3FE">Heads, Sam W.</mods:namePart>
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<subSubSection id="70D2653EFFCF8D0229B0C8BE2C4EFCE7" pageId="2" pageNumber="58" type="nomenclature">
<paragraph id="387736B5FFCF8D0229B0C8BE2C62FCBF" blockId="2.[214,728,739,792]" pageId="2" pageNumber="58">
<heading id="633F81D9FFCF8D0229B0C8BE2C62FCBF" centered="true" fontSize="9" level="2" pageId="2" pageNumber="58" reason="4">
<emphasis id="0ABCEAA7FFCF8D0229B0C8BE2F37FD50" bold="true" box="[214,333,739,761]" pageId="2" pageNumber="58">SUBTRIBE</emphasis>
<taxonomicName id="FFC84D36FFCF8D022830C8BE2CA2FD5F" ID-CoL="8PS5G" authority="HUBBELL, 1938" authorityName="HUBBELL" authorityYear="1938" box="[342,728,739,761]" class="Insecta" family="Phalangopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="subTribe" subTribe="Amphiacustina">
<emphasis id="0ABCEAA7FFCF8D022830C8BE2C60FD5F" bold="true" box="[342,538,739,760]" pageId="2" pageNumber="58">AMPHIACUSTINA</emphasis>
<bibRefCitation id="5C594B44FFCF8D022B42C8B92CA2FD5F" author="Hubbell TH" box="[548,728,739,761]" pageId="2" pageNumber="58" pagination="191 - 233" refId="ref6412" refString="Hubbell TH. 1938. New cave-crickets from Yucatan, with a review of the Pentacentrinae, and studies on the genus Amphiacusta (Orthoptera: Gryllidae). Carnegie Institute of Washington Publication 491: 191 - 233." type="journal article" year="1938">HUBBELL, 1938</bibRefCitation>
</taxonomicName>
<emphasis id="0ABCEAA7FFCF8D0228F0C95A2C62FCBF" bold="true" box="[406,536,771,792]" pageId="2" pageNumber="58">STAT. NOV.</emphasis>
</heading>
</paragraph>
<paragraph id="387736B5FFCF8D0229C2C9772C4EFCE7" blockId="2.[164,779,810,1903]" box="[164,564,810,832]" pageId="2" pageNumber="58">
Amphiacustae
<bibRefCitation id="5C594B44FFCF8D022837C9772C4AFCE7" author="Hubbell TH" box="[337,560,810,832]" pageId="2" pageNumber="58" pagination="191 - 233" refId="ref6412" refString="Hubbell TH. 1938. New cave-crickets from Yucatan, with a review of the Pentacentrinae, and studies on the genus Amphiacusta (Orthoptera: Gryllidae). Carnegie Institute of Washington Publication 491: 191 - 233." type="journal article" year="1938">Hubbell, 1938: 206</bibRefCitation>
.
