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<document id="3D6CDFDF66821BCE519371F08D70F353" ENCODING="UTF-8" ID-GBIF-Dataset="59725ac5-691b-41a4-9c12-431eca55fdb5" ModsDocID="z00928p001" checkinTime="1247067629905" checkinUser="thomas" docAuthor="Frans B. M. Vermeulen &amp; Tomas Hrbek" docDate="2005" docId="880CB37765B6AF878EF22818F2824F5F" docLanguage="en" docName="2005_Vermeulen_Hrbek_gg1.xml" docOrigin="Zootaxa 928" docSource="http://www.zoobank.org/urn:lsid:zoobank.org:pub:784CB6FB-0BEB-4C7A-99A4-45C85AE8A5B7" docTitle="Kryptolebias sepia Vermeulen &amp; Hrbek, 2005, n. sp." docType="treatment" docUuid="D6AC62C8-15CA-4620-A15A-A0AE945E5E67" docUuidSource="ZooBank" docVersion="13" lastPageNumber="18" masterDocId="ECF1493A57CCA5FDB089B078E95D4529" masterDocTitle="Kryptolebias sepia n. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish from the Tapanahony River drainage in southeast Surinam." masterLastPageNumber="20" masterPageNumber="1" pageNumber="4" updateTime="1701314126560" updateUser="plazi">
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<mods:title id="4A4E3864E4F2768F478FA3FA1A37E1E8">Kryptolebias sepia n. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish from the Tapanahony River drainage in southeast Surinam.</mods:title>
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<mods:name id="C28B4341E3631FDFF44091F4048C49D7" type="personal">
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<mods:namePart id="779CCB9F1AA3BDA54E2DC330F62BB04F">Frans B. M. Vermeulen</mods:namePart>
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<mods:namePart id="8C5433C693C2274EC40EE94FEA60A08C">Tomas Hrbek</mods:namePart>
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<mods:typeOfResource id="42E7B529CA71D61C243C4FF3CAD1AA71">text</mods:typeOfResource>
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<mods:title id="071298529AE7C03461AEA4FB54229935">Zootaxa</mods:title>
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<mods:date id="22781ADF3F29F539126B2FD981791FE9">2005</mods:date>
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<mods:number id="860704BFB3EEB28C7E12820B9053F4ED">928</mods:number>
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<mods:location id="9029FED51576CE6CF91F7A774B0EF661">
<mods:url id="C53CA25EBC85E06021A87A32B3AE3C24">http://www.zoobank.org/urn:lsid:zoobank.org:pub:784CB6FB-0BEB-4C7A-99A4-45C85AE8A5B7</mods:url>
</mods:location>
<mods:classification id="7A4B4D623A454746E999ADC62925B306">journal article</mods:classification>
<mods:identifier id="6741CB941DCE415D2E121BDB141E01B8" type="Plazi-Custom">z00928p001</mods:identifier>
<mods:identifier id="DF6609F30B67472AB6EA24404846C141" type="ZooBank">784CB6FB-0BEB-4C7A-99A4-45C85AE8A5B7</mods:identifier>
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<treatment id="880CB37765B6AF878EF22818F2824F5F" ID-DOI="http://doi.org/10.5281/zenodo.6265245" ID-GBIF-Taxon="100124109" ID-Zenodo-Dep="6265245" LSID="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" httpUri="http://treatment.plazi.org/id/880CB37765B6AF878EF22818F2824F5F" lastPageNumber="18" pageNumber="4">
<subSubSection id="CB4BF3506A99F65924BA82E2FDFC6EBE" type="nomenclature">
<paragraph id="AC01D906F4AD7D329892CB6BBF783F04" pageNumber="4">
<taxonomicName id="41DBFA38675B98C48722CB29E9816162" ID-CoL="3RF29" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
,
<taxonomicNameLabel id="E10F572C19FC7C56E817E328FBBD2397" rank="species">n. sp.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="73842EC3186D385D94C49E5547248FF9" type="description">
<paragraph id="9B31075E7D2DD83BFC8AB12FB843DD2C" pageNumber="4">(Figs. 1, 2)</paragraph>
</subSubSection>
<subSubSection id="76C6ED9BCE04B025000187DCF07F6E76" type="nomenclature">
<paragraph id="219D2A0404202C2D2BEAC3A57758D5B8" pageNumber="4">
<materialsCitation id="A7F65F6BF5A4F56A54583D64C0A4D7CD" ID-GBIF-Occurrence="923893718" country="Surinam">
<typeStatus id="A1D0407925ACB8AA04F7ED67D81AF09A">Holotype</typeStatus>
-
<collectionCode id="5E585EE1459FAEACD612DCF8AA368CA2" collectionName="ZMA">ZMA</collectionCode>
123.714 male, 61.8 mm:
<collectingCountry id="E747EFDB726510D0A9FFE9980DFF04D2">Surinam</collectingCountry>
, Upper Marowijne system, Tapanahony River, 5 km downriver from Palumeu on the right bank, a hunting trail leads to a creek 15 minutes walk from the river;
<geoCoordinate id="FE45CBEDEF9217690C5919E624F34EB8" direction="north" orientation="latitude" value="3.3786113">03° 22’ 43”N</geoCoordinate>
<geoCoordinate id="2445B1FA11EE29747632162FF4DF89E9" direction="west" orientation="longitude" value="-55.411392">055° 24’ 41”W</geoCoordinate>
, coll. F.B.M. Vermeulen,
<date id="80AC4EBE3DF70245FDE7B2B4B5854AB0" day="14" month="07" value="2003-07-14" year="2003">14 July 2003</date>
. Station number SU 2003-02.
