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<document id="2B4C43EB553512A8318BCEA2F5C940D9" ID-DOI="10.5281/zenodo.10881992" ID-ISSN="0936-9902" ID-Zenodo-Dep="10881992" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatmentCitations_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1711474576113" checkinUser="juliana" docAuthor="Weitzman, Stanley H. &amp; Malabarba, Luiz R." docDate="1999" docId="03DA87B22A39FFF8FF5CF97448C1FB7A" docLanguage="en" docName="IchthyolExplorFres.10.1.1-43.pdf.imf" docOrigin="Ichthyol. Explor. Freshwaters 10 (1)" docTitle="Spintherobolus Eigenmann 1911" docType="treatment" docVersion="3" lastPageNumber="20" masterDocId="FFE3FFCA2A33FFEBFFB0FF824B09FF9B" masterDocTitle="Systematics of Spintherobolus (Teleostei: Characidae: Cheirodontinae) from eastern Brazil" masterLastPageNumber="43" masterPageNumber="1" pageNumber="11" updateTime="1711475764197" updateUser="ExternalLinkService">
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<mods:title id="3676C9C93454D1AA5DE5B678433C1CDE">Systematics of Spintherobolus (Teleostei: Characidae: Cheirodontinae) from eastern Brazil</mods:title>
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<mods:namePart id="C2630955F08C11C4C724513354A05DB7">Weitzman, Stanley H.</mods:namePart>
<mods:affiliation id="84531816B437049C61882B62BBDD23F0">Division of Fishes, Department of Vertebrate Zoology, MRC- 159, Smithsonian Institution, Washington D. C. 20560, USA.</mods:affiliation>
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<mods:namePart id="8A63229DC1B2070EA3E429E0022E1901">Malabarba, Luiz R.</mods:namePart>
<mods:affiliation id="E95656730E5B932FAFBE55443E70F444">Laboratório de Ictiologia, Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Av. Ipiranga, 6681, CEP 90619 - 900, Porto Alegre, RS, Brazil, and Departamento de Zoologia, IB, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil.</mods:affiliation>
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<mods:title id="2C842B34A77EF2C94904B9FF0D67374E">Ichthyol. Explor. Freshwaters</mods:title>
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<mods:date id="95BEAEACFD01A968B596E6DF88CACAAB">1999</mods:date>
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<mods:identifier id="0461F3913F3BF2BE333D4D57967A1F2D" type="DOI">10.5281/zenodo.10881992</mods:identifier>
<mods:identifier id="2E68B78E524D86848A20E87115BD1941" type="ISSN">0936-9902</mods:identifier>
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<treatment id="03DA87B22A39FFF8FF5CF97448C1FB7A" ID-DOI="http://doi.org/10.5281/zenodo.10881805" ID-Zenodo-Dep="10881805" LSID="urn:lsid:plazi:treatment:03DA87B22A39FFF8FF5CF97448C1FB7A" httpUri="http://treatment.plazi.org/id/03DA87B22A39FFF8FF5CF97448C1FB7A" lastPageId="19" lastPageNumber="20" pageId="10" pageNumber="11">
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<paragraph id="8BCC36A42A39FFE1FF5CF97449B6F880" blockId="10.[236,703,1782,1819]" box="[236,703,1782,1819]" pageId="10" pageNumber="11">
<heading id="D08481C82A39FFE1FF5CF97449B6F880" box="[236,703,1782,1819]" centered="true" fontSize="9" level="4" pageId="10" pageNumber="11" reason="1">
<taxonomicName id="4C734D272A39FFE1FF5CF97449B6F880" ID-CoL="7L95" ID-ENA="930386" authority="Eigenmann" authorityName="Eigenmann" authorityYear="1911" box="[236,703,1782,1819]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A39FFE1FF5CF9744AE1F880" box="[236,488,1782,1819]" italics="true" pageId="10" pageNumber="11">Spintherobolus</emphasis>
Eigenmann
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<paragraph id="8BCC36A42A39FFE1FFF1F8D249A4F8EC" blockId="10.[65,875,1872,2001]" box="[65,685,1872,1911]" pageId="10" pageNumber="11">
<treatmentCitationGroup id="AB63118A2A39FFE1FFF1F8D249A4F8EC" box="[65,685,1872,1911]" pageId="10" pageNumber="11">
<treatmentCitation id="0AD210B52A39FFE1FFF1F8D249A4F8EC" author="Eigenmann, C. H." box="[65,685,1872,1911]" page="167" pageId="10" pageNumber="11" year="1911">
<taxonomicName id="4C734D272A39FFE1FFF1F8D249A4F8EC" ID-CoL="7L95" authority="Eigenmann, 1911: 167" authorityName="Eigenmann" authorityPageNumber="167" authorityYear="1911" box="[65,685,1872,1911]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A39FFE1FFF1F8D24A28F8EC" box="[65,289,1872,1911]" italics="true" pageId="10" pageNumber="11">Spintherobolus</emphasis>
<bibRefCitation id="EFE24B552A39FFE1FE87F8D249A4F8EC" author="Eigenmann, C. H." box="[311,685,1872,1911]" pageId="10" pageNumber="11" pagination="164 - 181" refId="ref29012" refString="Eigenmann, C. H. 1911. New characins in the collections of the Carnegie Museum. Ann. Carnegie Mus., 8: 164 - 181." type="journal article" year="1911">Eigenmann, 1911: 167</bibRefCitation>
</taxonomicName>
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</paragraph>
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<subSubSection id="C369652F2A39FFE1FD70F8D249EBF84A" pageId="10" pageNumber="11" type="type_taxon">
<paragraph id="8BCC36A42A39FFE1FD70F8D249EBF84A" blockId="10.[65,875,1872,2001]" pageId="10" pageNumber="11">
<treatmentCitationGroup id="AB63118A2A39FFE1FD70F8D249EBF84A" pageId="10" pageNumber="11">
(
<typeStatus id="54C888062A39FFE1FD7CF8D2481DF8EC" box="[716,788,1872,1911]" pageId="10" pageNumber="11">type</typeStatus>
species:
<treatmentCitation id="0AD210B52A39FFE1FF64F8FF4839F83E" author="Eigenmann, C. H." box="[212,816,1917,1957]" page="167" pageId="10" pageNumber="11" year="1911">
<taxonomicName id="4C734D272A39FFE1FF64F8FF4839F83E" ID-CoL="4Z63S" authority="Eigenmann, 1911: 167" authorityName="Eigenmann" authorityPageNumber="167" authorityYear="1911" box="[212,816,1917,1957]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A39FFE1FF64F8FF4BE4F83E" box="[212,237,1917,1957]" italics="true" pageId="10" pageNumber="11">S.</emphasis>
<emphasis id="B907EAB62A39FFE1FF4AF8FF4AAAF83E" box="[250,419,1917,1957]" italics="true" pageId="10" pageNumber="11">papilliferus</emphasis>
<bibRefCitation id="EFE24B552A39FFE1FE08F8FF4839F83E" author="Eigenmann, C. H." box="[440,816,1917,1957]" pageId="10" pageNumber="11" pagination="164 - 181" refId="ref29012" refString="Eigenmann, C. H. 1911. New characins in the collections of the Carnegie Museum. Ann. Carnegie Mus., 8: 164 - 181." type="journal article" year="1911">Eigenmann, 1911: 167</bibRefCitation>
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by monotypy and original designation)
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C369652F2A39FFE1FFF2F78648F0FA46" pageId="10" pageNumber="11" type="diagnosis">
<paragraph id="8BCC36A42A39FFE1FFF2F78648F0FA46" blockId="10.[63,874,2052,2588]" lastBlockId="10.[932,1737,1236,1501]" pageId="10" pageNumber="11">
Diagnosis. Jaw teeth elongate and conical or tricuspid (
<figureCitation id="13482A212A39FFE1FF76F7B04A38F7C3" box="[198,305,2098,2136]" captionStart="Fig" captionStartId="10.[933,981,1063,1097]" captionTargetBox="[933,1740,197,1020]" captionTargetPageId="10" captionText="Fig. 12. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Jaws &amp; teeth." figureDoi="http://doi.org/10.5281/zenodo.10882020" httpUri="https://zenodo.org/record/10882020/files/figure.png" pageId="10" pageNumber="11">Fig. 12</figureCitation>
); anterior ventral procurrent caudal-fin rays with proximal ends anterior to haemal spine of antepenultimate vertebrae fused to one another proximally; symphyseal dentary ar ­ ticulations are smooth, lacking intercalated folded bony surfaces common to most characids; neuromasts on head obvious and present in shallow channels and distributed on head in a char ­ acteristic manner in which one horizontal series in the infraorbital region is crossed by several more or less vertical series (
<figureCitation id="13482A212A39FFE1FD8DF67749AEF587" box="[573,679,2549,2588]" captionStart="Fig" captionStartId="6.[70,119,841,876]" captionTargetBox="[68,882,219,802]" captionTargetPageId="6" captionText="Fig. 4. Spintherobolus broccae, USNM 287324, adult male, 17.9 mm SL. Head and anterior body region, right side. Two pseudotympanums, black areas are located posterior to the dorsal portion of the opercular opening. The posterior pseudotympanum is homologous with the single pseudotympanum of other Cheirodontines while the anterior pseudotympanum is found only in Spintherobolus. The small dark dots arranged in rows arer the neuromast pattern over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region and on the dentary." figureDoi="http://doi.org/10.5281/zenodo.10882001" httpUri="https://zenodo.org/record/10882001/files/figure.png" pageId="10" pageNumber="11">Figs 4</figureCitation>
,
<figureCitation id="13482A212A39FFE1FD76F67749E0F587" box="[710,745,2549,2588]" captionStart="Fig" captionStartId="9.[170,218,740,776]" captionTargetBox="[167,836,218,695]" captionTargetPageId="9" captionText="Fig. 10. Spintherobolus papilliferus, MNRJ 4237, female, 34.7 mm SL, head region, left side, Brazil, São Paulo, rio Ipiranga. Rows of neuromasts occur over the infraorbital, preopercular, and opercular region. These are distributed in shallow grooves that can be seen in the photograph. The neuromasts are also distributed on the scale borders posterior to the head and operculum but these are hard to distinguish from dark chromatophores in the photograph. A row of neuromasts along the ventral border of the dentary is hidden in a deep groove. The small dark dots arranged in rows are the neuromast patterns over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region." figureDoi="http://doi.org/10.5281/zenodo.10882016" httpUri="https://zenodo.org/record/10882016/files/figure.png" pageId="10" pageNumber="11">10</figureCitation>
&amp;
<figureCitation id="13482A212A39FFE1FC81F6774859F587" box="[817,848,2549,2588]" captionStart="Fig" captionStartId="9.[169,217,1134,1180]" captionTargetBox="[848,1844,271,681]" captionTargetPageId="9" captionText="Fig. 11. Spintherobolus papilliferus, MZUSP 51021, female 60.8 mm SL. Dorsum of head and back nearly to dorsal-fin origin. Neuromasts are present in the irregular grooves between the eyes and the supraoccipital region but these are nearly impossible to see in the photograph. A bilaterally distributed row of neuromasts runs along the long axis of the body between the nares. This row of neuromasts shows more clearly on the left side than on the right side of the top of the head." figureDoi="http://doi.org/10.5281/zenodo.10882018" httpUri="https://zenodo.org/record/10882018/files/figure.png" pageId="10" pageNumber="11">11</figureCitation>
); dentary with a large anterior fenestra (
<figureCitation id="13482A212A39FFE1F9E3FB564DB5FB66" box="[1619,1724,1236,1277]" captionStart="Fig" captionStartId="10.[933,981,1063,1097]" captionTargetBox="[933,1740,197,1020]" captionTargetPageId="10" captionText="Fig. 12. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Jaws &amp; teeth." figureDoi="http://doi.org/10.5281/zenodo.10882020" httpUri="https://zenodo.org/record/10882020/files/figure.png" pageId="10" pageNumber="11">Fig. 12</figureCitation>
) associated with large epidermal, papilla-like structure bearing several neuromasts; anterior pseudotympanum occurs anterior to first pleural rib (see character 21 below for detailed explanation).
</paragraph>
</subSubSection>
<subSubSection id="C369652F2A39FFFAFC16F9914A5CF583" lastPageId="17" lastPageNumber="18" pageId="10" pageNumber="11" type="description">
<paragraph id="8BCC36A42A39FFE1FC16F9914FB7F8EF" blockId="10.[930,1736,1555,2589]" pageId="10" pageNumber="11">
Monophyly of
<emphasis id="B907EAB62A39FFE1FB11F9914EADF9A0" box="[1185,1444,1555,1595]" italics="true" pageId="10" pageNumber="11">
<taxonomicName id="4C734D272A39FFE1FB11F9914EA9F9A0" authorityName="Eigenmann" authorityYear="1911" box="[1185,1440,1555,1595]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
The following 14 (15-27, 3
<collectionCode id="ED62AE612A39FFE1FB9EF9BD4F4FF9F3" box="[1070,1094,1599,1640]" pageId="10" pageNumber="11">R</collectionCode>
) synapomorphies indicate monophyly for
<emphasis id="B907EAB62A39FFE1FC53F9EC4EB4F90F" box="[995,1469,1646,1684]" italics="true" pageId="10" pageNumber="11">
<taxonomicName id="4C734D272A39FFE1FC53F9EC4FCAF90F" authorityName="Eigenmann" authorityYear="1911" box="[995,1219,1646,1684]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
<taxonomicName id="4C734D272A39FFE1FB6EF9EC4EB4F90F" authorityName="Eigenmann" authorityYear="1911" box="[1246,1469,1646,1684]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
</emphasis>
consists of two sister groups, one (a) containing
<taxonomicName id="4C734D272A39FFE1FA41F91E4DCEF959" authorityName="Eigenmann" authorityYear="1911" box="[1521,1735,1692,1730]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A39FFE1FA41F91E4D02F959" box="[1521,1547,1692,1730]" italics="true" pageId="10" pageNumber="11">S.</emphasis>
<emphasis id="B907EAB62A39FFE1F990F91E4DCEF959" box="[1568,1735,1692,1730]" italics="true" pageId="10" pageNumber="11">papilliferus</emphasis>
</taxonomicName>
and the other (b) the remaining three species. This information is provided at this point to facilitate discussion of the fourteen synapomorphies described below.
