treatments-xml/data/6F/24/DA/6F24DA781DC5CBA5B425CD81169C03CC.xml

<document 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. &amp; 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 &amp; Godoy 1998" docType="treatment" docVersion="3" 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="1668156099637" updateUser="ExternalLinkService">
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<mods:title>Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Ball, George E.</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Shpeley, Danny</mods:namePart>
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<mods:date>2013</mods:date>
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<mods:number>315</mods:number>
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<mods:start>17</mods:start>
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<mods:url>http://dx.doi.org/10.3897/zookeys.315.5293</mods:url>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.315.5293</mods:identifier>
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<treatment ID-GBIF-Taxon="152045953" LSID="urn:lsid:plazi:treatment:6F24DA781DC5CBA5B425CD81169C03CC" httpUri="http://treatment.plazi.org/id/6F24DA781DC5CBA5B425CD81169C03CC" lastPageId="16" lastPageNumber="32" pageId="10" pageNumber="27">
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<paragraph pageId="10" pageNumber="27">
<taxonomicName LSID="http://species-id.net/wiki/Coarazuphium" authority="Gnaspini, Vanin &amp; Godoy, 1998" authorityName="Gnaspini, Vanin &amp; 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 &amp; Godoy, 1998</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="27" type="reference_group">
<paragraph pageId="10" pageNumber="27">
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium" order="Coleoptera" pageId="10" pageNumber="27" phylum="Arthropoda" rank="genus">Parazuphium</taxonomicName>
; 
<bibRefCitation 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>
: 795 (not 
<bibRefCitation pageId="10" pageNumber="27">Jeannel 1942</bibRefCitation>
).
</paragraph>
<paragraph pageId="10" pageNumber="27">
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphium" order="Coleoptera" pageId="10" pageNumber="27" phylum="Arthropoda" rank="genus">Zuphium</taxonomicName>
; 
<bibRefCitation 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>
: 486 (not 
<bibRefCitation pageId="10" pageNumber="27">Latreille 1805</bibRefCitation>
).
</paragraph>
<paragraph pageId="10" pageNumber="27">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="10" pageNumber="27" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
<bibRefCitation author="Alvares, ESS" journalOrPublisher="Lundiana" pageId="23" pageNumber="40" pagination="41 - 43" title="Coarazuphium pains, a new species of troglobitic beetle from Brazil (Coleoptera: Carabidae: Zuphiini)." volume="3" year="2002">
Gnaspini, Vanin &amp; Godoy, 1998: 298. 
<normalizedToken originalValue="Álvares">Alvares</normalizedToken>
and Ferreira 2002
</bibRefCitation>
: 41. 
<bibRefCitation author="Lorenz, W" journalOrPublisher="Published by Author, Tutzing" pageId="25" pageNumber="42" title="Systematic List of extant Ground Beetles of the World (Insecta Coleoptera &quot; Geadephaga &quot;: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Ed. 2." year="2005">Lorenz 2005</bibRefCitation>
: 507. 
<bibRefCitation 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>
: 39. 
<bibRefCitation 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>
: 47.
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="27" type="type species">
<paragraph pageId="10" pageNumber="27">Type species.</paragraph>
<paragraph pageId="10" pageNumber="27">
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium tessai" order="Coleoptera" pageId="10" pageNumber="27" phylum="Arthropoda" rank="species" species="tessai">Parazuphium tessai</taxonomicName>
Godoy &amp; Vanin, 1990 (designated by 
<bibRefCitation 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>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="11" lastPageNumber="28" pageId="10" pageNumber="27" type="generic name">
<paragraph pageId="10" pageNumber="27">Generic name.</paragraph>
<paragraph lastPageId="11" lastPageNumber="28" pageId="10" pageNumber="27">
As explained by 
<bibRefCitation 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>
: 299), the name 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">
<pageBreakToken pageId="11" pageNumber="28" start="start">Coarazuphium</pageBreakToken>
</taxonomicName>
is a compound Latinized neuter noun derived from 
<normalizedToken originalValue="“coara”">&quot;coara&quot;</normalizedToken>
, meaning hole or cave in the Tupi language (Brazilian, native tongue), plus 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Zuphium</taxonomicName>
. The name refers to the troglophilic or troglobitic habits of the species of this genus, and their basic nominotypical 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Zuphium</taxonomicName>
-like features.
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="28" type="recognition">
<paragraph pageId="11" pageNumber="28">Recognition.</paragraph>
<paragraph pageId="11" pageNumber="28">
With character states of Western Hemisphere 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Zuphium</taxonomicName>
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 
<normalizedToken originalValue="A–">A-</normalizedToken>
10C): phallus without dorsal paraostial sclerites. Female genital tract (Fig. 12A, 12B): without secondary spermathecal gland.