</materialsCitation>
</paragraph>
<paragraph id="47E4EE9128C5B169CCDC48CC0CBD8810" pageNumber="4">
<materialsCitation id="38442C1A8A75C1297F33E4F99996ED42" ID-GBIF-Occurrence="923893720" country="Surinam">
Allotype -
<collectionCode id="5A2A5ABD7E62ED071BCA9A54030EA831" collectionName="ZMA">ZMA</collectionCode>
123.715 female, 65.1 mm.
<collectingCountry id="3010068EF0D1C844DA1CC5DA298D9F46">Surinam</collectingCountry>
, Upper Marowijne system, Palumeu River 20 km upstream from the confluence of the Palumeu and Tapanahony Rivers, small creek 20 minute walk from riverbank,
<geoCoordinate id="5D84114A207DDDDF2B4B9D782267C0A7" direction="north" orientation="latitude" value="3.2308333">03° 13’ 51”N</geoCoordinate>
<geoCoordinate id="E4EDE2F6456D864C15A6331896582983" direction="west" orientation="longitude" value="-55.396114">055° 23’ 46” W</geoCoordinate>
coll. F.B.M. Vermeulen,
<date id="DCC15E96F8953EED77449B3E30785212" day="16" month="07" value="2003-07-16" year="2003">16 July 2003</date>
. Station number SU 2003-06.
</materialsCitation>
</paragraph>
<paragraph id="6817BD1DAF7B624794C0BDD565543615" country="Surinam " pageNumber="4">
<materialsCitation id="5BD4E419099D7A0B175BA5CA4F3E9C76" ID-GBIF-Occurrence="923893717" country="Surinam">
<typeStatus id="7E8AAA4AF4D904165EDEB8104C0C4D68">Paratypes</typeStatus>
-
<collectionCode id="7E226262376E80F73D19EA244D984492" collectionName="ZMA">ZMA</collectionCode>
123.716, 2 juveniles, 32.9-36.3 mm specimens collected by S. Sladkowsky and F.B.M. Vermeulen,
<date id="A6A6A864A8325B9979B6E31C9FAD749C" day="20" month="08" value="2000-08-20" year="2000">20 August 2000</date>
. Station number SVS 2000-02 (same collection locality as holotype).
</materialsCitation>
<materialsCitation id="2F345F873170D70551682271E95D0ABE" ID-GBIF-Occurrence="923893719" country="Surinam">
<collectionCode id="A8788F72A449FD9980283292C9AB3215" collectionName="ZMA">ZMA</collectionCode>
123.717, 6 juvenile 31.5-38.1 mm specimens collected by F.B.M. Vermeulen, same collection data as holotype.
</materialsCitation>
<materialsCitation id="748DDA4200A5C9772815142234D29B22" ID-GBIF-Occurrence="923893715" country="Surinam">
<collectionCode id="3E4C6DFA0EEF62E3A455C25C8107C8FE" collectionName="Suriname, Paramaraibo, University, National Zoological Collection of Suriname">NZCS</collectionCode>
F6418, (originally identified as
<taxonomicName id="9AB587B9F6B8D700CB089D26D9D2F8C4" LSID-ZBK="urn:lsid:zoobank.org:act:D1F0E8BA-67BE-4135-BBD9-9A4459817679" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus lanceolatus Eigenmann 1909:51" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="lanceolatus">Rivulus cf. lanceolatus Eigenmann, 1909</taxonomicName>
); Ulemari tributary, 13 km upstream of confluence with Litani River,
<geoCoordinate id="43B8406BE834F61E147578E1334FF217" direction="north" orientation="latitude" value="3.2230556">03° 13’ 23”N</geoCoordinate>
<geoCoordinate id="AC82533A1089779363F69515009CAA58" direction="west" orientation="longitude" value="-54.260555">054° 15’ 38”W</geoCoordinate>
, collected by P. E. Ouboter, S. Sahdew and S. Ramanand, 3-
<date id="49128D6F65FF9F8281FB265B71009B8A" day="05" month="04" value="1998-04-05" year="1998">5 April 1998</date>
.
</materialsCitation>
<materialsCitation id="924367686B8B2AC24B79F7E2C37B5BE7" ID-GBIF-Occurrence="923893716" country="Surinam">
<collectionCode id="3053B9B1AEDC9B5DB825F61481C0FE70" collectionName="Suriname, Paramaraibo, University, National Zoological Collection of Suriname">NZCS</collectionCode>
F7063-F7066, 4 juvenile specimens, same collection data as holotype.
</materialsCitation>
<materialsCitation id="A1200DA78C2AD1BD2C808A8F3559C0FA" ID-GBIF-Occurrence="923893721" country="Surinam">
<collectionCode id="C84B1B9291111C828C788659C629CC7A" collectionName="France, Paris, Museum National d'Histoire Naturelle">MNHN</collectionCode>
2004-3096, 1 specimen, same collection data as holotype.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="F93FCBDF98201B3140AF3A13D250C403" type="diagnosis">
<paragraph id="FD7729937135819480FB5A48F6A8CFD5" pageNumber="4">
Diagnosis -Distinguished from other
<taxonomicName id="71E5375A8D9D1EE4F805449C6CD4E929" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
and
<taxonomicName id="CCA094275BDCFCDD08026648AE52DB51" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
species by a combination of characters including low number of LL scales, high number of anal rays, short predorsal length and a robust body (Table 1), and unique mitochondrial DNA gene sequences.
<taxonomicName id="0D5B781AD59E56694651EF7ABD8C631B" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is further distinguished by remarkably strong melanistic pigmentation and the ability to change this coloration into several distinct patterns, lack of obvious sexual dimorphism, strong dichromatism between juveniles and adults, and absence of the “ Rivulus ” spot in females and juveniles.