</paragraph>
<paragraph id="8BCC36A42A39FFE1FC50F8F94EA8F65A" blockId="10.[930,1736,1555,2589]" pageId="10" pageNumber="11">
(3
<collectionCode id="ED62AE612A39FFE1FC4EF8F94F1FF83F" box="[1022,1046,1915,1956]" pageId="10" pageNumber="11">R</collectionCode>
) The jaw teeth are elongate and conical or tricuspid. When tricuspid there are small lateral cusps near the distal apices of what are essentially teeth having an elongate pedicle and a slightly recurved apical cone (
<figureCitation id="13482A212A39FFE1FAA7F7B24E81F7C3" box="[1303,1416,2096,2136]" captionStart="Fig" captionStartId="10.[933,981,1063,1097]" captionTargetBox="[933,1740,197,1020]" captionTargetPageId="10" captionText="Fig. 12. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Jaws &amp; teeth." figureDoi="http://doi.org/10.5281/zenodo.10882020" httpUri="https://zenodo.org/record/10882020/files/figure.png" pageId="10" pageNumber="11">Fig. 12</figureCitation>
). The anterior dentary teeth of
<taxonomicName id="4C734D272A39FFE1FBDDF7DE4E42F71E" authorityName="Eigenmann" authorityYear="1911" box="[1133,1355,2140,2181]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A39FFE1FBDDF7DE4E42F71E" box="[1133,1355,2140,2181]" italics="true" pageId="10" pageNumber="11">Spintherobolus</emphasis>
</taxonomicName>
have three small cusps. Teeth bearing these small cusps are more numer ­ ous than the strictly conical teeth present in the large species,
<emphasis id="B907EAB62A39FFE1FB30F7664E5AF696" box="[1152,1363,2276,2317]" italics="true" pageId="10" pageNumber="11">
<taxonomicName id="4C734D272A39FFE1FB30F7664E46F696" authorityName="Eigenmann" authorityYear="1911" box="[1152,1359,2276,2317]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="papilliferus">S. papilliferus</taxonomicName>
.
</emphasis>
The premaxillary teeth are usually conical, but in
<taxonomicName id="4C734D272A39FFE1FAE7F6914D2DF6A1" authorityName="Eigenmann" authorityYear="1911" box="[1367,1572,2323,2362]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A39FFE1FAE7F6914D2DF6A1" box="[1367,1572,2323,2362]" italics="true" pageId="10" pageNumber="11">S. papilliferus</emphasis>
</taxonomicName>
they bear some very small lateral cusps in their somewhat laterally expanded distal portions. The maxillary teeth in all species are conical.
</paragraph>
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Tooth shape, tooth counts and the number of cusps have been extensively used for suggesting cheirodontine and other characid relationships since
<bibRefCitation id="EFE24B552A38FFE0FEB7FF7D4931FEBC" author="Eigenmann, C. H." bookContentInfo="pls. 1 - 17" box="[263,568,255,295]" journalOrPublisher="Mem. Carnegie Mus." pageId="11" pageNumber="12" pagination="1 - 99" part="7" refId="ref29042" refString="- 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Mem. Carnegie Mus., 7: 1 - 99, pls. 1 - 17." title="The Cheirodontinae, a subfamily of minute characid fishes of South America" type="journal article" year="1915">Eigenmann (1915)</bibRefCitation>
used these features in Cheirodontines and other small characids. As discussed by
<bibRefCitation id="EFE24B552A38FFE0FE3AFED948CAFE1A" author="Weitzman, S. H. &amp; W. L. Fink" box="[394,963,347,385]" pageId="11" pageNumber="12" pagination="339 - 395" refId="ref31246" refString="Weitzman, S. H. &amp; W. L. Fink. 1983. Relationships of the neon tetras, a group of South American freshwater fishes (Teleostei, Characidae), with comments on the phylogeny of the New World characiforms. Bull. Mus. Comp. Zool., 150: 339 - 395." type="journal article" year="1983">Weitzman &amp; Fink (1983: 341-353)</bibRefCitation>
, features such as numbers of rows of teeth, counts of the numbers of teeth in a particular jaw bone, the number of cusps in particular teeth, and the shape and form of the teeth and their cusps var ­ ies widely in various lineages of the American
<taxonomicName id="4C734D272A38FFE0FF12FDEA4A52FD0B" baseAuthorityName="Vari &amp; Siebert" baseAuthorityYear="1990" box="[162,347,616,656]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="family">Characidae</taxonomicName>
and provide data in support of phylogenetic hypotheses. However, as with many other characters, convergent evolution of such tooth features often cause problems in phylogenetic reconstruction, especially when teeth are not compared and described in enough detail to detect differences and similarities and are used for this purpose without the abundant use of other data.
</paragraph>
<paragraph id="8BCC36A42A38FFE0FF6CFC7E4A48F836" blockId="11.[154,971,209,2600]" pageId="11" pageNumber="12">
Conical and tricuspid teeth are found in a number of American characids. The clades within the
<taxonomicName id="4C734D272A38FFE0FEB6FBDA490EFB1B" box="[262,519,1112,1152]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="14" phylum="Chordata" rank="subFamily" subFamily="Cheirodontinae">Cheirodontinae</taxonomicName>
closest to
<taxonomicName id="4C734D272A38FFE0FD03FBDA489DFB1B" authorityName="Eigenmann" authorityYear="1911" box="[691,916,1112,1152]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0FD03FBDA489DFB1B" box="[691,916,1112,1152]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
on the basis of other synapomorphies have five or more cusps, indicating that the low cusp number in
<taxonomicName id="4C734D272A38FFE0FF7AFB634AA1FA93" authorityName="Eigenmann" authorityYear="1911" box="[202,424,1249,1288]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0FF7AFB634AA1FA93" box="[202,424,1249,1288]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
is derived. Some species belonging to other cheirodontine clades also have tricuspid teeth, but their teeth are never elongate and essentially conical in shape at their apices. For example,
<taxonomicName id="4C734D272A38FFE0FE38FA1A4A30FA70" authority="(Eigenmann, 1915: 80)" baseAuthorityName="Eigenmann" baseAuthorityPageNumber="80" baseAuthorityYear="1915" family="Characidae" genus="Odontostilbe" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="microdon">
<emphasis id="B907EAB62A38FFE0FE38FA1A49EBFA25" box="[392,738,1432,1470]" italics="true" pageId="11" pageNumber="12">Odontostilbe microdon</emphasis>
(
<bibRefCitation id="EFE24B552A38FFE0FCB0FA1A4A27FA70" author="Eigenmann, C. H." bookContentInfo="pls. 1 - 17" journalOrPublisher="Mem. Carnegie Mus." pageId="11" pageNumber="12" pagination="1 - 99" part="7" refId="ref29042" refString="- 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Mem. Carnegie Mus., 7: 1 - 99, pls. 1 - 17." title="The Cheirodontinae, a subfamily of minute characid fishes of South America" type="journal article" year="1915">Eigenmann, 1915: 80</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C734D272A38FFE0FEE5FA474BDBF982" authority="Eigenmann (1927: 44)" authorityName="Eigenmann" authorityPageNumber="44" authorityYear="1927" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="australe">
<emphasis id="B907EAB62A38FFE0FEE5FA474AE6FA70" box="[341,495,1477,1515]" italics="true" pageId="11" pageNumber="12">Cheirodon</emphasis>
<emphasis id="B907EAB62A38FFE0FDB4FA474977FA70" box="[516,638,1477,1515]" italics="true" pageId="11" pageNumber="12">australe</emphasis>
<bibRefCitation id="EFE24B552A38FFE0FD23FA474BDBF982" author="Eigenmann, C. H." journalOrPublisher="Mem. Mus. Comp. Zool." pageId="11" pageNumber="12" pagination="1 - 102" part="43" refId="ref29104" refString="- 1927. The fresh-water fishes of Chile. Mem. Mus. Comp. Zool., 43: 1 - 102." title="The fresh-water fishes of Chile." type="journal article" year="1927">Eigenmann (1927: 44)</bibRefCitation>
</taxonomicName>
, and C.
<taxonomicName id="4C734D272A38FFE0FED0FA724815F982" authority="Campos (1982: 154)" authorityName="Campos" authorityPageNumber="154" authorityYear="1982" box="[352,796,1520,1561]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="kiliani">
<emphasis id="B907EAB62A38FFE0FED0FA724AB6F982" box="[352,447,1520,1561]" italics="true" pageId="11" pageNumber="12">kiliani</emphasis>
Campos (1982: 154)
</taxonomicName>
, although having tricuspid teeth, have these two secondary cusps large enough so that the teeth are broad and the cusps are more or less equal. This cusp number is here considered independently derived, not because of the different tooth shapes, but because of the numerous synapomorphies indicating these
<taxonomicName id="4C734D272A38FFE0FE61F8AF497BF8CE" box="[465,626,1837,1877]" pageId="11" pageNumber="12">
<emphasis id="B907EAB62A38FFE0FE61F8AF497BF8CE" box="[465,626,1837,1877]" italics="true" pageId="11" pageNumber="12">Cheirodon</emphasis>
</taxonomicName>
species belong to cheirodontine clades different from that of
<emphasis id="B907EAB62A38FFE0FF10F80B4A48F836" box="[160,321,1929,1965]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C734D272A38FFE0FCC5F8D84A34F836" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0FCC5F8D848CCF819" box="[885,965,1882,1922]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
.
</emphasis>
</paragraph>
<paragraph id="8BCC36A42A38FFE0FF6AF8344D2FFCDD" blockId="11.[154,971,209,2600]" lastBlockId="11.[1021,1836,212,2601]" pageId="11" pageNumber="12">
(15) A complex, patterned series of exposed neuromasts are distributed on the head and body (
<figureCitation id="13482A212A38FFE0FF19F7924A1BF7AC" box="[169,274,2064,2103]" captionStart="Fig" captionStartId="6.[70,119,841,876]" captionTargetBox="[68,882,219,802]" captionTargetPageId="6" captionText="Fig. 4. Spintherobolus broccae, USNM 287324, adult male, 17.9 mm SL. Head and anterior body region, right side. Two pseudotympanums, black areas are located posterior to the dorsal portion of the opercular opening. The posterior pseudotympanum is homologous with the single pseudotympanum of other Cheirodontines while the anterior pseudotympanum is found only in Spintherobolus. The small dark dots arranged in rows arer the neuromast pattern over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region and on the dentary." figureDoi="http://doi.org/10.5281/zenodo.10882001" httpUri="https://zenodo.org/record/10882001/files/figure.png" pageId="11" pageNumber="12">Figs. 4</figureCitation>
,
<figureCitation id="13482A212A38FFE0FE9BF7924A44F7AC" box="[299,333,2064,2103]" captionStart="Fig" captionStartId="9.[170,218,740,776]" captionTargetBox="[167,836,218,695]" captionTargetPageId="9" captionText="Fig. 10. Spintherobolus papilliferus, MNRJ 4237, female, 34.7 mm SL, head region, left side, Brazil, São Paulo, rio Ipiranga. Rows of neuromasts occur over the infraorbital, preopercular, and opercular region. These are distributed in shallow grooves that can be seen in the photograph. The neuromasts are also distributed on the scale borders posterior to the head and operculum but these are hard to distinguish from dark chromatophores in the photograph. A row of neuromasts along the ventral border of the dentary is hidden in a deep groove. The small dark dots arranged in rows are the neuromast patterns over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region." figureDoi="http://doi.org/10.5281/zenodo.10882016" httpUri="https://zenodo.org/record/10882016/files/figure.png" pageId="11" pageNumber="12">10</figureCitation>
&amp;
<figureCitation id="13482A212A38FFE0FE37F7924AAFF7AC" box="[391,422,2064,2103]" captionStart="Fig" captionStartId="9.[169,217,1134,1180]" captionTargetBox="[848,1844,271,681]" captionTargetPageId="9" captionText="Fig. 11. Spintherobolus papilliferus, MZUSP 51021, female 60.8 mm SL. Dorsum of head and back nearly to dorsal-fin origin. Neuromasts are present in the irregular grooves between the eyes and the supraoccipital region but these are nearly impossible to see in the photograph. A bilaterally distributed row of neuromasts runs along the long axis of the body between the nares. This row of neuromasts shows more clearly on the left side than on the right side of the top of the head." figureDoi="http://doi.org/10.5281/zenodo.10882018" httpUri="https://zenodo.org/record/10882018/files/figure.png" pageId="11" pageNumber="12">11</figureCitation>
). All
<taxonomicName id="4C734D272A38FFE0FDBFF79249F9F7AC" box="[527,752,2064,2103]" pageId="11" pageNumber="12">
<emphasis id="B907EAB62A38FFE0FDBFF79249F9F7AC" box="[527,752,2064,2103]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
species lack most latero-sensory canals on the head and body. Instead there are series of exposed neuromasts distributed often in shallow channels, in a pattern unique to
<taxonomicName id="4C734D272A38FFE0FE01F741499CF771" authorityName="Eigenmann" authorityYear="1911" box="[433,661,2243,2282]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0FE01F741499CF771" box="[433,661,2243,2282]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
over the dorsal, ventral, and lateral surfaces of the head. The neuromasts also occur near the posterior border of the body scales and scales at the base of the caudal fin. The neuromast lines of the head for
<taxonomicName id="4C734D272A38FFE0FF2DF6254A27F655" box="[157,302,2471,2510]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A38FFE0FF2DF6254BBCF655" box="[157,181,2471,2510]" italics="true" pageId="11" pageNumber="12">S.</emphasis>
<emphasis id="B907EAB62A38FFE0FF72F6254A27F655" box="[194,302,2471,2510]" italics="true" pageId="11" pageNumber="12">broccae</emphasis>
</taxonomicName>
were mapped by
<bibRefCitation id="EFE24B552A38FFE0FDE8F6254816F655" author="Sarraf, A." box="[600,799,2471,2510]" pageId="11" pageNumber="12" pagination="27 - 30" refId="ref30602" refString="Sarraf, A. 1997. Redescription and distribution of Spintherobolus broccae Myers (Characiformes: Cha - racidae). Rev. Fr. Aquariol. Herpet., 24: 27 - 30." type="journal article" year="1997">Sarraf (1997</bibRefCitation>
: figs. 2-4), but their terminology equivalents to these lines in other fishes, for example in
<bibRefCitation id="EFE24B552A38FFE0FD39F5824F57FF60" author="Arratia, G. &amp; L. Huaquin" box="[649,1118,212,2600]" pageId="11" pageNumber="12" pagination="1 - 108" refId="ref28749" refString="Arratia, G. &amp; L. Huaquin. 1995. Morphology of the lateral line system and of the skin of diplomystid and certain primitive loricarioid catfishes and systematic and ecological considerations. Bonn. Zool. Monogr., 36: 1 - 108." type="journal article" year="1995">Arratia &amp; Huaquin (1995</bibRefCitation>
: fig. 13) were not attempted. The neuromast lines of the species of
<taxonomicName id="4C734D272A38FFE0F9B4FE824DEDFEB3" authorityName="Eigenmann" authorityYear="1911" box="[1540,1764,256,296]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0F9B4FE824DEDFEB3" box="[1540,1764,256,296]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
are quite complex and an accurate comparative study of their homology and the nomenclatural equivalency of these lines with those of other characins, catfishes and other teleosts would require a comparative study of the nerve branches serving their neuromasts, a study beyond the scope of the present work. Such a study would perhaps be a useful source of synapomorphies suggesting a phylogeny of the species of
<emphasis id="B907EAB62A38FFE0FA48FD144DE8FD25" box="[1528,1761,662,702]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C734D272A38FFE0FA48FD144DD5FD25" authorityName="Eigenmann" authorityYear="1911" box="[1528,1756,662,702]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
but we hesitate to use the neuromast lines of
<taxonomicName id="4C734D272A38FFE0F91CFD404F66FC83" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0F91CFD404F66FC83" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
for this purpose without a detailed study of their anatomy and homology.