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="28" type="description">
<paragraph pageId="11" pageNumber="28">Description.</paragraph>
<paragraph pageId="11" pageNumber="28">
None required here. See 
<bibRefCitation 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>
: 298-299), 
<bibRefCitation 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>
, and description of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium whiteheadi" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="whiteheadi">Coarazuphium whiteheadi</taxonomicName>
below.
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="28" type="geographical distribution">
<paragraph pageId="11" pageNumber="28">Geographical distribution.</paragraph>
<paragraph pageId="11" pageNumber="28">
Confined to the Neotropical Region, the seven species of this genus are known only from southeastern Brazil (
<bibRefCitation 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 Ferreria 2011b</bibRefCitation>
: 57, fig. 10) and southern Mexico (Map 1). This is an example of the &quot;Paleo-American distribution pattern&quot; (see 
<bibRefCitation author="Halffter, G" journalOrPublisher="Annual Review of Entomology" pageId="24" pageNumber="41" pagination="95 - 114" title="Biogeography of the montane entomofauna of Mexico and Central America." url="10.1146/annurev.en.32.010187.000523" volume="32" year="1987">Halffter (1987)</bibRefCitation>
and 
<bibRefCitation author="Liebherr, JK" journalOrPublisher="Canadian Entomologist" pageId="25" pageNumber="42" pagination="841 - 860" title="Biogeographic patterns of Montane Mexican and Central American Carabidae (Coleoptera)." url="10.4039/Ent126841-3" volume="126" year="1994">Liebherr (1994</bibRefCitation>
: 845)).
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="28" type="way of life">
<paragraph pageId="11" pageNumber="28">Way of life.</paragraph>
<paragraph pageId="11" pageNumber="28">
The previously described species of genus 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
inhabit caves in one of two types of substrate (
<bibRefCitation 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>
: 55): limestone, occupied by 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium bezerra" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="bezerra">Coarazuphium bezerra</taxonomicName>
Gnaspini, Vanin &amp; Godoy; 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium cessaima" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="cessaima">Coarazuphium cessaima</taxonomicName>
Gnaspini, Vanin &amp; Godoy; 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium formoso" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="formoso">Coarazuphium formoso</taxonomicName>
Pellegrini &amp; Ferreira; 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium pains" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="pains">Coarazuphium pains</taxonomicName>
<normalizedToken originalValue="Álvares">Alvares</normalizedToken>
&amp; Ferreira; and 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tessai" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tessai">Coarazuphium tessai</taxonomicName>
(Godoy &amp; Vanin); and iron ore, occupied by 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tapiaguassu" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tapiaguassu">Coarazuphium tapiaguassu</taxonomicName>
Pellegrini &amp; Ferreira. The caves in iron ore are described as 
<normalizedToken originalValue="“shallow”">&quot;shallow&quot;</normalizedToken>
. 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 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tapiaguassu" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tapiaguassu">Coarazuphium tapiaguassu</taxonomicName>
were found under rocks, on the cave floor. In contrast, known members of the Mexican species, 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium whiteheadi" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="whiteheadi">Coarazuphium whiteheadi</taxonomicName>
, new species, seem to be surface inhabitants (see below)
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="28" type="parasites">
<paragraph pageId="11" pageNumber="28">Parasites.</paragraph>
<paragraph pageId="11" pageNumber="28">
Some specimens of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tapiaguassu" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tapiaguassu">Coarazuphium tapiaguassu</taxonomicName>
were infested by laboulbenelian fungi. The fungi were not identified further. See this topic for 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium whiteheadi" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="whiteheadi">Coarazuphium whiteheadi</taxonomicName>
, below.
</paragraph>
</subSubSection>
<subSubSection lastPageId="16" lastPageNumber="33" pageId="11" pageNumber="28" type="evolutionary considerations">
<paragraph pageId="11" pageNumber="28">Evolutionary considerations.</paragraph>
<paragraph pageId="11" pageNumber="28">
The ultrastructural features (i.e., principally sensillar) observed with scanning electron microscopy by 
<bibRefCitation 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>
, 
<bibRefCitation 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">2011b</bibRefCitation>
) differ only slightly between the two species of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
that they studied, and between that genus and 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphioides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphioides" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Zuphioides</taxonomicName>
(cf. accompanying SEM illustrations of 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphioides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphioides mexicanum" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="mexicanum">Zuphioides mexicanum</taxonomicName>
). So, it seems to us unlikely that such features are or will be evolutionarily informative.