</paragraph>
</subSubSection>
<subSubSection id="77B4452D84826DE65950085AC4415C90" type="description">
<paragraph id="C2C9D8C68C61CE9D2FF17F6463F81C57" lastPageNumber="5" pageNumber="4">
Description -A medium sized
<taxonomicName id="97129F2824270A518C131232417225B9" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
, robust shape, with a non-annual lifecycle. Morphometric data of
<typeStatus id="F54C1A939B581D5FAE961CF252529EB9">holotype</typeStatus>
, allotype and eight
<typeStatus id="5CF003A502A91B2B7EA79DB0ECD7A432">paratypes</typeStatus>
from different localities are given in Table 2 in millimeters and in Table 3 as percent of standard length. Meristic data are in Table 4. Males differ little from females in their body coloration, shape and fins. In adult males yellow marking on the gill cover is slightly more prominent and some but not all males show a few yellow markings along the lateral line. Frontal scalation pattern is in general E-type, but one individual showed mixed E-D-type (Hoedeman 1958). The strong melanism observed in
<taxonomicName id="737BA9C8262343F618A5B1A29420AE4C" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
is also observed in at least three other species of
<taxonomicName id="52C7498DA0E5189384846265C2A2A041" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
-
<taxonomicName id="AA34EDEA9198B73E5B249AEF55430C57" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="caudomarginatus">K. caudomarginatus</taxonomicName>
,
<taxonomicName id="9A6349E530E6393F93F93FCBC7803C6F" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="brasiliensis">K. brasiliensis</taxonomicName>
and
<taxonomicName id="6B85FFFA7B5A8B5B11A89F58018CD7C8" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="marmoratus">K. marmoratus</taxonomicName>
- and in
<taxonomicName id="ADC955EBDE4E793E89F80DE182DFC9DD" LSID-ZBK="urn:lsid:zoobank.org:act:3449F866-C045-4154-92A0-F9A956507FD5" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus atratus Garman 1895:140" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="atratus">R. atratus</taxonomicName>
. However,
<taxonomicName id="9716B16C3258AC0E8F87032186C3DCE4" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
is differentiated from these species by morphometric data, lower dorsal fin ray count (7.8 vs. 8.6 to 9.0), high anal fin ray count (15.1 vs. 11.2 to 12.6), by the absence of clear sexual dimorphism, by a network of dark markings beginning behind the eye and continuing just posterior of gill cover, by an oblique pattern on the lateral sides, and by their extraordinary capability to change color pattern within seconds. Diagnostic differences are given in Table 1.
<taxonomicName id="76FFDEB46C38AD1AB648DA2AABE838FA" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
also differs by a strong orange body coloration of juveniles, which otherwise is only known in juvenile
<taxonomicName id="5A0261FAAD503DB70BBF94A8E7FE38E7" LSID-ZBK="urn:lsid:zoobank.org:act:26EFD8AE-A63A-4E0C-9F60-02CCFCD1D95C" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus amphoreus Huber 1979:68" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="amphoreus">R. amphoreus</taxonomicName>
. With the co-occurring
<taxonomicName id="350554CA9C9F95395C044294D04B458F" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="urophthalmus">R. urophthalmus</taxonomicName>
/
<taxonomicName id="3220904A313A5614FC8BEF428053A903" LSID-ZBK="urn:lsid:zoobank.org:act:561FF4A2-6736-4ABA-80D0-5AE89957E232" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus stagnatus Eigenmann 1909:50" order="Cyprinodontiformes" pageNumber="4" phylum="Chordata" rank="species" species="stagnatus">R. stagnatus</taxonomicName>
group they share the lower number of dorsal rays and low number of LL scales but differ by a higher number of anal rays (15.1 vs. 12.3), by slightly shorter predorsal length to SL (76.6 % vs. 78.0 %), the absence of red spots forming lateral rows, and other diagnostic characters.
</paragraph>
<paragraph id="F7A0255557534B2215AD34D7A7728A3A" pageNumber="6">Adult males -See Figures 1 and 2. Body grayish, dark brown near the dorsum, pale white near the belly. Scales on lateral sides show dark pigments forming a network pattern, scale centers are brighter forming five to six light rows from post operculum to the end of peduncle. Eight to 10 oblique bars are shown in subadults and adults depending on their mood. These oblique markings become more permanent and intense after reaching full maturity. Some scales along the lateral line on the anterior part of body show bright yellow color in sexually active males. Head dark brown to reddish brown, in the suprapectoral region a bluish shine is only visible with striking light, cheek pale white without markings. On the operculum, posterior to the eye, a network of deep black blotches start on partly yellow background forming two to three bands. On the lateral sides posterior to the gill opening the black blotches continue in an irregular branched pattern. All fins rounded without any filaments or extensions. Caudal fin shows a wine-red hue that is more intense near the caudal base. Around the caudal fin base a large number of small speckles, mostly situated along the rays, form a reticulated pattern. These reticulations become more intense with age. A dark wine-red margin is often present. Dorsal transparent with reddish hue and irregular wine-red dots, some speckles near the base form a reticulated pattern as in caudal but more subdued. Anal iridescent green, near the base fading to bluish white with numerous fine wine-red speckles forming five to six diagonal lines, more compact towards the fin tip. A wine-red band is weakly visible near the anal fin margin. Ventral fins iridescent pale blue, no markings. Pectorals clear but orange towards the base; color becomes more intense with age. Eyes with a golden-brown iris, more golden when sexually active.</paragraph>
<paragraph id="025A638B18BC3E08A3F6D722868C8267" pageNumber="6">
Adult females -See Figures 3 and 4. Same coloration as in adult males. Body graybrown toward the dorsum and whitish gray to white toward the ventrum. Depending on the mood of the female, eight to 10 dark melanistic oblique bars may appear or disappear. Also depending on mood, head black to grayish black, dorsal portion being reddish brown. Chin whitish, operculum black or with black markings on a not very distinct yellow background. All fins rounded without any filaments and with the same colors and patterns as adult males. Eyes same as in males. The so-called “ Rivulus ” spot is normally present on the dorsal portion of the caudal peduncle at its junction with the caudal fin, and is a major identifying feature of
<taxonomicName id="6C35F03731EE77C2AA3A06D9E6B37CF9" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="6" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
females; however, it is not present in this species at any lifestage.