</paragraph>
<paragraph id="8BCC36A42A38FFE0FB8EFCC84D51F709" blockId="11.[1021,1836,212,2601]" pageId="11" pageNumber="12">
The neuromasts of
<taxonomicName id="4C734D272A38FFE0FA24FCC84D7AFCE9" authorityName="Eigenmann" authorityYear="1911" box="[1428,1651,842,882]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A38FFE0FA24FCC84D7AFCE9" box="[1428,1651,842,882]" italics="true" pageId="11" pageNumber="12">Spintherobolus</emphasis>
</taxonomicName>
were first identified as “tactile papillae” by
<bibRefCitation id="EFE24B552A38FFE0F9D9FCFB4FB1FC57" author="Eigenmann, C. H." pageId="11" pageNumber="12" pagination="164 - 181" refId="ref29012" refString="Eigenmann, C. H. 1911. New characins in the collections of the Carnegie Museum. Ann. Carnegie Mus., 8: 164 - 181." type="journal article" year="1911">Eigenmann (1911: 167)</bibRefCitation>
. We examined the surface of these organs at 200 x but have no histological evidence to confirm that these are neuromasts. However, these organs are similar in gross anatomy to the neuromasts described for
<taxonomicName id="4C734D272A38FFE0FA1AFBD94DE8FB19" baseAuthorityName="De Filippi" baseAuthorityYear="1853" box="[1450,1761,1115,1154]" family="Characidae" genus="Astyanax" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="mexicanus">
<emphasis id="B907EAB62A38FFE0FA1AFBD94DE8FB19" box="[1450,1761,1115,1154]" italics="true" pageId="11" pageNumber="12">Astyanax mexicanus</emphasis>
</taxonomicName>
and histologically examined by
<bibRefCitation id="EFE24B552A38FFE0FA45FB044C16FB34" author="Schemmel, C." box="[1525,1823,1158,1199]" pageId="11" pageNumber="12" pagination="255 - 316" refId="ref30702" refString="Schemmel, C. 1967. Vergleichende Untersuchungen an den Hautsinnesorganen ober- und unterirdisch lebender As ty anax- Formen - ein Beitrag zur Evolution der Cavernicolen. Ztschr. Morph. Tiere, 61: 255 - 316." type="journal article" year="1967">Schemmel (1967)</bibRefCitation>
. They are found distributed in a manner somewhat similar to those figured by
<bibRefCitation id="EFE24B552A38FFE0F9A6FB624F77FAAC" author="Schemmel, C." pageId="11" pageNumber="12" pagination="255 - 316" refId="ref30702" refString="Schemmel, C. 1967. Vergleichende Untersuchungen an den Hautsinnesorganen ober- und unterirdisch lebender As ty anax- Formen - ein Beitrag zur Evolution der Cavernicolen. Ztschr. Morph. Tiere, 61: 255 - 316." type="journal article" year="1967">Schemmel (1967: 266-267</bibRefCitation>
, figs. 4-5) along body scales of this species. In
<emphasis id="B907EAB62A38FFE0FB22FABE4E74FAFF" box="[1170,1405,1340,1380]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C734D272A38FFE0FB22FABE4E71FAFF" authorityName="Eigenmann" authorityYear="1911" box="[1170,1400,1340,1380]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
these organs are small, rounded, slightly elevated discs that are translucent or have a brown,grange, or red, color. The color intensity varies from collection to collection, ranging from no perceptible color to a relatively dark brown or red. The color possibly depends upon length of time in and the conditions of the preservative. It is also possible that the color is an environmental contaminant because specimens of some collections that have brown or red sediment caught in the mucus around their scales and fins are the ones that most often have colored neuromasts. The disc’ s color, when present, is due to several small, circular areas of color at the surface of the neuromast. These spots of color appear to correspond to sensory cells, each probably bearing a sensory filament. However, sensory filaments or cupulae are not detectable under a dissecting microscope.
</paragraph>
<paragraph id="8BCC36A42A38FFE7FB89F714498AF990" blockId="11.[1021,1836,212,2601]" lastBlockId="12.[78,892,198,2541]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
Many of the neuromasts are located in shallow epidermal channels of
<emphasis id="B907EAB62A38FFE0FA39F7464D79F777" box="[1417,1648,2244,2284]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C734D272A38FFE0FA39F7464D62F777" authorityName="Eigenmann" authorityYear="1911" box="[1417,1643,2244,2284]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
especially on the head. The distribution pattern of these organs is approximately the same in all four species except that there are more vertical channels on the cheek (infraorbital region) in
<taxonomicName id="4C734D272A38FFE0F9E6F6FB4C2DF639" authorityName="Eigenmann" authorityYear="1911" box="[1622,1828,2425,2466]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A38FFE0F9E6F6FB4D66F639" box="[1622,1647,2425,2466]" italics="true" pageId="11" pageNumber="12">S.</emphasis>
<emphasis id="B907EAB62A38FFE0F9CBF6FB4C2DF639" box="[1659,1828,2425,2466]" italics="true" pageId="11" pageNumber="12">papilliferus</emphasis>
</taxonomicName>
than in the other species (
<figureCitation id="13482A212A38FFE0FA70F6244D78F654" box="[1472,1649,2470,2511]" captionStart-0="Fig" captionStart-1="Fig" captionStartId-0="9.[170,218,740,776]" captionStartId-1="9.[169,217,1134,1180]" captionTargetBox-0="[167,836,218,695]" captionTargetBox-1="[848,1844,271,681]" captionTargetPageId-0="9" captionTargetPageId-1="9" captionText-0="Fig. 10. Spintherobolus papilliferus, MNRJ 4237, female, 34.7 mm SL, head region, left side, Brazil, São Paulo, rio Ipiranga. Rows of neuromasts occur over the infraorbital, preopercular, and opercular region. These are distributed in shallow grooves that can be seen in the photograph. The neuromasts are also distributed on the scale borders posterior to the head and operculum but these are hard to distinguish from dark chromatophores in the photograph. A row of neuromasts along the ventral border of the dentary is hidden in a deep groove. The small dark dots arranged in rows are the neuromast patterns over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region." captionText-1="Fig. 11. Spintherobolus papilliferus, MZUSP 51021, female 60.8 mm SL. Dorsum of head and back nearly to dorsal-fin origin. Neuromasts are present in the irregular grooves between the eyes and the supraoccipital region but these are nearly impossible to see in the photograph. A bilaterally distributed row of neuromasts runs along the long axis of the body between the nares. This row of neuromasts shows more clearly on the left side than on the right side of the top of the head." figureDoi-0="http://doi.org/10.5281/zenodo.10882016" figureDoi-1="http://doi.org/10.5281/zenodo.10882018" httpUri-0="https://zenodo.org/record/10882016/files/figure.png" httpUri-1="https://zenodo.org/record/10882018/files/figure.png" pageId="11" pageNumber="12">Figs. 10-11</figureCitation>
). Laterally on the cheek a longitudinal curved neuromast channel extends from the posterior border of maxilla to the posterior margin of preopercle. From this channel, at intervals, extend more or less vertical channels, each dorsally reaching the ventral margin of the orbit, and ventrally reaching to near the ventral border of the preopercle. The opercle has an anterior, vertical linear channel near the posterior preopercular border. Posterior to this the opercle bears a series of shallow channels containing neuromasts. Dorsally on the head these organs are mainly distributed in somewhat irregular transverse grooves between the orbits and in two small longitudinal channels between the nares (
<figureCitation id="13482A212A3FFFE7FE13FD604907FC90" box="[419,526,738,779]" captionStart="Fig" captionStartId="9.[169,217,1134,1180]" captionTargetBox="[848,1844,271,681]" captionTargetPageId="9" captionText="Fig. 11. Spintherobolus papilliferus, MZUSP 51021, female 60.8 mm SL. Dorsum of head and back nearly to dorsal-fin origin. Neuromasts are present in the irregular grooves between the eyes and the supraoccipital region but these are nearly impossible to see in the photograph. A bilaterally distributed row of neuromasts runs along the long axis of the body between the nares. This row of neuromasts shows more clearly on the left side than on the right side of the top of the head." figureDoi="http://doi.org/10.5281/zenodo.10882018" httpUri="https://zenodo.org/record/10882018/files/figure.png" pageId="12" pageNumber="13">Fig. 11</figureCitation>
). There is a well-developed ventrally facing skin flap extending posteriorly from the area between the dentary and maxilla to the posteroventral end of the preoper ­ cle. This flap partly covers an elongate groove running along its length. The groove with its neuromasts has several medially directed branches. The dentary bears ventral grooves containing neuromasts, these being numerous in the gular area between the two halves of the dentary. Two to six vertically aligned neuromasts are located near the posterior border of many of the body scales. Although many of the scales bear neuromasts, they are not present on all scales. We were unable to detect any repetitive pattern of distribution of neuromasts o^ the body scales. See
<bibRefCitation id="EFE24B552A3FFFE7FFE0FA3A4AF2FA44" author="Arratia, G. &amp; L. Huaquin" box="[80,507,1464,1503]" pageId="12" pageNumber="13" pagination="1 - 108" refId="ref28749" refString="Arratia, G. &amp; L. Huaquin. 1995. Morphology of the lateral line system and of the skin of diplomystid and certain primitive loricarioid catfishes and systematic and ecological considerations. Bonn. Zool. Monogr., 36: 1 - 108." type="journal article" year="1995">Arratia &amp; Huaquin (1995)</bibRefCitation>
for citation of much of the literature on fish neuromasts.
</paragraph>
<paragraph id="8BCC36A42A3FFFE7FF3BF9934A59F676" blockId="12.[78,892,198,2541]" pageId="12" pageNumber="13">
What appear to be similar organs, also with complex patterns of distribution on the head, accompanied by an absence of laterosensory canals, were described by
<bibRefCitation id="EFE24B552A3FFFE7FE41F91B4866F924" author="Poll, M. &amp; J. Lambert" box="[497,879,1689,1727]" pageId="12" pageNumber="13" pagination="336 - 340" refId="ref30351" refString="Poll, M., &amp; J. Lambert. 1964 a. Congocharax gossei sp. n. du Gabon (Ogoou e) (Pisces, Citharinidae). Rev. Zool. Bot. Afr., 70: 336 - 340." type="journal article" year="1964">Poll &amp; Lambert (1964a</bibRefCitation>
: 337,1964b: 407) for certain species of the African citharinid characiform
<taxonomicName id="4C734D272A3FFFE7FE59F9704B95F8D2" authority="Steindachner, 1894" authorityName="Steindachner" authorityYear="1894" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FE59F9704972F881" box="[489,635,1778,1818]" italics="true" pageId="12" pageNumber="13">Neolebias</emphasis>
Steindachner, 1894
</taxonomicName>
. These organs were identified as “lignes d’organes sensoriels ponctiformes”. Note: Although Poll &amp; Lambert used
<taxonomicName id="4C734D272A3FFFE7FDE5F8F94BE7F84B" authority="Matthes, 1964" authorityName="Matthes" authorityYear="1964" family="Distichodontidae" genus="Congocharax" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FDE5F8F94813F83F" box="[597,794,1915,1956]" italics="true" pageId="12" pageNumber="13">Congocharax</emphasis>
Matthes, 1964
</taxonomicName>
for the generic name of those species of
<taxonomicName id="4C734D272A3FFFE7FFFFF8564BD7F866" authorityName="Steindachner" authorityYear="1894" box="[79,222,2004,2045]" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FFFFF8564BD7F866" box="[79,222,2004,2045]" italics="true" pageId="12" pageNumber="13">Neolebias</emphasis>
</taxonomicName>
readily showing the neuromasts, we follow
<bibRefCitation id="EFE24B552A3FFFE7FF70F7814AB0F7B1" author="Vari, R. P." box="[192,441,2051,2090]" pageId="12" pageNumber="13" pagination="261 - 344" refId="ref30871" refString="Vari, R. P. 1979. Anatomy, relationships and classifica - tion of the families Citharinidae and Distichodonti - dae (Pisces, Characoidea). Bull. Br. Mus. Nat. Hist., Zool., 36: 261 - 344." type="journal article" year="1979">Vari (1979: 330)</bibRefCitation>
in recognizing
<taxonomicName id="4C734D272A3FFFE7FD05F7814870F7B1" authorityName="Matthes" authorityYear="1964" box="[693,889,2051,2090]" family="Distichodontidae" genus="Congocharax" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FD05F7814870F7B1" box="[693,889,2051,2090]" italics="true" pageId="12" pageNumber="13">Congocharax</emphasis>
</taxonomicName>
as a junior synonym of
<taxonomicName id="4C734D272A3FFFE7FE4EF7AD4984F7CC" authorityName="Steindachner" authorityYear="1894" box="[510,653,2095,2135]" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FE4EF7AD4984F7CC" box="[510,653,2095,2135]" italics="true" pageId="12" pageNumber="13">Neolebias</emphasis>
</taxonomicName>
even though
<bibRefCitation id="EFE24B552A3FFFE7FFE0F7DF4AB6F71F" author="Poll, M. &amp; J. P. Gosse" box="[80,447,2141,2180]" pageId="12" pageNumber="13" pagination="1 - 8" refId="ref30281" refString="Poll, M. &amp; J. P. Gosse. 1982. Rehabilitation des genres Congocharax Matthes, 1964 et Dundocharax Poll, 1967 (Pisces, Distichodontidae) mis en synonymie par R. P. Vari, 1979, avec Neolebias Steindachner, 1894. Bull. Inst. R. Sci. Nat. Belg., 54 (2): 1 - 8." type="journal article" year="1982">Poll &amp; Gosse (1982: 5)</bibRefCitation>
provided ample evidence of the different features that can be used to distinguish these nominal genera. We do this because recognition of
<taxonomicName id="4C734D272A3FFFE7FE73F7664983F690" authorityName="Matthes" authorityYear="1964" box="[451,650,2276,2315]" family="Distichodontidae" genus="Congocharax" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FE73F7664983F690" box="[451,650,2276,2315]" italics="true" pageId="12" pageNumber="13">Congocharax</emphasis>
</taxonomicName>
would make
<taxonomicName id="4C734D272A3FFFE7FFFFF6924BD2F6A2" authorityName="Steindachner" authorityYear="1894" box="[79,219,2320,2361]" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FFFFF6924BD2F6A2" box="[79,219,2320,2361]" italics="true" pageId="12" pageNumber="13">Neolebias</emphasis>
</taxonomicName>
paraphyletic according to the phylogenetic evidence provided by Vari and not contradicted by additional data extracted from Poll &amp; Gosse and cladistically analyzed along with the data from Vari.