</paragraph>
<paragraph pageId="11" pageNumber="28">
In contrast, the standard structural troglobitic features of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
(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, 
<bibRefCitation 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>
: 303) proposed that 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium cessaima" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="cessaima">Coarazuphium cessaima</taxonomicName>
showed the more modified features, compared to 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tessai" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tessai">Coarazuphium tessai</taxonomicName>
and 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium bezerra" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="bezerra">Coarazuphium bezerra</taxonomicName>
, the only other species of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
known at that time. 
<bibRefCitation author="Alvares, ESS" journalOrPublisher="Lundiana" pageId="23" pageNumber="40" pagination="41 - 43" title="Coarazuphium pains, a new species of troglobitic beetle from Brazil (Coleoptera: Carabidae: Zuphiini)." volume="3" year="2002">
<normalizedToken originalValue="Álvares">Alvares</normalizedToken>
and Ferreira (2002
</bibRefCitation>
: 43) proposed that, based on the features noted above, their newly described 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium pains" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="pains">Coarazuphium pains</taxonomicName>
would occupy a position intermediate between 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium cessaima" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="cessaima">Coarazuphium cessaima</taxonomicName>
and 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tessai" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="tessai">Coarazuphium tessai</taxonomicName>
+ 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium bezerra" order="Coleoptera" pageId="11" pageNumber="28" phylum="Arthropoda" rank="species" species="bezerra">Coarazuphium bezerra</taxonomicName>
.
</paragraph>
<paragraph lastPageId="13" lastPageNumber="30" pageId="12" pageNumber="29">
<pageBreakToken pageId="12" pageNumber="29" start="start">The</pageBreakToken>
external features of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="genus">
<pageBreakToken pageId="13" pageNumber="30" start="start">Coarazuphium</pageBreakToken>
</taxonomicName>
evidently evolved in parallel with, and independent of, four other zuphiine troglobite taxa: the remarkable Spanish 
<taxonomicName class="Insecta" family="Carabidae" genus="Ildobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ildobates neboti" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="species" species="neboti">Ildobates neboti</taxonomicName>
<normalizedToken originalValue="Español">Espanol</normalizedToken>
, 1966; the Canary Islands 
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium feloi" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="species" species="feloi">Parazuphium feloi</taxonomicName>
Machado (1997: 163); the Australian Nullarbor 
<taxonomicName class="Insecta" family="Carabidae" genus="Speozuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Speozuphium poulteri" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="species" species="poulteri">Speozuphium poulteri</taxonomicName>
Moore (1995: 159) and 
<taxonomicName class="Insecta" family="Carabidae" genus="Speothalpius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Speothalpius grayi" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="species" species="grayi">Speothalpius grayi</taxonomicName>
Moore (1995: 160). Another Canary Islands 
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="genus">Parazuphium</taxonomicName>
(
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium damascenum subsp. canariense" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="subSpecies" species="damascenum" subSpecies="canariense">Parazuphium damascenum canariense</taxonomicName>
Machado, 1992: 580) exhibits these same reductive (though less developed) features, but it is evidently conspecific with the continental Palaearctic 
<taxonomicName class="Insecta" family="Carabidae" genus="Parazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Parazuphium damascenum subsp. damascenum" order="Coleoptera" pageId="13" pageNumber="30" phylum="Arthropoda" rank="subSpecies" species="damascenum" subSpecies="damascenum">Parazuphium damascenum damascenum</taxonomicName>
(Fairmaire, 1897). For a more detailed discussion of the matter see 
<bibRefCitation author="Machado, A" journalOrPublisher="Instituto de Estudios Canarias, La Laguna" pageId="25" pageNumber="42" title="Monografia de los Carabidos de las Islas Canarias." year="1992">Machado (1992</bibRefCitation>
: 581-582).
</paragraph>
<paragraph lastPageId="15" lastPageNumber="32" pageId="13" pageNumber="30">
<bibRefCitation 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>
: 308-309) proposed for the Brazilian species of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="14" pageNumber="31" phylum="Arthropoda" rank="genus">
<pageBreakToken pageId="14" pageNumber="31" start="start">Coarazuphium</pageBreakToken>
</taxonomicName>
that &quot;..... these highly derived troglobitic features are due to a long-term 
<pageBreakToken pageId="15" pageNumber="32" start="start">isolation</pageBreakToken>
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 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
] 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&quot;.