</paragraph>
<paragraph id="B801AE1D3C9CBC564D6A2E3A51D9D33D" pageNumber="6">Juveniles -Head and body same as in adults, but the pale white color of the belly is replaced by orange white. Lateral pattern depends on the mood of the individual; see Figures5, 6 and 7. During the night and when frightened, a fine oblique barred pattern appears following the lines of the scales. However, this pattern is also sometimes exhibited in other situations. Often pale golden blotches form an irregular pattern on the sides.</paragraph>
<paragraph id="7B917732D8C0D2A6D32E680F1682D4A3" pageNumber="6">Color in alcohol -Males and females: body brown to yellow brown with barred pattern subdued but visible; chin, lower half of gill cover, and abdomen whitish; operculum with dark markings; unpaired fins with fine brown spots and markings.</paragraph>
</subSubSection>
<subSubSection id="0118E92D7D33E96ABC44021ADF7E2CAD" type="biology_ecology">
<paragraph id="38C4C608D8C66A4773A4ABDEE8A2D773" lastPageNumber="12" pageNumber="11">
Reproduction -Eggs are 2 mm in diameter and have no major filaments on the cambium membrane, but are covered with an elastic and sticky substance. They are placed one by one between roots at the waters edge, or just above it. Under aquarium conditions,
<taxonomicName id="93ABA95AECDF0C164B6532D5198995E0" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="11" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
deposits more than 50% of its eggs above the waterline. This behavior is occasionally observed in rivuline species of the genus
<taxonomicName id="A0ED139E4A9B3543C067ECFF8F581281" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="11" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
(FBMV pers. obs.). Very low water levels stimulate this species to mate, with males aggressively driving females. Decreasing water level to less than 3 cm instantly result in spawning activity. Color of eggs is dark amber matching the color of the spawning substrate. No differences in egg development time were observed under the three incubation conditions. In fully aquatic environment, eggs develop and hatch in 14 to 16 days at 24°C. On both wet and moist peat moss, eggs develop normally within the 14 to 16 day period, but do not hatch. If not allowed to hatch by immersing them in water, the embryos die within the following next seven days. This species is therefore unlikely to lay eggs capable of undergoing a developmental diapause (Wourms 1972), and the three week period potentially sets the upper time limit for temporary habitat desiccation.
<taxonomicName id="86230C571B73FF573DC532E25E832802" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="11" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is also a good jumper and will, as many of its family members do (
<normalizedToken id="AC0EDC1EDE686CA5D90289E1959DFA4B" originalValue="Lüling">Lueling</normalizedToken>
1971, Huber 1992), seek new water bodies by moving overland through leaf litter if its habitat becomes unsuitable.
</paragraph>
<paragraph id="38909F8D02ABA116C0055FFFCA22D343" pageNumber="13">
Habitat -See Figure 8.
<taxonomicName id="2EDFB418C8D62F2351D07A08B850F434" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
appears to live only in extremely shallow parts of small creeks in hilly areas, and is restricted to primary forests with a very dense canopy cover. The actual habitat of
<taxonomicName id="4F043FBD139DC9E91B007512BEFACCD9" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
is found outside the main stream in swampy areas directly adjacent to the creeks themselves. The layer of leaf litter is often thick, sometimes over 50 cm, and is especially thick in the lower parts of creeks and adjacent flooded areas.
<taxonomicName id="0D9E54A51105ED652F0FFC7340674906" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is found in only a few centimeters of water which stands above this leaf layer, and over 95% of the population appears to consist of juveniles. In case of disturbance of the biotope,
<taxonomicName id="3CE071A0E52A4DD6975D950A31302602" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">K. sepia seeks</taxonomicName>
shelter in the soft bottom layer or jumps out of the water. The creeks proper had permanent running water. The water was clear, mineral poor, but with high concentrations of tannic acid, the so called black water type (Sioli 1984). No aquatic vegetation was present. Water temperatures were not always taken but likely never exceed 23ºC; pH values varied between 4.1 and 5.4, and GH and KH hardness were both &lt;0.5 DH.