</paragraph>
<paragraph id="8BCC36A42A3FFFE7FC40FF454D44F976" blockId="12.[944,1753,199,2585]" pageId="12" pageNumber="13">
As pointed out by
<bibRefCitation id="EFE24B552A3FFFE7FA8DFF454D58FF76" author="Sarraf, A." box="[1341,1617,199,237]" pageId="12" pageNumber="13" pagination="27 - 30" refId="ref30602" refString="Sarraf, A. 1997. Redescription and distribution of Spintherobolus broccae Myers (Characiformes: Cha - racidae). Rev. Fr. Aquariol. Herpet., 24: 27 - 30." type="journal article" year="1997">Sarraf (1997: 30)</bibRefCitation>
, similar organs are found in the species of
<taxonomicName id="4C734D272A3FFFE7F9B2FF714DD1FE80" authorityName="Eigenmann" authorityYear="1907" box="[1538,1752,243,283]" family="Characidae" genus="Phenacogaster" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7F9B2FF714DD1FE80" box="[1538,1752,243,283]" italics="true" pageId="12" pageNumber="13">Phenacogaster</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3FFFE7FC4EFEA24E27FEDD" authority="Gunther." authorityName="Gunther." box="[1022,1326,288,326]" family="Characidae" genus="Roeboides" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FC4EFEA24F99FEDD" box="[1022,1168,288,326]" italics="true" pageId="12" pageNumber="13">Roeboides</emphasis>
Gunther.
</taxonomicName>
These were called “cervical pit-lines” by
<bibRefCitation id="EFE24B552A3FFFE7FB50FECE4D15FEEE" author="Géry, J." box="[1248,1564,332,373]" journalOrPublisher="Stud. Neotop. Fauna" pageId="12" pageNumber="13" pagination="1 - 35" part="7" refId="ref29360" refString="- 1972. Corrected and supplemental descriptions of certain characoid fishes described by Henry W. Fowler, with revisions of several of their genera. Stud. Neotop. Fauna, 7: 1 - 35." title="Corrected and supplemental descriptions of certain characoid fishes described by Henry W. Fowler, with revisions of several of their genera" type="journal article" year="1972">Géry (1972: 10-11)</bibRefCitation>
. Although cervical is a word referring to a neck, these struc ­ tures are located in shallow channels mostly on the head and putatively assigned as a synapomorphy uniting these two genera by
<bibRefCitation id="EFE24B552A3FFFE7F99AFD834E1FFDCE" author="Malabarba, L. R. &amp; Z. M. S. Lucena" pageId="12" pageNumber="13" pagination="337 - 344" refId="ref29810" refString="Malabarba, L. R. &amp; Z. M. S. Lucena. 1995. Phenacogaster jancupa, new species, with comments on the relationships and a new diagnosis of the genus (Ostar - iophysi: Characidae). Ichthyol. Explor. Freshwaters, 6: 337 - 344." type="journal article" year="1995">Malabarba &amp; Lucena (1995: 342)</bibRefCitation>
. We examined these struc ­ tures in species of these two genera and found they have the same apparent gross anatomy and sometimes the same color characteristics as those of
<emphasis id="B907EAB62A3FFFE7FC50FD614FC0FC91" box="[992,1225,739,778]" italics="true" pageId="12" pageNumber="13">
<taxonomicName id="4C734D272A3FFFE7FC50FD614FCAFC91" authorityName="Eigenmann" authorityYear="1911" box="[992,1219,739,778]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
but with different distributional patterns on the head and body. We agree with
<bibRefCitation id="EFE24B552A3FFFE7FC02FCBF4F83FCFE" author="Sarraf, A." box="[946,1162,829,869]" pageId="12" pageNumber="13" pagination="27 - 30" refId="ref30602" refString="Sarraf, A. 1997. Redescription and distribution of Spintherobolus broccae Myers (Characiformes: Cha - racidae). Rev. Fr. Aquariol. Herpet., 24: 27 - 30." type="journal article" year="1997">Sarraf (1997)</bibRefCitation>
that these patterns are independently derived in
<taxonomicName id="4C734D272A3FFFE7FB64FCEB4EBAFC09" authorityName="Eigenmann" authorityYear="1911" box="[1236,1459,873,914]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FB64FCEB4EBAFC09" box="[1236,1459,873,914]" italics="true" pageId="12" pageNumber="13">Spintherobolus</emphasis>
</taxonomicName>
on the one hand and
<taxonomicName id="4C734D272A3FFFE7FC4CFC154FDDFC5B" authorityName="Eigenmann" authorityYear="1907" box="[1020,1236,919,960]" family="Characidae" genus="Phenacogaster" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FC4CFC154FDDFC5B" box="[1020,1236,919,960]" italics="true" pageId="12" pageNumber="13">Phenacogaster</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3FFFE7FA97FC154EB1FC5B" box="[1319,1464,919,960]" family="Characidae" genus="Roeboides" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FA97FC154EB1FC5B" box="[1319,1464,919,960]" italics="true" pageId="12" pageNumber="13">Roeboides</emphasis>
</taxonomicName>
on the other. The surface neuromasts of the laterosensory system and their characteristic distributional patterns found in some species of
<emphasis id="B907EAB62A3FFFE7FAFFFBA24EECFBDC" box="[1359,1509,1056,1095]" italics="true" pageId="12" pageNumber="13">
<taxonomicName id="4C734D272A3FFFE7FAFFFBA24EE9FBDC" authorityName="Steindachner" authorityYear="1894" box="[1359,1504,1056,1095]" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Neolebias</taxonomicName>
,
</emphasis>
represent similar but apparently also independently derived features from those present in
<taxonomicName id="4C734D272A3FFFE7FA77FBFA4DADFB3A" authorityName="Eigenmann" authorityYear="1911" box="[1479,1700,1144,1185]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FA77FBFA4DADFB3A" box="[1479,1700,1144,1185]" italics="true" pageId="12" pageNumber="13">Spintherobolus</emphasis>
</taxonomicName>
or
<taxonomicName id="4C734D272A3FFFE7FC02FB254F82FB56" authorityName="Eigenmann" authorityYear="1907" box="[946,1163,1191,1229]" family="Characidae" genus="Phenacogaster" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FC02FB254F82FB56" box="[946,1163,1191,1229]" italics="true" pageId="12" pageNumber="13">Phenacogaster</emphasis>
</taxonomicName>
and
<emphasis id="B907EAB62A3FFFE7FB59FB254E8DFB56" box="[1257,1412,1191,1229]" italics="true" pageId="12" pageNumber="13">
<taxonomicName id="4C734D272A3FFFE7FB59FB254E76FB56" box="[1257,1407,1191,1229]" family="Characidae" genus="Roeboides" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Roeboides</taxonomicName>
.
</emphasis>
As
<bibRefCitation id="EFE24B552A3FFFE7FA60FB254DDFFB56" author="Vari, R. P." box="[1488,1750,1191,1229]" pageId="12" pageNumber="13" pagination="261 - 344" refId="ref30871" refString="Vari, R. P. 1979. Anatomy, relationships and classifica - tion of the families Citharinidae and Distichodonti - dae (Pisces, Characoidea). Bull. Br. Mus. Nat. Hist., Zool., 36: 261 - 344." type="journal article" year="1979">Vari (1979: 329)</bibRefCitation>
has shown, the species of
<taxonomicName id="4C734D272A3FFFE7FAC9FB504D0EFB60" authorityName="Steindachner" authorityYear="1894" box="[1401,1543,1234,1275]" family="Distichodontidae" genus="Neolebias" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FAC9FB504D0EFB60" box="[1401,1543,1234,1275]" italics="true" pageId="12" pageNumber="13">Neolebias</emphasis>
</taxonomicName>
bearing apparent neuromasts have a derived position in a clade of African characiforms. The exposed neuromasts described for
<taxonomicName id="4C734D272A3FFFE7FAFCFADC4D91FA1E" baseAuthorityName="De Filippi" baseAuthorityYear="1853" box="[1356,1688,1374,1413]" family="Characidae" genus="Astyanax" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="mexicanus">
<emphasis id="B907EAB62A3FFFE7FAFCFADC4EE8FA1E" box="[1356,1505,1374,1413]" italics="true" pageId="12" pageNumber="13">Astyanax</emphasis>
<emphasis id="B907EAB62A3FFFE7FA46FADC4D91FA1E" box="[1526,1688,1374,1413]" italics="true" pageId="12" pageNumber="13">mexicanus</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFE24B552A3FFFE7FC02FA0B4FC3FA2A" author="Schemmel, C." box="[946,1226,1417,1457]" pageId="12" pageNumber="13" pagination="255 - 316" refId="ref30702" refString="Schemmel, C. 1967. Vergleichende Untersuchungen an den Hautsinnesorganen ober- und unterirdisch lebender As ty anax- Formen - ein Beitrag zur Evolution der Cavernicolen. Ztschr. Morph. Tiere, 61: 255 - 316." type="journal article" year="1967">Schemmel (1967)</bibRefCitation>
seems to be also independently derived from those present in
<taxonomicName id="4C734D272A3FFFE7FA27FA344D7AFA44" authorityName="Eigenmann" authorityYear="1911" box="[1431,1651,1462,1503]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FA27FA344D7AFA44" box="[1431,1651,1462,1503]" italics="true" pageId="12" pageNumber="13">Spintherobolus</emphasis>
</taxonomicName>
(compare the neuromast distributional pattern of
<taxonomicName id="4C734D272A3FFFE7FC02F9934F76F9A3" baseAuthorityName="De Filippi" baseAuthorityYear="1853" box="[946,1151,1553,1592]" family="Characidae" genus="Astyanax" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="mexicanus">
<emphasis id="B907EAB62A3FFFE7FC02F99348DAF9A3" box="[946,979,1553,1592]" italics="true" pageId="12" pageNumber="13">A.</emphasis>
<emphasis id="B907EAB62A3FFFE7FC51F9934F76F9A3" box="[993,1151,1553,1592]" italics="true" pageId="12" pageNumber="13">mexicanus</emphasis>
</taxonomicName>
in
<bibRefCitation id="EFE24B552A3FFFE7FB0BF9934D11F9A3" author="Schemmel, C." box="[1211,1560,1553,1592]" pageId="12" pageNumber="13" pagination="255 - 316" refId="ref30702" refString="Schemmel, C. 1967. Vergleichende Untersuchungen an den Hautsinnesorganen ober- und unterirdisch lebender As ty anax- Formen - ein Beitrag zur Evolution der Cavernicolen. Ztschr. Morph. Tiere, 61: 255 - 316." type="journal article" year="1967">Schemmel (1967: 267</bibRefCitation>
, fig. 5) and of
<taxonomicName id="4C734D272A3FFFE7FC6BF9BC4FB1F9FD" box="[987,1208,1598,1638]" pageId="12" pageNumber="13">
<emphasis id="B907EAB62A3FFFE7FC6BF9BC4FB1F9FD" box="[987,1208,1598,1638]" italics="true" pageId="12" pageNumber="13">Spintherobolus</emphasis>
</taxonomicName>
species; Figs. 4,10). None of these outgroup characiform taxa bearing obvious neuromasts and discussed above have all the synapomorphies listed for the
<taxonomicName id="4C734D272A3FFFE7FAF5F9464D4CF976" box="[1349,1605,1732,1773]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="14" phylum="Chordata" rank="subFamily" subFamily="Cheirodontinae">Cheirodontinae</taxonomicName>
.
</paragraph>
<paragraph id="8BCC36A42A3FFFE7FC5EF9734FC8F582" blockId="12.[944,1753,199,2585]" pageId="12" pageNumber="13">
<bibRefCitation id="EFE24B552A3FFFE7FC5EF9734E0FF880" author="Bohlke, J. E." box="[1006,1286,1777,1819]" pageId="12" pageNumber="13" refId="ref28796" refString="Bohlke, J. E. 1954. Studies on the phylogeny and systematics of fishes of the family Characidae. Unpubl. Dissertation, University Microfilms, Ann Arbor, v + 209 pp." type="book" year="1954">Böhlke (1954: 25)</bibRefCitation>
reported “rows of papillae” on the head of
<taxonomicName id="4C734D272A3FFFE7FB01F89D4ED3F8D3" authorityName="Gunther" authorityYear="1863" box="[1201,1498,1823,1864]" family="Crenuchidae" genus="Crenuchus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="spilurus">
<emphasis id="B907EAB62A3FFFE7FB01F89D4ED3F8D3" box="[1201,1498,1823,1864]" italics="true" pageId="12" pageNumber="13">Crenuchus spilurus</emphasis>
</taxonomicName>
and suggested this genus should be examined when consider ­ ing the relationships of either
<taxonomicName id="4C734D272A3FFFE7FA75F8F84DAAF839" authorityName="Eigenmann" authorityYear="1911" box="[1477,1699,1914,1954]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7FA75F8F84DAAF839" box="[1477,1699,1914,1954]" italics="true" pageId="12" pageNumber="13">Spintherobolus</emphasis>
</taxonomicName>
or
<emphasis id="B907EAB62A3FFFE7FC03F8254F6BF855" box="[947,1122,1959,1998]" italics="true" pageId="12" pageNumber="13">
<taxonomicName id="4C734D272A3FFFE7FC03F8254F53F855" box="[947,1114,1959,1998]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">Grundulus</taxonomicName>
.
</emphasis>
We have examined the heads of several specimens of
<taxonomicName id="4C734D272A3FFFE7FB66F8544E72F867" authorityName="Gunther" authorityYear="1863" box="[1238,1403,2006,2044]" family="Crenuchidae" genus="Crenuchus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="spilurus">
<emphasis id="B907EAB62A3FFFE7FB66F8544E72F867" box="[1238,1403,2006,2044]" italics="true" pageId="12" pageNumber="13">C. spilurus</emphasis>
</taxonomicName>
and fail to find struc ­ tures that we can confidently identify as being similar to the organs we tentatively identify as neuromasts. There are, however, scattered small papillose structures having a central depression or opening. These might be taste buds, but histological examination is needed to confirm such an identification. The Crenuchinae is phylogenetically related to the Characidiinae and both placed in the
<taxonomicName id="4C734D272A3FFFE7FB9CF6E84FF6F608" box="[1068,1279,2410,2451]" family="Crenuchidae" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Crenuchidae</taxonomicName>
by
<bibRefCitation id="EFE24B552A3FFFE7FAFEF6E84D96F608" author="Buckup, P. A." box="[1358,1695,2410,2451]" pageId="12" pageNumber="13" pagination="225 - 245" refId="ref28832" refString="Buckup, P. A. 1993 a. The monophyly of the Characidiinae, a neotropical group of characiform fishes (Teleostei: Ostariophysi). Zool. J. Linn. Soc., 108: 225 - 245." type="journal article" year="1993">Buckup (1993a: 229</bibRefCitation>
; in press) and we found no evidence to contradict Buckup ’ s hypothesis or suggest that
<taxonomicName id="4C734D272A3FFFE7F9A2F6464DBAF675" authorityName="Günther" authorityYear="1863" box="[1554,1715,2500,2542]" family="Crenuchidae" genus="Crenuchus" kingdom="Animalia" order="Characiformes" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3FFFE7F9A2F6464DBAF675" box="[1554,1715,2500,2542]" italics="true" pageId="12" pageNumber="13">Crenuchus</emphasis>
</taxonomicName>
is a cheirodontine.
</paragraph>
<caption id="DF0C662C2A3EFFE6FF0FFB5D4836FAEE" ID-DOI="http://doi.org/10.5281/zenodo.10882022" ID-Zenodo-Dep="10882022" httpUri="https://zenodo.org/record/10882022/files/figure.png" pageId="13" pageNumber="14" startId="13.[191,252,1247,1291]" targetBox="[187,1269,268,1188]" targetPageId="13" targetType="figure">
<paragraph id="8BCC36A42A3EFFE6FF0FFB5D4836FAEE" blockId="13.[188,1265,1247,1397]" pageId="13" pageNumber="14">
Fig. 13.