</paragraph>
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<paragraph id="387736B5FFCF8D0229C2C9142CF6FCF9" blockId="2.[164,779,810,1903]" box="[164,652,840,862]" pageId="2" pageNumber="58">
<emphasis id="0ABCEAA7FFCF8D0229C2C9142F50FCF9" box="[164,298,841,862]" italics="true" pageId="2" pageNumber="58">
<typeStatus id="E7738817FFCF8D0229C2C9142EA2FCF9" box="[164,216,841,862]" pageId="2" pageNumber="58">Type</typeStatus>
genus:
</emphasis>
<taxonomicName id="FFC84D36FFCF8D022855C9152CF2FCF9" authority="Saussure, 1874" authorityName="Saussure" authorityYear="1874" box="[307,648,840,862]" class="Insecta" family="Haglotettigoniidae" genus="Amphiacusta" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D022855C9152FB3FCFA" box="[307,457,840,861]" italics="true" pageId="2" pageNumber="58">Amphiacusta</emphasis>
<bibRefCitation id="5C594B44FFCF8D0228B2C9142CF2FCF9" author="de Saussure H" box="[468,648,841,862]" pageId="2" pageNumber="58" refId="ref6855" refString="de Saussure H. 1874. Mission scientifique au Mexique et dans l'Amerique centrale. Sixieme partie. Etudes sur les Myriapodes et les Insectes Orthopteres. Geneva: Imprimerie Imperiale." type="book" year="1874">Saussure, 1874</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="387736B5FFCF8D0229C2C93A2C90FAAC" blockId="2.[164,779,810,1903]" pageId="2" pageNumber="58">
<emphasis id="0ABCEAA7FFCF8D0229C2C93A2F75FCDB" box="[164,271,871,892]" italics="true" pageId="2" pageNumber="58">Remarks:</emphasis>
The priority of family-group names based on the
<typeStatus id="E7738817FFCF8D02299FC9DB2F50FC3C" box="[249,298,902,923]" pageId="2" pageNumber="58">type</typeStatus>
genus
<taxonomicName id="FFC84D36FFCF8D0228E4C9DB2C62FC3C" authorityName="Saussure" authorityYear="1874" box="[386,536,902,923]" class="Insecta" family="Haglotettigoniidae" genus="Amphiacusta" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D0228E4C9DB2C62FC3C" box="[386,536,902,923]" italics="true" pageId="2" pageNumber="58">Amphiacusta</emphasis>
</taxonomicName>
dates from Amphiacustae
<bibRefCitation id="5C594B44FFCF8D02299CC9F92FD8FC1E" author="Hubbell TH" box="[250,418,932,954]" pageId="2" pageNumber="58" pagination="191 - 233" refId="ref6412" refString="Hubbell TH. 1938. New cave-crickets from Yucatan, with a review of the Pentacentrinae, and studies on the genus Amphiacusta (Orthoptera: Gryllidae). Carnegie Institute of Washington Publication 491: 191 - 233." type="journal article" year="1938">Hubbell, 1938</bibRefCitation>
. This name has subsequently been used to refer to a group of genera comprising
<taxonomicName id="FFC84D36FFCF8D0229C2C9BC2F40FC51" authorityName="Saussure" authorityYear="1874" box="[164,314,993,1014]" class="Insecta" family="Haglotettigoniidae" genus="Amphiacusta" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D0229C2C9BC2F40FC51" box="[164,314,993,1014]" italics="true" pageId="2" pageNumber="58">Amphiacusta</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D02282EC9BC2F9AFC51" authorityName="Desutter-Grandcolas &amp; Hubbell" authorityYear="1993" box="[328,480,993,1014]" class="Insecta" family="Haglotettigoniidae" genus="Arachnopsita" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D02282EC9BC2F9AFC51" box="[328,480,993,1014]" italics="true" pageId="2" pageNumber="58">Arachnopsita</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D022888C9BC2C1FFC51" authorityName="Desutter-Grandcolas" authorityYear="1994" box="[494,613,993,1014]" class="Insecta" family="Haglotettigoniidae" genus="Cantrallia" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D022888C9BC2C1FFC51" box="[494,613,993,1014]" italics="true" pageId="2" pageNumber="58">Cantrallia</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D022B12C9BC2D7EFC51" authorityName="Desutter-Grandcolas" authorityYear="1994" box="[628,772,993,1014]" class="Insecta" family="Haglotettigoniidae" genus="Leptopedetes" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D022B12C9BC2D7EFC51" box="[628,772,993,1014]" italics="true" pageId="2" pageNumber="58">Leptopedetes</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D0229C2CE5C2F59FBB1" authorityName="Desutter-Grandcolas &amp; Hubbell" authorityYear="1993" box="[164,291,1025,1046]" class="Insecta" family="Haglotettigoniidae" genus="Longuripes" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D0229C2CE5C2F59FBB1" box="[164,291,1025,1046]" italics="true" pageId="2" pageNumber="58">Longuripes</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D022859CE5D2FB4FBB2" authorityName="Desutter-Grandcolas &amp; Hubbell" authorityYear="1993" box="[319,462,1024,1045]" class="Insecta" family="Haglotettigoniidae" genus="Mayagryllus" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D022859CE5D2FB4FBB2" box="[319,462,1024,1045]" italics="true" pageId="2" pageNumber="58">Mayagryllus</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D02288CCE5C2CFCFBB1" authorityName="Desutter-Grandcolas &amp; Hubbell" authorityYear="1993" box="[490,646,1025,1046]" class="Insecta" family="Haglotettigoniidae" genus="Nemoricantor" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D02288CCE5C2CFCFBB1" box="[490,646,1025,1046]" italics="true" pageId="2" pageNumber="58">Nemoricantor</emphasis>
</taxonomicName>
,
<taxonomicName id="FFC84D36FFCF8D022BC4CE5C2D7EFBB1" authorityName="Desutter-Grandcolas &amp; Hubbell" authorityYear="1993" box="[674,772,1025,1046]" class="Insecta" family="Haglotettigoniidae" genus="Noctivox" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D022BC4CE5C2D7EFBB1" box="[674,772,1025,1046]" italics="true" pageId="2" pageNumber="58">Noctivox</emphasis>
</taxonomicName>
, and
<taxonomicName id="FFC84D36FFCF8D0229B0CE422F0FFB93" authorityName="Desutter-Grandcolas" authorityYear="1993" box="[214,373,1055,1076]" class="Insecta" family="Phalangopsidae" genus="Prolonguripes" higherTaxonomySource="CoL" kingdom="Animalia" order="Orthoptera" pageId="2" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="0ABCEAA7FFCF8D0229B0CE422F0FFB93" box="[214,373,1055,1076]" italics="true" pageId="2" pageNumber="58">Prolonguripes</emphasis>
</taxonomicName>
(
<bibRefCitation id="5C594B44FFCF8D0228E2CE422CC3FB93" author="Desutter-Grandcolas L" box="[388,697,1055,1077]" pageId="2" pageNumber="58" pagination="1 - 82" refId="ref5975" refString="Desutter-Grandcolas L. 1993. The cricket fauna of Chipanecan caves (Mexico), systematics, phylogeny and the evolution towards troglobitic life (Orthoptera, Grylloidea, Phalangopsidae, Luzarinae). International Journal of Speleology 22: 1 - 82." type="journal article" year="1993">Desutter-Grandcolas, 1993</bibRefCitation>
,
<bibRefCitation id="5C594B44FFCF8D022BA3CE422C85FB92" author="Desutter-Grandcolas L" box="[709,767,1055,1077]" pageId="2" pageNumber="58" pagination="1 - 19" refId="ref6090" refString="Desutter-Grandcolas L. 1999. Are troglobitic taxa troglobiomorphic? A tes using phylogenetic inference. International Journal of Speleology 26: 1 - 19." type="journal article" year="1999">1999</bibRefCitation>
). The monophyly of this group is well supported by the apical migration of sclerites in the epiphallic arms (termed the C-sclerites by
<bibRefCitation id="5C594B44FFCF8D022B71CE272EA1FB08" author="Desutter-Grandcolas L" pageId="2" pageNumber="58" pagination="1 - 82" refId="ref5975" refString="Desutter-Grandcolas L. 1993. The cricket fauna of Chipanecan caves (Mexico), systematics, phylogeny and the evolution towards troglobitic life (Orthoptera, Grylloidea, Phalangopsidae, Luzarinae). International Journal of Speleology 22: 1 - 82." type="journal article" year="1993">Desutter-Grandcolas, 1993</bibRefCitation>
) and the endophallic apodeme without a mediodorsal crest. The group includes some 84 species in the aforementioned genera, distributed throughout Central America and the West Indies.