</paragraph>
<paragraph id="3FE26089BF771F40D0F51EFB5E238075" lastPageNumber="14" pageNumber="13">
<taxonomicName id="E127B38C0374861692226EC3322C0C08" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is found syntopically with
<taxonomicName id="256F62536E07C20050BD9EB7053FA59F" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="urophthalmus">Rivulus aff. urophthalmus</taxonomicName>
and frequently with
<taxonomicName id="B3CE4814604BE3D44246F9E73D4A313F" LSID-ZBK="urn:lsid:zoobank.org:act:FBA3D378-1C49-443A-A0E2-E8F9EE0E5BE4" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus geayi Vaillant 1899:156 [3]" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="geayi">Rivulus aff. geayi</taxonomicName>
. On two occasions a third species of
<taxonomicName id="AC038382B8D81FA2CEBEA15ECA26F491" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
was found together with
<taxonomicName id="0414F9AAE4A20BF2AF434CD0C58FDF0C" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
. This species was identified as
<taxonomicName id="C62D7CA710495F1D3177C7A5402170C6" LSID-ZBK="urn:lsid:zoobank.org:act:36459AF1-9B00-41E9-8F26-8D8497124B36" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus holmiae Eigenmann 1909:50" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="holmiae">R. holmiae</taxonomicName>
by Hoedeman (1961) but moved to
<taxonomicName id="C7CB5745A300FBCC58A40B6A845630E3" LSID-ZBK="urn:lsid:zoobank.org:act:074615A2-A324-4E15-8AAA-345FBF2C308D" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus igneus Huber 1991:68" order="Cyprinodontiformes" pageNumber="13" phylum="Chordata" rank="species" species="igneus">R. igneus</taxonomicName>
by Huber (1991). This species, however, does not fit well with original descriptions of either species, thus its true identity needs further study. The only non-rivulid fish fauna included the rare
<taxonomicName id="B31D3E836BE75D9ED5A818A6588AE4AC" family="Lebiasinidae" genus="Pyrrhulina" higherTaxonomySource="CoL" kingdom="Animalia" order="Characiformes" pageNumber="13" phylum="Chordata" rank="species" species="laeta">Pyrrhulina laeta</taxonomicName>
and on one occasion
<taxonomicName id="7BB5E9FB2AF4B6F30EA0EF150905F84A" family="Auchenipteridae" genus="Tatia" higherTaxonomySource="CoL" kingdom="Animalia" order="Siluriformes" pageNumber="13" phylum="Chordata" rank="species" species="creutzbergi">Tatia aff. creutzbergi</taxonomicName>
. Besides these species, no other fish were found in the creek or surrounding leaf litter.
</paragraph>
<paragraph id="9775D42C5E5C50D0AF5F46424302CA23" pageNumber="14">
It is likely
<taxonomicName id="94CAE3AA7D82FDF87BA2E78909C2C9DC" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is a predator of small fishes and invertebrates. However, aquarium observations suggest they are not cannibalistic. The authors’ field observations suggest that in general all large and medium sized rivulids feed on tadpoles, ants and mosquito larvae and in many areas they probably contribute to controlling malaria outbreaks.
</paragraph>
</subSubSection>
<subSubSection id="E71DFE310A66B2195C0BCA7E4E9D9C01" type="distribution">
<paragraph id="0295C699229B8E29DEE154B4F51EB34B" pageNumber="14">
Distribution -
<taxonomicName id="201306B778C4F2DEA58F5DEA4C181093" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
was found only in creeks emptying into the Tapanahony and Palumeu Rivers and in the Ulemari River area (
<collectionCode id="F82FC8DDDA2C9AC09DFFFFE23349FE4C" collectionName="Suriname, Paramaraibo, University, National Zoological Collection of Suriname">NZCS</collectionCode>
F 6418-fish collected by P. E. Ouboter, S. Sahdew and S. Ramanand in
<date id="CCEEA6878955F666F2C7C2033CF5F6EF" month="04" value="1998-04" year="1998">April 1998</date>
). Additional collections are needed, including the upper Litani and Marowini River basins, to confirm the authors hypothesis that
<taxonomicName id="427D8616DF8B6A5277F1D99F44CC1A60" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
is wide spread in southeastern Surinam and southwestern French Guiana.
</paragraph>
</subSubSection>
<subSubSection id="563F4E9ECF2B79C5F6554701F109BD20" type="etymology">
<paragraph id="841EA01B8E6EAC832354484E11534500" pageNumber="14">Etymology -The name sepia alludes to the family Sepiidae, the cuttlefish, whose chromatophores also can cause rapid changes in color.</paragraph>
</subSubSection>
<subSubSection id="104413A5E83968F73E5C028ABC11A26A" type="discussion">
<paragraph id="5E97AFF1969D60D0A80D26A466BD0A2A" pageNumber="14">
Phylogenetic relationships -The maximum likelihood and the Bayesian likelihood analysis yielded identical topologies. Due to the size of the data set, it was not practical to estimate nodal support of the maximum likelihood topology via bootstrapping. Therefore, we report the maximum likelihood topology, conveying information on branch lengths, and indicate support for individual nodes with Bayesian posterior probability values (Fig. 9).