<taxonomicName id="4C734D272A3EFFE6FED5FB5D4824FA90" box="[357,813,1247,1291]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A3EFFE6FED5FB5D4964FA90" box="[357,621,1247,1291]" italics="true" pageId="13" pageNumber="14">Spintherobolus</emphasis>
<emphasis id="B907EAB62A3EFFE6FD31FB5D4824FA90" box="[641,813,1247,1291]" italics="true" pageId="13" pageNumber="14">ankoseion</emphasis>
</taxonomicName>
, infraorbital bones. a, USNM 297935, adult female, 24.6 mm SL; b, USNM 297935, adult male, 24.5 mm SL.
</paragraph>
</caption>
<paragraph id="8BCC36A42A3EFFE6FEBCFA7D4A86F61B" blockId="13.[185,1265,1535,3456]" pageId="13" pageNumber="14">
(16) The dentary has a large anterior fenestra (
<figureCitation id="13482A212A3EFFE6FF7DF9B94A69F9EB" box="[205,352,1595,1648]" captionStart="Fig" captionStartId="10.[933,981,1063,1097]" captionTargetBox="[933,1740,197,1020]" captionTargetPageId="10" captionText="Fig. 12. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Jaws &amp; teeth." figureDoi="http://doi.org/10.5281/zenodo.10882020" httpUri="https://zenodo.org/record/10882020/files/figure.png" pageId="13" pageNumber="14">Fig. 12</figureCitation>
). Associated with this fenestra is a large epidermal, papilla-like structure surrounded by a deep groove that has its deep internal portion lodged in the dentary fenestra. The external sur ­ face of this papilla bears several exposed neuromasts. The ventral face of the dentary bone, posterior to the fenestra is concave (more so in
<taxonomicName id="4C734D272A3EFFE6FF0DF8664AC4F783" authorityName="Eigenmann" authorityYear="1911" box="[189,461,2020,2072]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="39" phylum="Chordata" rank="species" species="papilliferus">
S.
<emphasis id="B907EAB62A3EFFE6FF5FF8664AC4F783" box="[239,461,2020,2072]" italics="true" pageId="13" pageNumber="14">papilliferus</emphasis>
</taxonomicName>
than the other species; see also the dentary characters noted below under the synapomorphies that hypothesize subgroup b in Fig ­ ure 3). We suggest the fenestra and the associated nervous and supporting tissues are a derived sensory organ and consider all to be one complex character.
</paragraph>
<paragraph id="8BCC36A42A3EFFE6FEB9F60B4802F4BE" blockId="13.[185,1265,1535,3456]" pageId="13" pageNumber="14">
<bibRefCitation id="EFE24B552A3EFFE6FEB9F60B49D2F627" author="Travassos, H." box="[265,731,2441,2492]" journalOrPublisher="An. Acad. Bras. Ci e nc." pageId="13" pageNumber="14" pagination="505 - 517" part="25" refId="ref30826" refString="- 1953. Fauna do Distrito Federal. III. S o bre o g e nero &quot; Spintherobolus &quot; Eigenmann, 1911 (Cypriniformes- Characoidei). An. Acad. Bras. Ci e nc., 25: 505 - 517." title="Fauna do Distrito Federal. III. S o bre o g e nero &quot; Spintherobolus &quot; Eigenmann, 1911 (Cypriniformes- Characoidei)" type="journal article" year="1953">Travassos (1953: 506)</bibRefCitation>
first noted the fenestra, but not the associated modified soft tissue. He considered the fenestra to demonstrate a close relationship between
<taxonomicName id="4C734D272A3EFFE6FD27F5BC4852F5E9" box="[663,859,2622,2674]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A3EFFE6FD27F5BC49BEF5E9" box="[663,695,2622,2674]" italics="true" pageId="13" pageNumber="14">S.</emphasis>
<emphasis id="B907EAB62A3EFFE6FD7BF5BC4852F5E9" box="[715,859,2622,2674]" italics="true" pageId="13" pageNumber="14">broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3EFFE6FC7DF5BC4FE8F5E9" authorityName="Eigenmann" authorityYear="1911" box="[973,1249,2622,2674]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3EFFE6FC7DF5BC48E6F5E9" box="[973,1007,2622,2674]" italics="true" pageId="13" pageNumber="14">S.</emphasis>
<emphasis id="B907EAB62A3EFFE6FBB0F5BC4FE4F5E9" box="[1024,1261,2622,2674]" italics="true" pageId="13" pageNumber="14">papilliferus.</emphasis>
</taxonomicName>
All four species examined herein have this feature, which is absent in other cheirodontines or characids examined by us.
</paragraph>
<paragraph id="8BCC36A42A3EFFE6FEBBF4AF4257FE1B" blockId="13.[185,1265,1535,3456]" lastBlockId="13.[1339,2422,271,3456]" pageId="13" pageNumber="14">
(17) The adipose fin is absent. The adipose fin loss in
<taxonomicName id="4C734D272A3EFFE6FE71F4EB49E4F406" authorityName="Eigenmann" authorityYear="1911" box="[449,749,2921,2973]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FE71F4EB49E4F406" box="[449,749,2921,2973]" italics="true" pageId="13" pageNumber="14">Spintherobolus</emphasis>
</taxonomicName>
is unique within the monophyletic subfamily
<taxonomicName id="4C734D272A3EFFE6FD5FF4274F4EF441" box="[751,1095,2981,3034]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="subFamily" subFamily="Cheirodontinae">Cheirodontinae</taxonomicName>
as rec ­ ognized by us; see also Malabarba (in press). We suggest the absence of an adipose fin in
<taxonomicName id="4C734D272A3EFFE6FB33F3A24A8DF315" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FB33F3A24A8DF315" italics="true" pageId="13" pageNumber="14">Spintherobolus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3EFFE6FE4FF3DE49D0F315" box="[511,729,3164,3214]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FE4FF3DE49D0F315" box="[511,729,3164,3214]" italics="true" pageId="13" pageNumber="14">Grundulus</emphasis>
</taxonomicName>
is homoplastic because of the relationships hypothesized here for
<emphasis id="B907EAB62A3EFFE6FB33F31A4A99F292" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="4C734D272A3EFFE6FB33F31A4A85F292" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
<bibRefCitation id="EFE24B552A3EFFE6FE1DF357487CF292" author="Eigenmann, C. H." bookContentInfo="pls. 1 - 17" box="[429,885,3285,3337]" journalOrPublisher="Mem. Carnegie Mus." pageId="13" pageNumber="14" pagination="1 - 99" part="7" refId="ref29042" refString="- 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Mem. Carnegie Mus., 7: 1 - 99, pls. 1 - 17." title="The Cheirodontinae, a subfamily of minute characid fishes of South America" type="journal article" year="1915">Eigenmann (1915: 4</bibRefCitation>
, 14) briefly discussed the lack of an adipose fin in
<taxonomicName id="4C734D272A3EFFE6FBA0F2924FE5F2DF" box="[1040,1260,3344,3396]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FBA0F2924FE5F2DF" box="[1040,1260,3344,3396]" italics="true" pageId="13" pageNumber="14">Grundulus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3EFFE6FEAFF2CE494CF21B" authorityName="Eigenmann" authorityYear="1911" box="[287,581,3404,3456]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FEAFF2CE494CF21B" box="[287,581,3404,3456]" italics="true" pageId="13" pageNumber="14">Spintherobolus</emphasis>
</taxonomicName>
and, in part, used this feature to unite these genera in a phyletic group.
<bibRefCitation id="EFE24B552A3EFFE6F6B9FE8D4D25FE1B" author="Géry, J." bookContentInfo="T. F. H. Publications, Neptune City, New Jersey" pageId="13" pageNumber="14" pagination="672" refId="ref29398" refString="- 1977. Characoids of the World. T. F. H. Publications, Neptune City, New Jersey, 672 pp." title="Characoids of the World" type="book" year="1977">Géry (1977: 607)</bibRefCitation>
followed Eigenmann in this regard.
</paragraph>
<paragraph id="8BCC36A42A3EFFE6FA3FFE0A423FF937" blockId="13.[1339,2422,271,3456]" pageId="13" pageNumber="14">
Adipose fin loss has occurred several times within hypothesized independent clades of characiforms, for example the Characinae
<emphasis id="B907EAB62A3EFFE6F6B0FD83427DFDAE" box="[2304,2420,513,565]" italics="true" pageId="13" pageNumber="14">
(
<taxonomicName id="4C734D272A3EFFE6F6A1FD834CA4FDE8" authority="Weitzman &amp; Vari" authorityName="Weitzman &amp; Vari" authorityYear="1987" family="Characidae" genus="Priocharax" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FA8FFDBD4EC5FDE8" box="[1343,1484,575,627]" italics="true" pageId="13" pageNumber="14">Priocharax</emphasis>
Weitzman &amp; Vari
</taxonomicName>
</emphasis>
),
<taxonomicName id="4C734D272A3EFFE6F856FDBD427BFDE8" box="[2022,2418,575,627]" pageId="13" pageNumber="43" rank="tribe" tribe="Xenurobryconini">Xenurobryconini</taxonomicName>
(
<taxonomicName id="4C734D272A3EFFE6FAE0FDF94DE9FD34" authority="Roberts" authorityName="Roberts" authorityYear="1973" box="[1360,1760,635,687]" family="Characidae" genus="Iotabrycon" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6FAE0FDF94D2AFD34" box="[1360,1571,635,687]" italics="true" pageId="13" pageNumber="14">Iotabrycon</emphasis>
Roberts
</taxonomicName>
,
<taxonomicName id="4C734D272A3EFFE6F949FDF94D7FFD73" authority="Myers &amp; P. Miranda-Ribeiro" authorityName="Myers &amp; P. Miranda-Ribeiro" family="Characidae" genus="Xenurobrycon" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F949FDF9431BFD34" box="[1785,2066,635,687]" italics="true" pageId="13" pageNumber="14">Xenurobrycon</emphasis>
Myers &amp; P. Miranda-Ribeiro
</taxonomicName>
, and
<taxonomicName id="4C734D272A3EFFE6F944FD37439DFD73" authority="Fowler" authorityName="Fowler" authorityYear="1913" box="[1780,2196,693,744]" family="Characidae" genus="Tyttocharax" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F944FD374CEDFD73" box="[1780,2020,693,744]" italics="true" pageId="13" pageNumber="14">Tyttocharax</emphasis>
Fowler
</taxonomicName>
), the
<taxonomicName id="4C734D272A3EFFE6F6A1FD374D6DFCBC" family="Gasteropelecidae" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="family">Gasteropelecidae</taxonomicName>
<emphasis id="B907EAB62A3EFFE6F931FD734C7DFCBC" box="[1665,1908,753,807]" italics="true" pageId="13" pageNumber="14">
(
<taxonomicName id="4C734D272A3EFFE6F923FD734399FCBC" authority="Eigenmann" authorityName="Eigenmann" authorityYear="1909" box="[1683,2192,753,807]" family="Gasteropelecidae" genus="Carnegiella" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">Carnegiella Eigenmann</taxonomicName>
</emphasis>
), and the
<taxonomicName id="4C734D272A3EFFE6FA8FFCAD4D62FCFA" baseAuthorityName="Weitzman" baseAuthorityYear="1966" box="[1343,1643,815,865]" family="Lebiasinidae" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="family">Lebiasinidae</taxonomicName>
(some species of
<taxonomicName id="4C734D272A3EFFE6F7F8FCAD4EF6FC04" authority="Gunther" authorityName="Gunther" family="Lebiasinidae" genus="Nannostomus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F7F8FCAD427BFCFA" box="[2120,2418,815,865]" italics="true" pageId="13" pageNumber="14">Nannostomus</emphasis>
Gunther
</taxonomicName>
); see also discussion by
<bibRefCitation id="EFE24B552A3EFFE6F7E1FCEB4DD9FC42" author="Weitzman, S. H. &amp; H. Ortega" pageId="13" pageNumber="14" pagination="129 - 148" refId="ref31394" refString="Weitzman, S. H. &amp; H. Ortega. 1995. A new species of Tyttocharax (Teleostei: Characidae: Glandulocaudinae: Xenurobryconini) from the Rio Madre de Dios basin of Peru. Ichthyol. Explor. Freshwaters, 6: 129 - 148." type="journal article" year="1995">Weitzman &amp; Ortega (1995: 147)</bibRefCitation>
. In these genera the absence of an adipose fin is correlated with small to miniature size. The absence of an adipose fin in the American characiforms is not always associated with miniature size. For example the large species of
<emphasis id="B907EAB62A3EFFE6FA65FB574D92FA9C" box="[1493,1691,1237,1287]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="4C734D272A3EFFE6FA65FB574D9FFA9C" authorityName="Valenciennes" authorityYear="1847" box="[1493,1686,1237,1287]" family="Lebiasinidae" genus="Lebiasina" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">Lebiasina</taxonomicName>
,
</emphasis>
family
<taxonomicName id="4C734D272A3EFFE6F8F6FB574368FA9C" baseAuthorityName="Weitzman" baseAuthorityYear="1966" box="[1862,2145,1237,1287]" family="Lebiasinidae" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="family">Lebiasinidae</taxonomicName>
, and species of the
<taxonomicName id="4C734D272A3EFFE6FA58FA8D4C01FAD8" box="[1512,1800,1295,1347]" family="Erythrinidae" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="family">Erythrinidae</taxonomicName>
lack an adipose fin. The American characid genus
<taxonomicName id="4C734D272A3EFFE6F8CCFAC943CBFAE4" authority="Ellis" authorityName="Ellis" authorityYear="1911" box="[1916,2242,1355,1407]" family="Characidae" genus="Hasemania" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F8CCFAC9435FFAE4" box="[1916,2134,1355,1407]" italics="true" pageId="13" pageNumber="14">Hasemania</emphasis>
Ellis
</taxonomicName>
was for the most part defined as being like
<taxonomicName id="4C734D272A3EFFE6F7EEFA0A4D3CFA6C" authority="Durbin" authorityName="Durbin" authorityYear="1908" family="Characidae" genus="Hyphessobrycon" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F7EEFA0A4E76FA6C" italics="true" pageId="13" pageNumber="14">Hyphessobrycon</emphasis>
Durbin
</taxonomicName>
but without an adipose fin. As currently conceived
<taxonomicName id="4C734D272A3EFFE6F900F98143F3F9AF" authority="(Gery, 1977: 518)" baseAuthorityName="Gery" baseAuthorityPageNumber="518" baseAuthorityYear="1977" box="[1712,2298,1539,1588]" family="Characidae" genus="Hasemania" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3EFFE6F900F9814C83F9AF" box="[1712,1930,1539,1588]" italics="true" pageId="13" pageNumber="14">Hasemania</emphasis>
(
<bibRefCitation id="EFE24B552A3EFFE6F819F98143E0F9AF" author="Géry, J." bookContentInfo="T. F. H. Publications, Neptune City, New Jersey" box="[1961,2281,1539,1588]" pageId="13" pageNumber="14" pagination="672" refId="ref29398" refString="- 1977. Characoids of the World. T. F. H. Publications, Neptune City, New Jersey, 672 pp." title="Characoids of the World" type="book" year="1977">Géry, 1977:518</bibRefCitation>
)
</taxonomicName>
lacks miniature species. We agree with
<bibRefCitation id="EFE24B552A3EFFE6F7DAF9B94E9EF937" author="Géry, J." bookContentInfo="T. F. H. Publications, Neptune City, New Jersey" pageId="13" pageNumber="14" pagination="672" refId="ref29398" refString="- 1977. Characoids of the World. T. F. H. Publications, Neptune City, New Jersey, 672 pp." title="Characoids of the World" type="book" year="1977">Géry (1977: 518)</bibRefCitation>
that this genus is probably polyphyletic.