</paragraph>
<paragraph id="387736B5FFCF8D0229DACF4E2DB9FE48" blockId="2.[164,779,810,1903]" lastBlockId="2.[826,1441,198,495]" pageId="2" pageNumber="58">
<bibRefCitation id="5C594B44FFCF8D0229DACF4E2F0AFA8D" author="Hubbell TH" box="[188,368,1299,1322]" pageId="2" pageNumber="58" pagination="191 - 233" refId="ref6412" refString="Hubbell TH. 1938. New cave-crickets from Yucatan, with a review of the Pentacentrinae, and studies on the genus Amphiacusta (Orthoptera: Gryllidae). Carnegie Institute of Washington Publication 491: 191 - 233." type="journal article" year="1938">Hubbell (1938)</bibRefCitation>
formed the name Amphiacustae using his standard suffix for family-group names beneath the rank of subfamily and although the group has been intensively studied since then (
<bibRefCitation id="5C594B44FFCF8D0229CDCFD32F90FA04" author="Desutter-Grandcolas L" box="[171,490,1422,1444]" pageId="2" pageNumber="58" pagination="1 - 82" refId="ref5975" refString="Desutter-Grandcolas L. 1993. The cricket fauna of Chipanecan caves (Mexico), systematics, phylogeny and the evolution towards troglobitic life (Orthoptera, Grylloidea, Phalangopsidae, Luzarinae). International Journal of Speleology 22: 1 - 82." type="journal article" year="1993">Desutter-Grandcolas, 1993</bibRefCitation>
, 1997, 1999; Desutter- Grandcolas &amp; Otte, 1997), it has never been assigned a rank. Thus,
<bibRefCitation id="5C594B44FFCF8D022820CF962C7BFA46" author="Hubbell TH" box="[326,513,1483,1505]" pageId="2" pageNumber="58" pagination="191 - 233" refId="ref6412" refString="Hubbell TH. 1938. New cave-crickets from Yucatan, with a review of the Pentacentrinae, and studies on the genus Amphiacusta (Orthoptera: Gryllidae). Carnegie Institute of Washington Publication 491: 191 - 233." type="journal article" year="1938">Hubbells (1938)</bibRefCitation>
original spelling of the name (using the suffix -ae) has never been questioned. The suffixes of family-group names are regulated by the International Code of Zoological Nomenclature (
<collectionCode id="5ED9AE70FFCF8D0229CDCC1B2E96F9FB" box="[171,236,1606,1628]" pageId="2" pageNumber="58">ICZN</collectionCode>
) at the ranks of superfamily, family, subfamily, tribe, and subtribe (
<collectionCode id="5ED9AE70FFCF8D0228FDCC392FA4F9DE" box="[411,478,1636,1657]" pageId="2" pageNumber="58">ICZN</collectionCode>
, 1999: article 29.2). The suffixes of names at other ranks within the family group are not formally regulated, but do have widely accepted standard suffixes (e.g. -oidae for an epifamily, -iti for a supertribe, -ita for an infratribe, etc.). The suffixes of rankless taxa are likewise unregulated by the
<collectionCode id="5ED9AE70FFCF8D02299ECD412F40F896" box="[248,314,1820,1841]" pageId="2" pageNumber="58">ICZN</collectionCode>
, and their use has become increasingly desirable in recent years because of the widespread use of cladistic methodology and the need to name important nodes within phylogenies; particularly in cases where the assignment of a specific rank would be superfluous. Nevertheless, the use of the name Amphiacustae as a rankless taxon retaining its original suffix is somewhat problematic because of the similarity of the name with that of the
<typeStatus id="E7738817FFCF8D022C7CCB022B31FED3" box="[1306,1355,351,372]" pageId="2" pageNumber="58">type</typeStatus>
genus. This has the potential to cause confusion, and so the group is here assigned the rank of subtribe and the suffix is emended to -ina, in accordance with
<collectionCode id="5ED9AE70FFCF8D022C04CBE62BDBFE77" box="[1378,1441,443,464]" pageId="2" pageNumber="58">ICZN</collectionCode>
regulations.
</paragraph>
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