<taxonomicName id="F550EF86CC63AD61C94FC4FCDA122230" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
shows a statistically well supported sister taxon relationship to
<taxonomicName id="96DCF9D18626E91236314C46DA4C2BFE" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="brasiliensis">K. brasiliensis</taxonomicName>
(Figs. 9, 10). The monophyly of
<taxonomicName id="CBF490208BE9D6FA859154DF8C5BDDB2" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
whose members in addition to
<taxonomicName id="770A6F834C3A69A8967A1978E129DABB" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
are
<taxonomicName id="49EE2890F9CB39FF3C56BE2E8FDCFD2F" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="marmoratus">K. marmoratus</taxonomicName>
,
<taxonomicName id="36B9AE0694A1BF3F8AD8C704681BE0C3" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="caudomarginatus">K. caudomarginatus</taxonomicName>
, and
<taxonomicName id="35001B5BC7B6D9373AB15EBB055C606B" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="brasiliensis">K. brasiliensis</taxonomicName>
, is also strongly supported (Figs. 9, 10). These relationships are recovered in all analyses, and strongly support the hypothesis that
<taxonomicName id="F44A742BC42B7D8DE0044F192C33F79C" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
is indeed a phylogenetic member of the genus
<taxonomicName id="54C45D8B82F89939EBA4CB8B5AA5382F" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
. Costa (2004b) also placed
<taxonomicName id="BF40123E16227717B0116F305481C5F1" LSID-ZBK="urn:lsid:zoobank.org:act:F88985D8-B482-4EBD-A7AB-97270DDC9F8F" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus campelloi Costa 1990:42" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="campelloi">R. campelloi</taxonomicName>
into
<taxonomicName id="B57712D3F2743CA9E3380F41B4BA88CC" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
, although this species was not included in the analysis on which this taxonomic revision was based; it was reassigned without being listed as examined. The maximum parsimony topology (Fig. 10) suggests that
<taxonomicName id="338B52A35ABD4DF155FF42E9CCC80699" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
is sister to all other Rivulidae as hypothesized by Hrbek &amp; Larson (1999), while the maximum likelihood topology (Fig. 9) suggests that
<taxonomicName id="BBD1E2B25591A97CDA89C1775B08159B" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
is sister to Rivulinae as hypothesized by Hrbek et al. (2004). The phylogenetic placement of
<taxonomicName id="25DE5B79D6908D82924BF7F4E12A5892" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
observed in the maximum likelihood hypothesis (Fig. 9) is also supported by Costa’s (2004b) recent maximum parsimony analysis of morphological data. Thus the most likely phylogenetic hypothesis for
<taxonomicName id="89D58691042EF73A1CDB7726A3963581" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
will need further investigation, but its monophyly and differentiation from
<taxonomicName id="E100093EC301CD4560861AED9AE13808" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
is undisputed. The two species (
<taxonomicName id="34A310E9EF92C641E5AC0C07E03B1071" LSID-ZBK="urn:lsid:zoobank.org:act:FBA3D378-1C49-443A-A0E2-E8F9EE0E5BE4" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus geayi Vaillant 1899:156 [3]" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="geayi">Rivulus aff. geayi</taxonomicName>
and
<taxonomicName id="741E4270EFB65C88FDDFFD3F5DECA055" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="urophthalmus">Rivulus aff. urophthalmus</taxonomicName>
) with which
<taxonomicName id="4B496D65846160042FC544F3CCCDC5ED" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
co-occurs, are members of a phylogenetically very distinct Amazonian clade of
<taxonomicName id="921DC42D050BF735A0BEB40A47C5D703" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
(Hrbek &amp; Larson 1999), and are not closely related to
<taxonomicName id="2E979A016D8B2050754E4B46FE52FC89" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="14" phylum="Chordata" rank="species" species="sepia">K. sepia</taxonomicName>
(Figs. 9, 10).
</paragraph>
<paragraph id="81ECE7D7E782038D614E2FFFCF0DDFD0" pageNumber="17">Discussion</paragraph>
<paragraph id="B9F1192DB840A78DE0DD8141D3375A6A" pageNumber="17">
In the Tapanahony and Palumeu River drainages, as well as in other areas of northern South America there is a pattern of co-appearance of robust and small
<taxonomicName id="9FD6E1B7BC35D8EFD315EE49244619AB" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
species. The robust and the small species form reciprocally monophyletic groups which were labeled as the Guyana Coast and Guyana Shield clades, respectively (Hrbek &amp; Larson 1999). Cooccurrence of robust and small species of
<taxonomicName id="BD1D1CF2D8E156006F882F6D647C66CE" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
is usually observed in shaded primary forest of the mountainous highland plateau. These biotopes are characterized by very shallow, slowly running or stagnant water, low pH (3.5-5.1), low mineral concentrations (hardness &lt;0.5 DH), and thick canopy with diffuse light. These marginal habitats are unoccupied by other fish species except for occasional
<taxonomicName id="EFD4F89C0E498F96CF80C267D37E84BE" LSID-ZBK="urn:lsid:zoobank.org:act:1658C0D0-184E-4A8B-9DC9-9B076B2751EE" family="Lebiasinidae" genus="Pyrrhulina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pyrrhulina Valenciennes 1846:535" order="Characiformes" pageNumber="17" phylum="Chordata" rank="genus">Pyrrhulina</taxonomicName>
spp.
</paragraph>
<paragraph id="C71AEB0F10F2900B83FD8AFCF7EB5487" pageNumber="17">
Typical community compositions include
<taxonomicName id="40E0367F955DD95EF17E391527033167" LSID-ZBK="urn:lsid:zoobank.org:act:074615A2-A324-4E15-8AAA-345FBF2C308D" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus igneus Huber 1991:68" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="igneus">R. igneus</taxonomicName>
with
<taxonomicName id="C1F1FA1FB60C4B746A8C952B73463583" LSID-ZBK="urn:lsid:zoobank.org:act:50858393-2EDF-4987-B505-2ABD204B1ED8" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus cladophorus Huber 1991:71" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="cladophorus">R. cladophorus</taxonomicName>
or