</paragraph>
<paragraph id="8BCC36A42A3EFFE6FA3FF936424BF406" blockId="13.[1339,2422,271,3456]" pageId="13" pageNumber="14">
(18, state 1) The infraorbital bones (
<figureCitation id="13482A212A3EFFE6F73DF9364213F972" box="[2189,2330,1716,1769]" captionStart="Fig" captionStartId="13.[191,252,1247,1291]" captionTargetBox="[187,1269,268,1188]" captionTargetPageId="13" captionText="Fig. 13. Spintherobolus ankoseion, infraorbital bones. a, USNM 297935, adult female, 24.6 mm SL; b, USNM 297935, adult male, 24.5 mm SL." figureDoi="http://doi.org/10.5281/zenodo.10882022" httpUri="https://zenodo.org/record/10882022/files/figure.png" pageId="13" pageNumber="14">Fig. 13</figureCitation>
a-b) are somewhat variable, but some are always reduced in size and number and others possibly fused. They are usually altered as follows. One large bony element is located in the usual charac ­ id position of infraorbital 2. In some specimens (
<figureCitation id="13482A212A3EFFE6FAFEF7A24ED2F7CF" box="[1358,1499,2080,2132]" captionStart="Fig" captionStartId="13.[191,252,1247,1291]" captionTargetBox="[187,1269,268,1188]" captionTargetPageId="13" captionText="Fig. 13. Spintherobolus ankoseion, infraorbital bones. a, USNM 297935, adult female, 24.6 mm SL; b, USNM 297935, adult male, 24.5 mm SL." figureDoi="http://doi.org/10.5281/zenodo.10882022" httpUri="https://zenodo.org/record/10882022/files/figure.png" pageId="13" pageNumber="14">Fig. 13</figureCitation>
a-b) this bone has an anterodorsal portion that is in the position of infraorbital 1of other characids and a posterior part that may represent infraorbital 2. Thus this bony element may consist of infraorbital 1 and 2 fused to each other. In some specimens this is the only infraorbital bone present. Another infraorbital, much reduced in size and apparently absent in
<taxonomicName id="4C734D272A3EFFE6F80AF64A4373F662" box="[1978,2170,2504,2553]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A3EFFE6F80AF64A4373F662" box="[1978,2170,2504,2553]" italics="true" pageId="13" pageNumber="14">S. broccae</emphasis>
</taxonomicName>
(18, state 2; see diagnosis under
<emphasis id="B907EAB62A3EFFE6F8A9F5834CFFF5AD" box="[1817,2038,2561,2614]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="4C734D272A3EFFE6F8A9F5834CEAF5AD" box="[1817,2019,2561,2614]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="broccae">S. broccae</taxonomicName>
),
</emphasis>
is placed in the position of infraorbital 3 (
<figureCitation id="13482A212A3EFFE6F839F5C2433FF5EA" box="[1929,2102,2624,2673]" captionStart="Fig" captionStartId="13.[191,252,1247,1291]" captionTargetBox="[187,1269,268,1188]" captionTargetPageId="13" captionText="Fig. 13. Spintherobolus ankoseion, infraorbital bones. a, USNM 297935, adult female, 24.6 mm SL; b, USNM 297935, adult male, 24.5 mm SL." figureDoi="http://doi.org/10.5281/zenodo.10882022" httpUri="https://zenodo.org/record/10882022/files/figure.png" pageId="13" pageNumber="14">Fig. 13a</figureCitation>
). Infraorbitals 4 through 6 are almost always absent, at least as ossifications, but in
<specimenCount id="9D75FD2D2A3EFFE6F94DF53A4323F572" box="[1789,2090,2744,2793]" count="1" pageId="13" pageNumber="14" type="generic">one specimen</specimenCount>
of
<taxonomicName id="4C734D272A3EFFE6F7CAF53A4267F572" box="[2170,2414,2744,2793]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A3EFFE6F7CAF53A4392F572" box="[2170,2203,2744,2793]" italics="true" pageId="13" pageNumber="14">S.</emphasis>
<emphasis id="B907EAB62A3EFFE6F71DF53A4267F572" box="[2221,2414,2744,2793]" italics="true" pageId="13" pageNumber="14">ankoseion</emphasis>
</taxonomicName>
a fourth infraorbital bone appears to be present (
<figureCitation id="13482A212A3EFFE6FAFDF4B24EF7F4FA" box="[1357,1534,2864,2913]" captionStart="Fig" captionStartId="13.[191,252,1247,1291]" captionTargetBox="[187,1269,268,1188]" captionTargetPageId="13" captionText="Fig. 13. Spintherobolus ankoseion, infraorbital bones. a, USNM 297935, adult female, 24.6 mm SL; b, USNM 297935, adult male, 24.5 mm SL." figureDoi="http://doi.org/10.5281/zenodo.10882022" httpUri="https://zenodo.org/record/10882022/files/figure.png" pageId="13" pageNumber="14">Fig. 13b</figureCitation>
). A well developed antorbital is present in the ventro-lateral wall of the nasal capsule.
</paragraph>
<paragraph id="8BCC36A42A3EFFE5FA3CF4274F16FC36" blockId="13.[1339,2422,271,3456]" lastBlockId="14.[95,1176,891,3472]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
Although reductions in the size and apparent loss of infraorbitals are observed in some Cheirodontines, for example
<taxonomicName id="4C734D272A3EFFE6F8F8F3A24EB7F316" authority="Eigenmann" authorityName="Eigenmann" authorityYear="1927" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="australe">
<emphasis id="B907EAB62A3EFFE6F8F8F3A243C3F3CF" box="[1864,2250,3104,3156]" italics="true" pageId="13" pageNumber="14">Cheirodon australe</emphasis>
Eigenmann
</taxonomicName>
, no other Cheirodontines have the infraorbitals with the anatomy described here for
<emphasis id="B907EAB62A3EFFE6FA8DF3564D7DF293" box="[1341,1652,3284,3336]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="4C734D272A3EFFE6FA8DF3564D79F293" authorityName="Eigenmann" authorityYear="1911" box="[1341,1648,3284,3336]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
A reduction of the posterior infraorbital bones also occurs in
<taxonomicName id="4C734D272A3EFFE6F86AF2934D82F21B" authority="(Humboldt)" baseAuthorityName="Humboldt" baseAuthorityYear="1821" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3EFFE6F86AF29343BCF2DF" box="[2010,2229,3345,3396]" italics="true" pageId="13" pageNumber="14">Grundulus</emphasis>
<emphasis id="B907EAB62A3EFFE6F775F2934267F2DF" box="[2245,2414,3345,3396]" italics="true" pageId="13" pageNumber="14">
<emphasis id="B907EAB62A3EFFE6FA8BF2CE4E62F21B" box="[1339,1387,3404,3456]" italics="true" pageId="13" pageNumber="14">bogotensis</emphasis>
</emphasis>
(Humboldt)
</taxonomicName>
but in that species infraorbital 2 and a large infraorbital 3 are well ossified.
</paragraph>
<caption id="DF0C662C2A3DFFE5FFD1FD094888FD70" ID-DOI="http://doi.org/10.5281/zenodo.10882025" ID-Zenodo-Dep="10882025" httpUri="https://zenodo.org/record/10882025/files/figure.png" pageId="14" pageNumber="15" startId="14.[97,163,651,695]" targetBox="[99,1178,287,598]" targetPageId="14" targetType="figure">
<paragraph id="8BCC36A42A3DFFE5FFD1FD094888FD70" blockId="14.[96,1177,651,747]" pageId="14" pageNumber="15">
Fig. 14.
<taxonomicName id="4C734D272A3DFFE5FEBCFD094876FD2C" authority=", USNM" authorityName="USNM" box="[268,895,651,695]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A3DFFE5FEBCFD0949E8FD2C" box="[268,737,651,695]" italics="true" pageId="14" pageNumber="15">Spintherobolus ankoseion,</emphasis>
USNM
</taxonomicName>
297935, adult male, 24.5 mm SL. Pelvic girdle and fin.
</paragraph>
</caption>
<paragraph id="8BCC36A42A3DFFE5FF1FFC354ACDF96D" blockId="14.[95,1176,891,3472]" pageId="14" pageNumber="15">
Infraorbital bone reduction and apparent loss are common in small and miniature characiforms of divergent clades; see
<bibRefCitation id="EFE24B552A3DFFE5FDC8FBAD4F91FBF8" author="Weitzman, S. H. &amp; W. L. Fink" box="[632,1176,1071,1123]" pageId="14" pageNumber="15" pagination="339 - 395" refId="ref31246" refString="Weitzman, S. H. &amp; W. L. Fink. 1983. Relationships of the neon tetras, a group of South American freshwater fishes (Teleostei, Characidae), with comments on the phylogeny of the New World characiforms. Bull. Mus. Comp. Zool., 150: 339 - 395." type="journal article" year="1983">Weitzman &amp; Fink (1983)</bibRefCitation>
and
<bibRefCitation id="EFE24B552A3DFFE5FF71FBEE49D6FB07" author="Weitzman, S. H. &amp; S. V. Fink" box="[193,735,1132,1180]" pageId="14" pageNumber="15" pagination="1 - 121" refId="ref31146" refString="Weitzman, S. H. &amp; S. V. Fink. 1985. Xenurobryconin phylogeny and putative pheromone pumps in glandulocaudine fishes (Teleostei: Characidae). Smithson. Contr. Zool., 421: 1 - 121, figs. 1 - 81." type="journal article" year="1985">Weitzman &amp; Fink (1985)</bibRefCitation>
for documentation. However,
<taxonomicName id="4C734D272A3DFFE5FEF7FB254952FB47" authorityName="Eigenmann" authorityYear="1911" box="[327,603,1191,1244]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3DFFE5FEF7FB254952FB47" box="[327,603,1191,1244]" italics="true" pageId="14" pageNumber="15">S. papilliferus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3DFFE5FD7CFB2548DEFB47" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[716,983,1191,1244]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3DFFE5FD7CFB2549FEFB47" box="[716,759,1191,1244]" italics="true" pageId="14" pageNumber="15">G.</emphasis>
<emphasis id="B907EAB62A3DFFE5FCB9FB2548DEFB47" box="[777,983,1191,1244]" italics="true" pageId="14" pageNumber="15">bogotensis</emphasis>
</taxonomicName>
are both comparatively large and were in part suggested as related by shared infraorbital loss by
<bibRefCitation id="EFE24B552A3DFFE5FBBCFA9C4AD8FA0B" author="Eigenmann, C. H." bookContentInfo="pls. 1 - 17" journalOrPublisher="Mem. Carnegie Mus." pageId="14" pageNumber="15" pagination="1 - 99" part="7" refId="ref29042" refString="- 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Mem. Carnegie Mus., 7: 1 - 99, pls. 1 - 17." title="The Cheirodontinae, a subfamily of minute characid fishes of South America" type="journal article" year="1915">Eigenmann (1915: 5-6</bibRefCitation>
, 14) and
<bibRefCitation id="EFE24B552A3DFFE5FD08FADC4F26FA0B" author="Géry, J." bookContentInfo="T. F. H. Publications, Neptune City, New Jersey" box="[696,1071,1374,1424]" pageId="14" pageNumber="15" pagination="672" refId="ref29398" refString="- 1977. Characoids of the World. T. F. H. Publications, Neptune City, New Jersey, 672 pp." title="Characoids of the World" type="book" year="1977">Géry (1977: 607)</bibRefCitation>
. Although the pattern of infraorbital loss in these species differs, this would not preclude consider ­ ing such loss a synapomorphy for the two species. However,
<taxonomicName id="4C734D272A3DFFE5FE0FF9C84992F91B" box="[447,667,1610,1664]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FE0FF9C84992F91B" box="[447,667,1610,1664]" italics="true" pageId="14" pageNumber="15">Grundulus</emphasis>
</taxonomicName>
lacks the synapomorphies uniting the species of
<taxonomicName id="4C734D272A3DFFE5FD68F9044F0AF927" authorityName="Eigenmann" authorityYear="1911" box="[728,1027,1670,1724]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FD68F9044F0AF927" box="[728,1027,1670,1724]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
to the
<taxonomicName id="4C734D272A3DFFE5FFD0F9464AC9F96D" box="[96,448,1732,1782]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="14" phylum="Chordata" rank="subFamily" subFamily="Cheirodontinae">Cheirodontinae</taxonomicName>
.
</paragraph>
<caption id="DF0C662C2A3DFFE5FB55FBEB4EB9FAF1" ID-DOI="http://doi.org/10.5281/zenodo.10882027" ID-Zenodo-Dep="10882027" httpUri="https://zenodo.org/record/10882027/files/figure.png" pageId="14" pageNumber="15" startId="14.[1253,1319,1129,1173]" targetBox="[1250,2328,286,1052]" targetPageId="14" targetType="figure">
<paragraph id="8BCC36A42A3DFFE5FB55FBEB4EB9FAF1" blockId="14.[1252,2324,1129,1386]" pageId="14" pageNumber="15">
Fig. 15.
<taxonomicName id="4C734D272A3DFFE5FA26FBEB4CFBFB0E" authority=", USNM" authorityName="USNM" box="[1430,2034,1129,1173]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A3DFFE5FA26FBEB4D97FB0E" box="[1430,1694,1129,1173]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
<emphasis id="B907EAB62A3DFFE5F908FBEB4C5BFB0E" box="[1720,1874,1129,1173]" italics="true" pageId="14" pageNumber="15">leptoura,</emphasis>
USNM
</taxonomicName>
346386, adult male, paratype, 21.8 mm SL. Anal fin, lateral view, left side, showing the distribution of pigment pattern on the fin and the derived nature of branched rays 2 through 5.