</paragraph>
<paragraph pageId="15" pageNumber="32">
Discovery of the Mexican species adds important details to the story of evolution of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
. 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 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
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 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium whiteheadi" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="species" species="whiteheadi">Coarazuphium whiteheadi</taxonomicName>
suggests that the basic troglobitic features of 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
(microphthalmy, brachyptery, depigmentation, etc.) evolved in surface habitats, though probably in forested deep leaf litter locations, and were in effect preadaptations for cave life.
</paragraph>
<paragraph pageId="15" pageNumber="32">
An additional observation relating to evolutionary history of the 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Zuphium</taxonomicName>
genus groupis that shared features of 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphioides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphioides" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Zuphioides</taxonomicName>
and 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
indicate that these two genera may be adelphotaxa, with 
<taxonomicName class="Insecta" family="Carabidae" genus="Zuphioides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuphioides" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Zuphioides</taxonomicName>
retaining mostly ancestral features including life in lowland hygrophilous or mesophilous situations.
</paragraph>
<paragraph pageId="15" pageNumber="32">
Key to species of genus 
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="genus">Coarazuphium</taxonomicName>
Gnaspini, Vanin &amp; Godoy
</paragraph>
<paragraph lastPageId="16" lastPageNumber="33" pageId="15" pageNumber="32">
<table lastPageId="16" lastPageNumber="33" pageId="15" pageNumber="32">
<tr pageId="15" pageNumber="32">
<td colspan="1" pageId="15" pageNumber="32" rowspan="1">
<bibRefCitation 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>
<bibRefCitation 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>
</td>
<td colspan="1" pageId="15" pageNumber="32" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium cessaima" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="species" species="cessaima">Coarazuphium cessaima</taxonomicName>
</td>
</tr>
<tr pageId="15" pageNumber="32">
<td colspan="1" pageId="15" pageNumber="32" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tapiaguassu" order="Coleoptera" pageId="15" pageNumber="32" phylum="Arthropoda" rank="species" species="tapiaguassu">Coarazuphium tapiaguassu</taxonomicName>
</td>
</tr>
<tr pageId="16" pageNumber="33">
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<bibRefCitation author="Alvares, ESS" journalOrPublisher="Lundiana" pageId="23" pageNumber="40" pagination="41 - 43" title="Coarazuphium pains, a new species of troglobitic beetle from Brazil (Coleoptera: Carabidae: Zuphiini)." volume="3" year="2002">
<pageBreakToken pageId="16" pageNumber="33" start="start">
<normalizedToken originalValue="Álvares">Alvares</normalizedToken>
</pageBreakToken>
and Ferreira 2002
</bibRefCitation>
<bibRefCitation author="Alvares, ESS" journalOrPublisher="Lundiana" pageId="23" pageNumber="40" pagination="41 - 43" title="Coarazuphium pains, a new species of troglobitic beetle from Brazil (Coleoptera: Carabidae: Zuphiini)." volume="3" year="2002">
<normalizedToken originalValue="Álvares">Alvares</normalizedToken>
and Ferreira 2002
</bibRefCitation>
</td>
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium pains" order="Coleoptera" pageId="16" pageNumber="33" phylum="Arthropoda" rank="species" species="pains">Coarazuphium pains</taxonomicName>
</td>
</tr>
<tr pageId="16" pageNumber="33">
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">Fig. 5AFig. 9BFig. 10C</td>
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium whiteheadi" order="Coleoptera" pageId="16" pageNumber="33" phylum="Arthropoda" rank="species" species="whiteheadi">Coarazuphium whiteheadi</taxonomicName>
</td>
</tr>
<tr pageId="16" pageNumber="33">
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<bibRefCitation 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>
</td>
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium formoso" order="Coleoptera" pageId="16" pageNumber="33" phylum="Arthropoda" rank="species" species="formoso">Coarazuphium formoso</taxonomicName>
</td>
</tr>
<tr pageId="16" pageNumber="33">
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<bibRefCitation 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>
<bibRefCitation 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>
</td>
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium bezerra" order="Coleoptera" pageId="16" pageNumber="33" phylum="Arthropoda" rank="species" species="bezerra">Coarazuphium bezerra</taxonomicName>
</td>
</tr>
<tr pageId="16" pageNumber="33">
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<bibRefCitation 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>
<bibRefCitation 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>
</td>
<td colspan="1" pageId="16" pageNumber="33" rowspan="1">
<taxonomicName class="Insecta" family="Carabidae" genus="Coarazuphium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coarazuphium tessai" order="Coleoptera" pageId="16" pageNumber="33" phylum="Arthropoda" rank="species" species="tessai">Coarazuphium tessai</taxonomicName>
</td>
</tr>
</table>
</paragraph>
</subSubSection>
</treatment>
</document>