<documentid="90D8766B01DC807E441A5F9E85B4DF15"ID-CLB-Dataset="33953"ID-DOI="http://dx.doi.org/10.3897/zookeys.315.5293"ID-GBIF-Dataset="eed3d9a5-c6b2-45ac-af9f-79770c35df88"ID-PMC="PMC3713351"ID-Pensoft-Pub="1313-2970-315-17"ID-PubMed="23878508"ModsDocAuthor=""ModsDocDate="2013"ModsDocID="1313-2970-315-17"ModsDocOrigin="ZooKeys 315"ModsDocTitle="Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)"checkinTime="1451247157087"checkinUser="pensoft"docAuthor="Ball, George E. & Shpeley, Danny"docDate="2013"docId="6F24DA781DC5CBA5B425CD81169C03CC"docLanguage="en"docName="ZooKeys 315: 17-54"docOrigin="ZooKeys 315"docSource="http://dx.doi.org/10.3897/zookeys.315.5293"docTitle="Coarazuphium Gnaspini, Vanin & Godoy 1998"docType="treatment"docVersion="5"lastPageNumber="32"masterDocId="0F6AFFB1B44A505EFFB3FF89734EE542"masterDocTitle="Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)"masterLastPageNumber="54"masterPageNumber="17"pageNumber="27"updateTime="1732797769636"updateUser="ExternalLinkService">
<mods:titleid="678E085C9068B978E499771B88557439">Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)</mods:title>
<taxonomicNameid="100D5A3F9E7650F528E2D208390AAB00"ID-CoL="8KW7G"LSID="http://species-id.net/wiki/Coarazuphium"authority="Gnaspini, Vanin & Godoy, 1998"authorityName="Gnaspini, Vanin & Godoy"authorityYear="1998"class="Insecta"family="Carabidae"genus="Coarazuphium"higherTaxonomySource="CoL"kingdom="Animalia"lsidName="Coarazuphium"order="Coleoptera"pageId="10"pageNumber="27"phylum="Arthropoda"rank="genus">Coarazuphium Gnaspini,Vanin & Godoy, 1998</taxonomicName>
<bibRefCitationid="81C4F1458E2A79673930307948393301"author="Godoy, NM"journalOrPublisher="Revista Brasiliera de Entomologia"pageId="24"pageNumber="41"pagination="795 - 799"title="Parazuphium tessai, sp. n., a new cavernicolous beetle from Bahia, Brazil (Coleoptera, Carabidae, Zuphiini)."volume="34"year="1990">Godoy and Vanin 1990</bibRefCitation>
<bibRefCitationid="31666EEA8578E8C96995A2A817BB2D1B"author="Mateu, J"journalOrPublisher="Journal of African Zoology"pageId="25"pageNumber="42"pagination="485 - 503"title="Contribution a la connaissance du genre Parazuphium Jeannel en Afrique noir (Coleoptera Carabidae)."volume="107"year="1993">Mateu 1993</bibRefCitation>
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<bibRefCitationid="2CF66773EF1A219816BCCE46ACD5FFD9"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="39 - 49"title="Ultrastructural analysis of Coarazuphium formoso (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle."volume="2866"year="2011 a">Pellegrini and Ferreira 2011a</bibRefCitation>
<bibRefCitationid="296F28E3483FB135B627BAB7B13091F8"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="47 - 58"title="Coarazuphium tapiaguassu (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle, with ultrastructural analysis and ecological considerations."volume="3116"year="2011 b">Pellegrini and Ferreira 2011b</bibRefCitation>
<bibRefCitationid="4B3967134EF4C855E35CE582511481C8"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. 1998</bibRefCitation>
<bibRefCitationid="BC2C6F57004E340F963DFFA7EC602BFC"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. (1998</bibRefCitation>
genus-group, restrictedas follows: size small; pronotum and dorsal surface of elytra moderately densely setose, setae decumbent; body markedly depressed; integument pale; head narrow, oviform, with posteriolateral margins broadly rounded; antennae elongate, antennomere 1 as long or longer than antennomeres 2-4; humeri narrowed, slightly or markedly so; metasternum short, metepisternum quadrate. Brachypterous or apterous. Male genitalia (Figs 10
<bibRefCitationid="52FE15954E604A0CAC1408E58A793760"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. (1998</bibRefCitation>
<bibRefCitationid="7B7B5E6C58F6A473D58B8AFD1DD861FE"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="47 - 58"title="Coarazuphium tapiaguassu (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle, with ultrastructural analysis and ecological considerations."volume="3116"year="2011 b">Pellegrini and Ferreira (2011b)</bibRefCitation>
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<bibRefCitationid="C058BAB945086371A124B375029B7E0B"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="47 - 58"title="Coarazuphium tapiaguassu (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle, with ultrastructural analysis and ecological considerations."volume="3116"year="2011 b">Pellegrini and Ferreira 2011b</bibRefCitation>