</paragraph>
</caption>
<paragraph id="8BCC36A42A3DFFE5FF1FF8824956F683" blockId="14.[95,1176,891,3472]" pageId="14" pageNumber="15">
(19) There are one unbranched and five to six branched pelvic-fin rays. The outgroup number of pelvic-fin raysin the
<taxonomicName id="4C734D272A3DFFE5FC93F8F84F12F82B" baseAuthorityName="Vari &amp; Siebert" baseAuthorityYear="1990" box="[803,1051,1914,1968]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Characidae</taxonomicName>
is i,7, and, with rare exceptions, this count is found in all outgroup cheirodontine species. The count in
<taxonomicName id="4C734D272A3DFFE5FFD1F7B34A80F7FF" authorityName="Eigenmann" authorityYear="1911" box="[97,393,2097,2148]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FFD1F7B34A80F7FF" box="[97,393,2097,2148]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
is i,5 or i,
<quantity id="4C8B9B412A3DFFE5FDC8F7B349C4F7FF" box="[632,717,2097,2148]" metricMagnitude="-1" metricUnit="m" metricValue="1.524" pageId="14" pageNumber="15" unit="in" value="6.0">6 in</quantity>
<taxonomicName id="4C734D272A3DFFE5FD54F7B348FEF7FF" authorityName="Eigenmann" authorityYear="1911" box="[740,1015,2097,2148]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3DFFE5FD54F7B348FEF7FF" box="[740,1015,2097,2148]" italics="true" pageId="14" pageNumber="15">S. papilliferus</emphasis>
</taxonomicName>
and is always i,
<quantity id="4C8B9B412A3DFFE5FE99F7EE4A8DF73B" box="[297,388,2156,2208]" metricMagnitude="-1" metricUnit="m" metricValue="1.27" pageId="14" pageNumber="15" unit="in" value="5.0">5 in</quantity>
the other three species. This is a derived reductive feature for the species of
<taxonomicName id="4C734D272A3DFFE5FFD0F7674A81F683" authorityName="Eigenmann" authorityYear="1911" box="[96,392,2277,2328]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FFD0F7674A81F683" box="[96,392,2277,2328]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
(
<figureCitation id="13482A212A3DFFE5FE05F7674940F683" box="[437,585,2277,2328]" captionStart="Fig" captionStartId="14.[97,163,651,695]" captionTargetBox="[99,1178,287,598]" captionTargetPageId="14" captionText="Fig. 14. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Pelvic girdle and fin." figureDoi="http://doi.org/10.5281/zenodo.10882025" httpUri="https://zenodo.org/record/10882025/files/figure.png" pageId="14" pageNumber="15">Fig. 14</figureCitation>
).
</paragraph>
<paragraph id="8BCC36A42A3DFFE5FF1DF6A3489AF435" blockId="14.[95,1176,891,3472]" pageId="14" pageNumber="15">
A low pelvic-fin ray count occurs in certain miniature or small sized characiforms, for example
<taxonomicName id="4C734D272A3DFFE5FF0CF6184AAAF656" authorityName="Eigenmann" authorityYear="1909" box="[188,419,2458,2509]" family="Gasteropelecidae" genus="Carnegiella" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FF0CF6184AAAF656" box="[188,419,2458,2509]" italics="true" pageId="14" pageNumber="15">Carnegiella</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFE24B552A3DFFE5FD80F6184F1CF656" author="Weitzman, S. H." box="[560,1045,2458,2509]" journalOrPublisher="Stanford Ichthyol. Bull." pageId="14" pageNumber="15" pagination="217 - 239" part="7" refId="ref31012" refString="- 1960. Further notes on the relationships and classification of the South American characid fishes of the subfamily Gasteropelecinae. Stanford Ichthyol. Bull., 7: 217 - 239." title="Further notes on the relationships and classification of the South American characid fishes of the subfamily Gasteropelecinae" type="journal article" year="1960">Weitzman, 1960: 217</bibRefCitation>
) and
<taxonomicName id="4C734D272A3DFFE5FFD0F65A48C2F59C" authority="Weitzman &amp; Vari (1987: 614)" authorityName="Weitzman &amp; Vari" authorityPageNumber="614" authorityYear="1987" box="[96,971,2520,2567]" family="Characidae" genus="Priocharax" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FFD0F65A4A3CF59C" box="[96,309,2520,2567]" italics="true" pageId="14" pageNumber="15">Priocharax</emphasis>
<bibRefCitation id="EFE24B552A3DFFE5FEF7F65A48C2F59C" author="Weitzman, S. H. &amp; R. P. Vari" box="[327,971,2520,2567]" pageId="14" pageNumber="15" pagination="640 - 652" refId="ref31443" refString="Weitzman, S. H. &amp; R. P. Vari. 1987. Two new species and a new genus of miniature characid fishes (Teleostei: Characiformes) from northern South Amer - ica. Proc. Biol. Soc. Wash., 100: 640 - 652." type="journal article" year="1987">Weitzman &amp; Vari (1987: 614)</bibRefCitation>
</taxonomicName>
, and one might therefore expect a low pelvic-fin ray count in a small species such as
<taxonomicName id="4C734D272A3DFFE5FD63F5CF4894F51A" box="[723,925,2637,2689]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A3DFFE5FD63F5CF49FDF51A" box="[723,756,2637,2689]" italics="true" pageId="14" pageNumber="15">S.</emphasis>
<emphasis id="B907EAB62A3DFFE5FCB7F5CF48ABF51A" box="[775,930,2637,2689]" italics="true" pageId="14" pageNumber="15">broccae.</emphasis>
</taxonomicName>
However,
<taxonomicName id="4C734D272A3DFFE5FFD0F50B4A79F525" authorityName="Eigenmann" authorityYear="1911" box="[96,368,2697,2750]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3DFFE5FFD0F50B4B76F525" box="[96,127,2697,2750]" italics="true" pageId="14" pageNumber="15">S.</emphasis>
<emphasis id="B907EAB62A3DFFE5FF20F50B4A79F525" box="[144,368,2697,2750]" italics="true" pageId="14" pageNumber="15">papilliferus</emphasis>
</taxonomicName>
although not especially small (to at least
<quantity id="4C8B9B412A3DFFE5FF7FF5444A80F561" box="[207,393,2758,2810]" metricMagnitude="-2" metricUnit="m" metricValue="6.08" pageId="14" pageNumber="15" unit="mm" value="60.8">60.8 mm</quantity>
<collectionCode id="ED62AE612A3DFFE5FE2BF5444ADFF561" box="[411,470,2758,2810]" pageId="14" pageNumber="15">SL</collectionCode>
) has a low pelvic-fin ray count.
<emphasis id="B907EAB62A3DFFE5FFD1F486492AF4AE" box="[97,547,2820,2869]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="4C734D272A3DFFE5FFD1F4864917F4AE" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[97,542,2820,2869]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bogotensis">Grundulus bogotensis</taxonomicName>
,
</emphasis>
also not especially small (to at least 67.0 mm
<collectionCode id="ED62AE612A3DFFE5FE43F4BE4921F4EB" box="[499,552,2876,2928]" pageId="14" pageNumber="15">SL</collectionCode>
) also has a somewhat reduced number of pelvic-fin rays, i,6.
</paragraph>
<paragraph id="8BCC36A42A3DFFE5FF1DF4374E8BF6C8" blockId="14.[95,1176,891,3472]" lastBlockId="14.[1246,2322,1493,3476]" pageId="14" pageNumber="15">
(20) The anal fin has iii-iv unbranched and 9- 16 branched rays (
<figureCitation id="13482A212A3DFFE5FDA9F47049AEF3B3" box="[537,679,3058,3112]" captionStart="Fig" captionStartId="8.[929,979,983,1017]" captionTargetBox="[929,1736,236,937]" captionTargetPageId="8" captionText="Fig. 9. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Anal-fin rays and associated pterygiophores, left side, anterior to right." figureDoi="http://doi.org/10.5281/zenodo.10882014" httpUri="https://zenodo.org/record/10882014/files/figure.png" pageId="14" pageNumber="15">Figs. 9</figureCitation>
,
<figureCitation id="13482A212A3DFFE5FD7FF4704808F3B3" box="[719,769,3058,3112]" captionStart="Fig" captionStartId="14.[1253,1319,1129,1173]" captionTargetBox="[1250,2328,286,1052]" captionTargetPageId="14" captionText="Fig. 15. Spintherobolus leptoura, USNM 346386, adult male, paratype, 21.8 mm SL. Anal fin, lateral view, left side, showing the distribution of pigment pattern on the fin and the derived nature of branched rays 2 through 5." figureDoi="http://doi.org/10.5281/zenodo.10882027" httpUri="https://zenodo.org/record/10882027/files/figure.png" pageId="14" pageNumber="15">15</figureCitation>
). The number of anal-fin rays in the
<taxonomicName id="4C734D272A3DFFE5FDA7F3AC4806F3FF" baseAuthorityName="Vari &amp; Siebert" baseAuthorityYear="1990" box="[535,783,3118,3172]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Characidae</taxonomicName>
is quite variable. The primitive state, however, can be hypothesized as having at least more than 15 branched rays as found in the apparently most primitive American characid species such as the species of
<taxonomicName id="4C734D272A3DFFE5FFD0F2DE49CAF20B" authority="Muller &amp; Troschell." authorityName="Muller &amp; Troschell." box="[96,707,3420,3472]" family="Bryconidae" genus="Brycon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FFD0F2DE4BE1F20B" box="[96,232,3420,3472]" italics="true" pageId="14" pageNumber="15">Brycon</emphasis>
Müller &amp; Troschell.
</taxonomicName>
Among the cheirodontine species the number of branched anal-fin rays varies from 16 to 24.
<taxonomicName id="4C734D272A3DFFE5F892F9904343F9DD" authorityName="Eigenmann" authorityYear="1911" box="[1826,2122,1554,1606]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5F892F9904343F9DD" box="[1826,2122,1554,1606]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
thus has a derived, reduced number and we suggest this as a synapomorphy uniting the species of the genus. A reduced number of anal-fin rays also occurs in the Chilean species of
<taxonomicName id="4C734D272A3DFFE5F82DF880436CF8A3" authorityName="Girard" authorityYear="1855" box="[1949,2149,1794,1848]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5F82DF880436CF8A3" box="[1949,2149,1794,1848]" italics="true" pageId="14" pageNumber="15">Cheirodon</emphasis>
</taxonomicName>
(C.
<taxonomicName id="4C734D272A3DFFE5F70EF8804D5CF8EE" authority="Eigenmann" authorityName="Eigenmann" authorityYear="1927" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="australe">
<emphasis id="B907EAB62A3DFFE5F70EF8804E35F8EE" italics="true" pageId="14" pageNumber="15">australe</emphasis>
Eigenmann
</taxonomicName>
,
<taxonomicName id="4C734D272A3DFFE5F9CAF8C0437CF8EE" authority="Eigenmann" authorityName="Eigenmann" authorityYear="1928" box="[1658,2165,1858,1909]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="galusdae">
<emphasis id="B907EAB62A3DFFE5F9CAF8C04DABF8EE" box="[1658,1698,1858,1909]" italics="true" pageId="14" pageNumber="15">C.</emphasis>
<emphasis id="B907EAB62A3DFFE5F902F8C04C68F8EE" box="[1714,1889,1858,1909]" italics="true" pageId="14" pageNumber="15">galusdae</emphasis>
Eigenmann
</taxonomicName>
(12-15 branched rays),
<taxonomicName id="4C734D272A3DFFE5F9D9F8FE4326F82A" authority="Girard" authorityName="Girard" authorityYear="1855" box="[1641,2095,1916,1969]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="pisciculus">
<emphasis id="B907EAB62A3DFFE5F9D9F8FE4D9BF82A" box="[1641,1682,1916,1969]" italics="true" pageId="14" pageNumber="15">C.</emphasis>
<emphasis id="B907EAB62A3DFFE5F900F8FE4C75F82A" box="[1712,1916,1916,1969]" italics="true" pageId="14" pageNumber="15">pisciculus</emphasis>
Girard
</taxonomicName>
,
<taxonomicName id="4C734D272A3DFFE5F7E6F8FE4218F82A" authorityName="Campos" authorityYear="1982" box="[2134,2321,1916,1969]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="kiliani">
<emphasis id="B907EAB62A3DFFE5F7E6F8FE4377F82A" box="[2134,2174,1916,1969]" italics="true" pageId="14" pageNumber="15">C.</emphasis>
<emphasis id="B907EAB62A3DFFE5F721F8FE4218F82A" box="[2193,2321,1916,1969]" italics="true" pageId="14" pageNumber="15">kiliani</emphasis>
</taxonomicName>
(9-13 )) but this is independently derived according to the hypothesis of cheirodontine relationships proposed herein.
<taxonomicName id="4C734D272A3DFFE5F941F7B343A3F7FD" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[1777,2218,2097,2150]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3DFFE5F941F7B34CC5F7FD" box="[1777,1996,2097,2150]" italics="true" pageId="14" pageNumber="15">Grundulus</emphasis>
<emphasis id="B907EAB62A3DFFE5F850F7B343A3F7FD" box="[2016,2218,2097,2150]" italics="true" pageId="14" pageNumber="15">bogotensis</emphasis>
</taxonomicName>
also has a reduced number of anal-fin rays, iv, 11 -12, but again evolved independently of
<emphasis id="B907EAB62A3DFFE5FB52F7644E2FF687" box="[1250,1318,2278,2332]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="4C734D272A3DFFE5F7A5F72E4E17F687" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5F7A5F72E4219F77B" box="[2069,2320,2220,2272]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
,
</emphasis>
based on its lack of synapomorphies of clades A toD.
</paragraph>
<paragraph id="8BCC36A42A3DFFE4FA81F6DC4FFCF44F" blockId="14.[1246,2322,1493,3476]" lastBlockId="15.[192,1272,2012,3451]" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15">
Branched anal-fin rays in
<taxonomicName id="4C734D272A3DFFE5F8CEF6DC43AFF60F" authorityName="Eigenmann" authorityYear="1911" box="[1918,2214,2398,2452]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5F8CEF6DC43AFF60F" box="[1918,2214,2398,2452]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
varies from 13- 16 (x = 14.1; state 1) in
<emphasis id="B907EAB62A3DFFE5F876F61F4398F656" box="[1990,2193,2461,2509]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="4C734D272A3DFFE5F876F61F4382F656" box="[1990,2187,2461,2509]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="broccae">S. broccae</taxonomicName>
,
</emphasis>
<quantity id="4C8B9B412A3DFFE5F712F61F4E03F597" metricMagnitude="-1" metricUnit="m" metricValue="3.175" metricValueMax="3.556" metricValueMin="2.794" pageId="14" pageNumber="15" unit="in" value="12.5" valueMax="14.0" valueMin="11.0">11 - 14 in</quantity>
<taxonomicName id="4C734D272A3DFFE5FA92F65A4D1CF597" box="[1314,1557,2520,2572]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A3DFFE5FA92F65A4D1CF597" box="[1314,1557,2520,2572]" italics="true" pageId="14" pageNumber="15">S. ankoseion</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3DFFE5F928F65A4C64F597" box="[1688,1901,2520,2572]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A3DFFE5F928F65A4C64F597" box="[1688,1901,2520,2572]" italics="true" pageId="14" pageNumber="15">S. leptoura</emphasis>
</taxonomicName>
(x = 12.6 for both species; state 2), and 9- 10 (x = 9.1; state 3) in
<taxonomicName id="4C734D272A3DFFE5FB52F5D24EFCF51F" authorityName="Eigenmann" authorityYear="1911" box="[1250,1525,2640,2692]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3DFFE5FB52F5D24E0BF51F" box="[1250,1282,2640,2692]" italics="true" pageId="14" pageNumber="15">S.</emphasis>
<emphasis id="B907EAB62A3DFFE5FAA4F5D24D08F51F" box="[1300,1537,2640,2692]" italics="true" pageId="14" pageNumber="15">papilliferus.</emphasis>
</taxonomicName>
We considered these three ranges and their means as different states in our analysis because all were significantly different and can thus be hypothesized as having a different genetic basis. All three states are derived according to outgroup comparison.