<taxonomicName id="614412261D01638442071429CB195842" LSID-ZBK="urn:lsid:zoobank.org:act:FBA3D378-1C49-443A-A0E2-E8F9EE0E5BE4" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus geayi Vaillant 1899:156 [3]" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="geayi">R. geayi</taxonomicName>
;
<taxonomicName id="DF8547BBD9D065A14E3C53A4507A174A" LSID-ZBK="urn:lsid:zoobank.org:act:074615A2-A324-4E15-8AAA-345FBF2C308D" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus igneus Huber 1991:68" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="igneus">Rivulus aff. igneus</taxonomicName>
with
<taxonomicName id="E3EF91C30D25416A95B85F7A32A79B25" LSID-ZBK="urn:lsid:zoobank.org:act:FBA3D378-1C49-443A-A0E2-E8F9EE0E5BE4" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus geayi Vaillant 1899:156 [3]" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="geayi">Rivulus aff. geayi</taxonomicName>
;
<taxonomicName id="57646797D2D942DE1EE1E1DF92D68882" LSID-ZBK="urn:lsid:zoobank.org:act:9E74E99A-0F37-473B-A742-8C1E1423A54A" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus waimacui Eigenmann 1909:50" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="waimacui">R. waimacui</taxonomicName>
with
<taxonomicName id="F6C766981388E0571B5C9EBD3CEAEF24" LSID-ZBK="urn:lsid:zoobank.org:act:28242994-BF55-43B9-AA5B-89413344A340" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus breviceps Eigenmann 1909:49" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="breviceps">R. breviceps</taxonomicName>
or
<taxonomicName id="759FCBAC26AB36A0E828DB873D33076F" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="sp. mahdia">R. sp. Mahdia</taxonomicName>
; and
<taxonomicName id="C9F9819A6E32043B9B8794699C29AF97" LSID-ZBK="urn:lsid:zoobank.org:act:DB932953-4461-47B1-9ABE-7FF68B5B63D9" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus immaculatus Thomerson, Nico &amp; Taphorn 1991:323" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="immaculatus">R. immaculatus</taxonomicName>
with
<taxonomicName id="7A94C55254A934C119AB9FDAC931D978" LSID-ZBK="urn:lsid:zoobank.org:act:6682190D-4BFC-4AB6-B2A1-3A19A95CD22D" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus gransabanae Lasso, Taphorn &amp; Thomerson 1992:298" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="gransabanae">R. gransabanae</taxonomicName>
or
<taxonomicName id="084CA11056DD4D6EE921219A7D655A13" LSID-ZBK="urn:lsid:zoobank.org:act:C1872DED-5792-41AB-AE50-EAFAAE1B5FE0" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus torrenticola Vermeulen &amp; Isbrücker 2000:186" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="torrenticola">R. torrenticola</taxonomicName>
. Sister clade to the Guyana Coast is the Amazon basin clade (Hrbek &amp; Larson 1999) whose members form similar associations with the eastern Guyana Shield subclade (Hrbek et al. 2004). These associations include
<taxonomicName id="CBC8BF6802054BC94005EE0B1F60CBA8" LSID-ZBK="urn:lsid:zoobank.org:act:89A5E0A8-6F08-4AC2-B367-8DB7EE1EF75E" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus ophiomimus Huber 1992:344" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="ophiomimus">R. ophiomimus</taxonomicName>
with
<taxonomicName id="E07357360F6FC77602FABEF2F10C7377" LSID-ZBK="urn:lsid:zoobank.org:act:65F512B3-6450-4C16-8292-22BECCD09F81" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus rectocaudatus Fels &amp; de Rham 1981:66" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="rectocaudatus">R. rectocaudatus</taxonomicName>
and
<taxonomicName id="F5B1317206B2682BAEE25AA290775C2C" LSID-ZBK="urn:lsid:zoobank.org:act:862F43D1-9464-44C3-AD55-3486C85230D8" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus compressus Henn 1916:111" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="compressus">R. compressus</taxonomicName>
with
<taxonomicName id="1F5F09E20F6500C5665F0E7C94AB15E4" LSID-ZBK="urn:lsid:zoobank.org:act:10B66DC4-3690-4416-B71D-672993F831B6" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus duckensis Hrbek, de Deus &amp; Farias 2004:571" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="duckensis">R. duckensis</taxonomicName>
. However, even within these communities, the robust species tends to be found in flooded swampy parts of the creek, while the smaller species tends to be found in the creek proper. Naturally some species within these clades do not fit this ecological pattern. These species include, for example,
<taxonomicName id="FE8FCBBDDB56E0899C121DE34F86CCDD" LSID-ZBK="urn:lsid:zoobank.org:act:C85063FD-B6D1-4505-A11A-2808251D256A" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus agilae Hoedeman 1954:203" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="agilae">R. agilae</taxonomicName>
which prefers open savannah and secondary forest habitats, and usually occurs alone, or
<taxonomicName id="DB916B23D24A2B7F12A7068AFDA0A7C6" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="hartii">R. hartii</taxonomicName>
which is found in a wide variety of habitats but seem to usually co-occur with poeciliids for which they are a major predator (Reznick 1982). The
<taxonomicName id="8A395C83E83D4AF82B99BB2CED269EBA" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="urophthalmus">R. urophthalmus species complex</taxonomicName>
, which includes
<taxonomicName id="316DEAE2C728C738BDC8359F64CA3BD2" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="urophthalmus">R. urophthalmus</taxonomicName>
,
<taxonomicName id="4EF4EE07354A1B3EECC5015753AAF66E" LSID-ZBK="urn:lsid:zoobank.org:act:561FF4A2-6736-4ABA-80D0-5AE89957E232" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus stagnatus Eigenmann 1909:50" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="stagnatus">R. stagnatus</taxonomicName>
,
<taxonomicName id="908AA187E0F753B9B9EB9F4713401A0B" LSID-ZBK="urn:lsid:zoobank.org:act:58A563DB-CD44-49E6-8757-EF2D9F1C0B95" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus deltaphilus Seegers 1983:39" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="deltaphilus">R. deltaphilus</taxonomicName>
and
<taxonomicName id="5B3E9DBCFAE98A16483AE420C45669D2" LSID-ZBK="urn:lsid:zoobank.org:act:DE0DECEC-8730-4112-A156-A628298E68D2" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus lungi Berkenkamp 1984:14" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="lungi">R. lungi</taxonomicName>
among others, occurs with rivulids as well as poeciliids.