. Members of those species occupying the caves in limestone substrate are described as freely walking over the soil, and presumably resting exposed, rather than resting (hiding?) under rock cover. In contrast, all of the specimens collected of
<bibRefCitationid="F11107ED3D83DBCD48647E1A2F194A7A"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="39 - 49"title="Ultrastructural analysis of Coarazuphium formoso (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle."volume="2866"year="2011 a">Pellegrini and Ferreria (2011a</bibRefCitation>
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(lengthening of antennae and legs, depigmentation, micro- or anophthalmy, and reduction of elytral length, elytral humeri, metathorax and metathoracic wings) plus details of male and female genitalia are evolutionarily informative. Based on eye loss and more elongate appendages,
<bibRefCitationid="E4613B11C0000CAF7328F3BE5438EEF2"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. (1998</bibRefCitation>
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<bibRefCitationid="17291E69F7E369EC55E4418B6C909913"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. (1998</bibRefCitation>
inside the subterranean environments, which took place under the drier climate to which the region was in the past and is still submitted in the present. It is largely accepted in the literature that cave arthropods are related to litter epigean and/or endogean ancestors, which already inhabited humid habitats. Therefore,... ... the ancestral species [of
] should have been epigean and lived in forested (or at least humid) areas, and occurred at least in part of the region where the genus occurs nowadays. From [such ancestral stock] several lineages invaded the caves from the northern Bambui Speleological Province, where they became isolated with the progressive shrinkage of humid environments. Thence, the origin of the genus takes back to a time when the area was not drying yet, which is probably the Tertiary".
. First, the marked geographical range extension (from southeastern South America to the southern part of the North American continent) shows that this genus was not confined to eastern Brazil. Further, the distribution pattern lends support to the hypothesis of Gnaspini et. al. (see above) that
originated in early Tertiary time (i.e., before the beginning of the drying trend that extended through much of the Cenozoic era). Second, the extra-speleal humid forest existence of
(microphthalmy, brachyptery, depigmentation, etc.) evolved in surface habitats, though probably in forested deep leaf litter locations, and were in effect preadaptations for cave life.
<bibRefCitationid="3DB9198E7A855E12C0A742787817EF3E"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. 1998</bibRefCitation>
<bibRefCitationid="5E1B5AA2C4A3B652881712434DC0F03A"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. 1998</bibRefCitation>
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<bibRefCitationid="0D07C750EE2B84C5BE74786FE93541C6"author="Pellegrini, TG"journalOrPublisher="Zootaxa"pageId="26"pageNumber="43"pagination="39 - 49"title="Ultrastructural analysis of Coarazuphium formoso (Coleoptera: Carabidae: Zuphiini), a new Brazilian troglobitic beetle."volume="2866"year="2011 a">Pellegrini and Ferreira 2011a</bibRefCitation>
<bibRefCitationid="337F1C4C2FC1C2CE7A024AE9DE68654F"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. 1998</bibRefCitation>
<bibRefCitationid="083A38FB86D7CEA2C0642BC53F134C1B"author="Gnaspini, P"journalOrPublisher="Papeis Avulsos de Zoologia Museu de Zoologia da Universidade de Sao Paulo"pageId="24"pageNumber="41"pagination="297 - 309"title="A new genus of troglobitic carabid beetle from Brazil (Coleoptera: Carabidae: Zuphiini)."volume="40"year="1998">Gnaspini et al. 1998</bibRefCitation>
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<bibRefCitationid="592E8FCF85D436CC90F77C1DE301AA0C"author="Godoy, NM"journalOrPublisher="Revista Brasiliera de Entomologia"pageId="24"pageNumber="41"pagination="795 - 799"title="Parazuphium tessai, sp. n., a new cavernicolous beetle from Bahia, Brazil (Coleoptera, Carabidae, Zuphiini)."volume="34"year="1990">Godoy and Vanin 1990</bibRefCitation>