</paragraph>
<paragraph id="1A06339A48148CBF0BF8B2020FFDF1BF" pageNumber="17">
The newly described
<taxonomicName id="ACA7AC68940145725F0CAE82558AB66E" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
also fits within this pattern of co-occurrence. Although it superficially resembles robust
<taxonomicName id="8FD52E63ACADAEC2332DD6208E963490" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
species, it is a member of a phylogenetically distinct clade, and appears to be ecologically differentiated as well. Formal taxonomic recognition of this clade did not occur until recently when Costa (2004b) described the genus
<taxonomicName id="979E473F50E769BDDAC80E1A71FCE0FD" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
. Costa (2004b) reanalyzed
<taxonomicName id="55165D8FBA3944B221B015A69EE91A71" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="brasiliensis">Rivulus brasiliensis</taxonomicName>
and its close relatives -all species analyzed in Costa (1998) as well -and no longer found
<taxonomicName id="80478A0EAB4F441E2E8E778ED8E23DEB" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="species" species="brasiliensis">R. brasiliensis</taxonomicName>
and its relatives to have synapomorphies which previously placed it within a monophyletic
<taxonomicName id="FF6699D12BE7FEBBE61D5D309DD4CEAA" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
(Costa 1998). This made
<taxonomicName id="644A3A575B3E5DB6C91D63E79F2AE2DA" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
non-monophyletic, a hypothesis that has been put forth by molecular phylogenetic analyses (Hrbek &amp; Larson 1999, Murphy et al. 1999), and by an unpublished molecular study reviewed by and cited in Costa (1998). Costa (2004b) therefore proposed the new generic name
<taxonomicName id="F92D9082020E1207861BD5D1E6E245D8" LSID-ZBK="urn:lsid:zoobank.org:act:C7A538BD-E800-4591-9D05-E094246E9BBA" family="Rivulidae" genus="Cryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Cryptolebias Costa 2004:115" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Cryptolebias</taxonomicName>
which, however, is preoccupied by a fossil cyprinodont fish (Gaudant 1978). As first reviser, Costa (2004a) substituted the name
<taxonomicName id="B5A6BB73CCFD4FE9B7EAE6FAB1A90124" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
for
<taxonomicName id="B4F4044758A3A17F89F0D7F546FD3CF1" LSID-ZBK="urn:lsid:zoobank.org:act:C7A538BD-E800-4591-9D05-E094246E9BBA" family="Rivulidae" genus="Cryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Cryptolebias Costa 2004:115" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Cryptolebias</taxonomicName>
. In spite of a somewhat complicated morphological history, all lines of evidence now clearly show that
<taxonomicName id="0683BBB97809E1983B2D1EC6584EA95C" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
is a clade distinct from the remaining
<taxonomicName id="F1B806AD5EABAF91B90DF746B025E146" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
species. Whether the remaining
<taxonomicName id="06041A999707ED68CAF7FBD83A5377D2" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
species form a clade (Costa 1998, 2004b) or are non-monophyletic (Hrbek &amp; Larson 1999, Murphy et al. 1999, Hrbek et al. 2004) remains to be resolved, but the taxonomic validity of
<taxonomicName id="9037AAD3EEA818EA87A6A78CAE1DA185" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="17" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
is upheld by this study.
</paragraph>
<paragraph id="4E1987C5C0B691050D43E2A96371AA10" pageNumber="18">Conclusion</paragraph>
<paragraph id="7947B104480F1842B541FA8F1C1EA7B8" pageNumber="18">
<taxonomicName id="82525B4DE1DF25603D05A608CA0F1D62" LSID-ZBK="urn:lsid:zoobank.org:act:D6AC62C8-15CA-4620-A15A-A0AE945E5E67" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias sepia Vermeulen &amp; Hrbek 2005:4" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="species" species="sepia">Kryptolebias sepia</taxonomicName>
is a distinct taxon that is morphologically, ecologically and phylogenetically differentiated from sympatrically occurring robust (
<taxonomicName id="1525AB5F69BBD3159C48F6F89B8C7C4B" LSID-ZBK="urn:lsid:zoobank.org:act:074615A2-A324-4E15-8AAA-345FBF2C308D" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus igneus Huber 1991:68" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="species" species="igneus">Rivulus aff. igneus</taxonomicName>
or
<taxonomicName id="3FAE26C518497FF382425A74D679B656" LSID-ZBK="urn:lsid:zoobank.org:act:65D0D6C9-7C66-46AA-9494-7FED50DE6DBF" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus urophthalmus Günther 1866:327" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="species" species="urophthalmus">R. urophthalmus</taxonomicName>
) and small (
<taxonomicName id="D3C91B768D5C6F6BB29B08364BA4520C" LSID-ZBK="urn:lsid:zoobank.org:act:FBA3D378-1C49-443A-A0E2-E8F9EE0E5BE4" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus geayi Vaillant 1899:156 [3]" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="species" species="geayi">Rivulus aff. geayi</taxonomicName>
) species. It occurs in low densities in a rarely collected primary forest habitat of the central mountains of Surinam, and is easily overlooked in usual surveys. This study clearly shows that a better understanding of rivuline biodiversity, and of the genera
<taxonomicName id="5D717F65EEA9FD02BB1B82816B0CB13A" LSID-ZBK="urn:lsid:zoobank.org:act:7737CC47-FB47-4DAF-87C2-18950BAD40FA" family="Rivulidae" genus="Rivulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulus Poey 1860:307" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="genus">Rivulus</taxonomicName>
and
<taxonomicName id="463508F043EB3298BB23075C42317B2F" LSID-ZBK="urn:lsid:zoobank.org:act:94FD1A9A-96E4-4EFE-820B-1313C8AB57F6" family="Rivulidae" genus="Kryptolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Kryptolebias Costa 2004" order="Cyprinodontiformes" pageNumber="18" phylum="Chordata" rank="genus">Kryptolebias</taxonomicName>
specifically, will require a focused collecting effort in appropriate habitats, as well as the combined use of molecular and morphological data in taxonomic and phylogenetic studies.
</paragraph>
</subSubSection>
</treatment>
</document>
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