<taxonomicName id="4C734D272A3DFFE5F958F4FC43ADF42B" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[1768,2212,2942,2992]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3DFFE5F958F4FC4CCDF42B" box="[1768,1988,2942,2992]" italics="true" pageId="14" pageNumber="15">Grundulus</emphasis>
<emphasis id="B907EAB62A3DFFE5F86AF4FC43ADF42B" box="[2010,2212,2942,2992]" italics="true" pageId="14" pageNumber="15">bogotensis</emphasis>
</taxonomicName>
also has a small number of anal-fin rays (11-12) and has been suggested as closely related to the species of
<taxonomicName id="4C734D272A3DFFE5FA22F3B04DB4F3FD" authorityName="Eigenmann" authorityYear="1911" box="[1426,1725,3122,3174]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FA22F3B04DB4F3FD" box="[1426,1725,3122,3174]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFE24B552A3DFFE5F89AF3B043ACF3FD" author="Géry, J." bookContentInfo="T. F. H. Publications, Neptune City, New Jersey" box="[1834,2213,3122,3174]" pageId="14" pageNumber="15" pagination="672" refId="ref29398" refString="- 1977. Characoids of the World. T. F. H. Publications, Neptune City, New Jersey, 672 pp." title="Characoids of the World" type="book" year="1977">Géry (1977: 607)</bibRefCitation>
, but again, its anal fin is different from those of the species of
<taxonomicName id="4C734D272A3DFFE5FA66F3284DF7F345" authorityName="Eigenmann" authorityYear="1911" box="[1494,1790,3242,3294]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5FA66F3284DF7F345" box="[1494,1790,3242,3294]" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
in that the anal fin in
<taxonomicName id="4C734D272A3DFFE5FB51F3644EE0F282" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[1249,1513,3302,3353]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3DFFE5FB51F3644E03F282" box="[1249,1290,3302,3353]" italics="true" pageId="14" pageNumber="15">G.</emphasis>
<emphasis id="B907EAB62A3DFFE5FAADF3644EE0F282" box="[1309,1513,3302,3353]" italics="true" pageId="14" pageNumber="15">bogotensis</emphasis>
</taxonomicName>
lacks the derived features of the anal-fin rays in the males of the species of
<taxonomicName id="4C734D272A3DFFE5F7A4F2A04E10F20F" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3DFFE5F7A4F2A04E10F20F" italics="true" pageId="14" pageNumber="15">Spintherobolus</emphasis>
</taxonomicName>
described here. Also, the anal-fin pterygiophores of
<taxonomicName id="4C734D272A3CFFE4FE07F85E499AF788" box="[439,659,2012,2067]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4FE07F85E499AF788" box="[439,659,2012,2067]" italics="true" pageId="15" pageNumber="16">Grundulus</emphasis>
</taxonomicName>
appear derived in that they are all short, having no elongate anterior pterygiophores as found in the species of
<taxonomicName id="4C734D272A3CFFE4FC7AF7D64FFCF711" authorityName="Eigenmann" authorityYear="1911" box="[970,1269,2132,2186]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4FC7AF7D64FFCF711" box="[970,1269,2132,2186]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
and in most characids. Thus the reduced number of anal-fin rays in
<taxonomicName id="4C734D272A3CFFE4FDD2F74C4F29F699" baseAuthorityName="Humboldt" baseAuthorityYear="1821" box="[610,1056,2254,2306]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bogotensis">
<emphasis id="B907EAB62A3CFFE4FDD2F74C4F29F699" box="[610,1056,2254,2306]" italics="true" pageId="15" pageNumber="16">Grundulus bogotensis</emphasis>
</taxonomicName>
seems to be derived, but based on morphology not homologous to the condition found in
<taxonomicName id="4C734D272A3CFFE4FC7AF6C54FFDF6E0" box="[970,1268,2375,2427]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B907EAB62A3CFFE4FC7AF6C54FFDF6E0" box="[970,1268,2375,2427]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
species. The most parsimonious hypothesis in the global analysis indicates state 3 (9-10) anal fin rays, is an autapomorphy for
<taxonomicName id="4C734D272A3CFFE4FC8EF6794F47F5AB" authorityName="Eigenmann" authorityYear="1911" box="[830,1102,2555,2608]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3CFFE4FC8EF6794856F5AB" box="[830,863,2555,2608]" italics="true" pageId="15" pageNumber="16">S.</emphasis>
<emphasis id="B907EAB62A3CFFE4FCDCF6794F53F5AB" box="[876,1114,2555,2608]" italics="true" pageId="15" pageNumber="16">papilliferus.</emphasis>
</taxonomicName>
State 1 (13-16 branched rays) is a synapomorphy for
<taxonomicName id="4C734D272A3CFFE4FF73F5F64AE6F532" authorityName="Eigenmann" authorityYear="1911" box="[195,495,2676,2729]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4FF73F5F64AE6F532" box="[195,495,2676,2729]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
with an additional reduction of 13-14 branched rays (state 2), as synapomorphic for
<taxonomicName id="4C734D272A3CFFE4FEBDF56C4AF6F4BB" box="[269,511,2798,2848]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A3CFFE4FEBDF56C4AF6F4BB" box="[269,511,2798,2848]" italics="true" pageId="15" pageNumber="16">S. ankoseion</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A3CFFE4FDDFF56C484CF4BB" box="[623,837,2798,2848]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A3CFFE4FDDFF56C499BF4BB" box="[623,658,2798,2848]" italics="true" pageId="15" pageNumber="16">S.</emphasis>
<emphasis id="B907EAB62A3CFFE4FD12F56C484CF4BB" box="[674,837,2798,2848]" italics="true" pageId="15" pageNumber="16">leptoura</emphasis>
</taxonomicName>
is equally parsimonious with the hypothesis that state 2 is a synapomorphy for
<taxonomicName id="4C734D272A3CFFE4FE6FF4E04800F402" authorityName="Eigenmann" authorityYear="1911" box="[479,777,2914,2969]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4FE6FF4E04800F402" box="[479,777,2914,2969]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
together with an increase (state 1) as an autapomorphy for
<taxonomicName id="4C734D272A3CFFE4FB95F41D4FE3F44F" box="[1061,1258,2975,3028]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A3CFFE4FB95F41D4F32F44F" box="[1061,1083,2975,3028]" italics="true" pageId="15" pageNumber="16">S</emphasis>
.
<emphasis id="B907EAB62A3CFFE4FBE9F41D4FFCF44F" box="[1113,1269,2975,3028]" italics="true" pageId="15" pageNumber="16">broccae.</emphasis>
</taxonomicName>
</paragraph>
<caption id="DF0C662C2A3CFFE4FF74FBD24871F8AD" ID-DOI="http://doi.org/10.5281/zenodo.10882030" ID-Zenodo-Dep="10882030" httpUri="https://zenodo.org/record/10882030/files/figure.png" pageId="15" pageNumber="16" startId="15.[196,262,1104,1150]" targetBox="[196,1276,275,1028]" targetPageId="15" targetType="figure">
<paragraph id="8BCC36A42A3CFFE4FF74FBD24871F8AD" blockId="15.[194,1273,1104,1846]" pageId="15" pageNumber="16">
Fig. 16.
<taxonomicName id="4C734D272A3CFFE4FED9FBD2485BFBE5" authority=", MZUSP" authorityName="MZUSP" box="[361,850,1104,1150]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferum">
<emphasis id="B907EAB62A3CFFE4FED9FBD2485BFBE5" box="[361,850,1104,1150]" italics="true" pageId="15" pageNumber="16">Spintherobolus papilliferum,</emphasis>
</taxonomicName>
MZUSP 49408, adult female, 50.4 mm SL. Area of the anterior and posterior pseudotympanums of left side, anterior is to the right. The skin and scales were removed, exposing the muscles of the lateral body wall, pleural ribs of the fifth and sixth vertebrae (first and second pleural ribs), and the black, pigmented wall of the swimbladder. The larger black area to the left represents the pseudotympanum present in all cheirodontines. The somewhat obscured apical black area in the right upper part of the photograph is the area of the anterior pseudotympanum characteristic of the species of
<emphasis id="B907EAB62A3CFFE4FCA3F91E4F2EF950" box="[787,1063,1692,1739]" italics="true" pageId="15" pageNumber="16">
<taxonomicName id="4C734D272A3CFFE4FCA3F91E4F15F950" authorityName="Eigenmann" authorityYear="1911" box="[787,1052,1692,1739]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
However, in adult
<taxonomicName id="4C734D272A3CFFE4FED7F9504951F89B" authorityName="Eigenmann" authorityYear="1911" box="[359,600,1746,1792]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3CFFE4FED7F9504A8AF89B" box="[359,387,1746,1792]" italics="true" pageId="15" pageNumber="16">S.</emphasis>
<emphasis id="B907EAB62A3CFFE4FE21F9504951F89B" box="[401,600,1746,1792]" italics="true" pageId="15" pageNumber="16">papilliferus</emphasis>
</taxonomicName>
the anterior pseudotympanum is, in part, obscured by muscle tissue.
</paragraph>
</caption>
<caption id="DF0C662C2A3CFFE4FAF5FBD34305F90F" ID-DOI="http://doi.org/10.5281/zenodo.10882032" ID-Zenodo-Dep="10882032" httpUri="https://zenodo.org/record/10882032/files/figure.png" pageId="15" pageNumber="16" startId="15.[1349,1417,1105,1150]" targetBox="[1346,2431,278,1044]" targetPageId="15" targetType="figure">
<paragraph id="8BCC36A42A3CFFE4FAF5FBD34305F90F" blockId="15.[1348,2428,1105,1684]" pageId="15" pageNumber="16">
Fig. 17.
<taxonomicName id="4C734D272A3CFFE4FA5EFBD34CD1FBE5" authority=", MZUSP" authorityName="MZUSP" box="[1518,2008,1105,1150]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3CFFE4FA5EFBD34DF1FBE5" box="[1518,1784,1105,1150]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
<emphasis id="B907EAB62A3CFFE4F8B8FBD34CD1FBE5" box="[1800,2008,1105,1150]" italics="true" pageId="15" pageNumber="16">papilliferus,</emphasis>
</taxonomicName>
MZUSP 51022, juvenile, 23.9 mm SL. Anterior and posterior pseudotympanums of left side, anterior is to the right. The skin and scales were removed, exposing the muscles of the lateral body wall, pleural ribs of the fifth and sixth vertebrae (first and second pleural ribs), and the black, pigmented wall of the swimbladder. The larger black area to the left represents thepseudotympanum present in all cheirodontines, while the black area to the right is the area of the anterior pseudotympanum character ­ istic of the species of
<taxonomicName id="4C734D272A3CFFE4F94BF9E74308F90F" authorityName="Eigenmann" authorityYear="1911" box="[1787,2049,1637,1684]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BCC36A42A3CFFE4FEA2F45D4CF0F84E" blockId="15.[192,1272,2012,3451]" lastBlockId="15.[1344,2426,1831,3450]" pageId="15" pageNumber="16">
(21) An anterior pseudotympanum lies anterior to the first pleural rib. This pseudotympanum consists of a muscle hiatus lateral to the anterior portion of the swimbladder. The juveniles of
<taxonomicName id="4C734D272A3CFFE4FE34F352499FF298" box="[388,662,3280,3331]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3CFFE4FE34F352499FF298" box="[388,662,3280,3331]" italics="true" pageId="15" pageNumber="16">S. papilliferus</emphasis>
</taxonomicName>
and all specimens of the other three species display both pseudotympanums (
<figureCitation id="13482A212A3CFFE4FED0F2CB491FF2E0" box="[352,534,3401,3451]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="6.[70,119,841,876]" captionStartId-1="6.[938,986,1407,1441]" captionStartId-2="6.[938,986,1809,1843]" captionTargetBox-0="[68,882,219,802]" captionTargetBox-1="[935,1741,218,801]" captionTargetPageId-0="6" captionTargetPageId-1="6" captionTargetPageId-2="6" captionText-0="Fig. 4. Spintherobolus broccae, USNM 287324, adult male, 17.9 mm SL. Head and anterior body region, right side. Two pseudotympanums, black areas are located posterior to the dorsal portion of the opercular opening. The posterior pseudotympanum is homologous with the single pseudotympanum of other Cheirodontines while the anterior pseudotympanum is found only in Spintherobolus. The small dark dots arranged in rows arer the neuromast pattern over the face and opercular region. In the region ventral to the pectoral fin a few neuromasts can be seen partly bordering the scales. Such neuromasts are also present in the dorsal or back region. However, these are sometimes hard to distinguish from the somewhat larger dark chromatophores that often have somewhat indistinct borders. The rows of neuromasts are especially clear in the infraorbital region and on the dentary." captionText-1="Fig. 5. Spintherobolus ankoseion, paratype, MCP 19253, adult female, 25.2 mm SL. Anterior and posterior pseudotympanal region of left side, anterior is to the left. The skin and scales were removed, exposing the muscles of the lateral body wall and the pleural ribs of the fifth and sixth vertebrae (first and second pleural ribs). The black, pigmented swimbladder walls (the actual pseudotympanums) of both the anterior and posterior pseudotympanal regions have been removed, showing the black interior of the swimbladder." captionText-2="Fig. 6. Spintherobolus leptoura, paratype, MCP 19254, adult female, 25.4 mm SL. Anterior and posterior pseudotympanums of left side. The skin and scales were removed, exposing the muscles of the lateral body wall, pleural ribs of the fifth and sixth vertebrae (first and second pleural ribs), and the black, pigmented wall of the swimbladder. The larger black area to the right represents the pseudotympanum present in all Cheirodontines, while the black area to the left is the area of the pseudotympanum characteristic of the species of Spintherobolus." figureDoi-0="http://doi.org/10.5281/zenodo.10882001" figureDoi-1="http://doi.org/10.5281/zenodo.10882003" figureDoi-2="http://doi.org/10.5281/zenodo.10882006" httpUri-0="https://zenodo.org/record/10882001/files/figure.png" httpUri-1="https://zenodo.org/record/10882003/files/figure.png" httpUri-2="https://zenodo.org/record/10882006/files/figure.png" pageId="15" pageNumber="16">Figs. 4-6</figureCitation>
, 16). However, in adult
<taxonomicName id="4C734D272A3CFFE4FBE0F2CB4EB6F8C6" box="[1104,1471,1831,3451]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A3CFFE4FBE0F2CB4FFCF2E0" box="[1104,1269,3401,3451]" italics="true" pageId="15" pageNumber="16">
S.
<emphasis id="B907EAB62A3CFFE4FAF4F8A54EB6F8C6" box="[1348,1471,1831,1885]" italics="true" pageId="15" pageNumber="16">papilliferus</emphasis>
</emphasis>
</taxonomicName>
the anterior pseudotympanum is partially filled by muscular tissue and is relatively obscure but still present (
<figureCitation id="13482A212A3CFFE4F8E1F8234CD6F84E" box="[1873,2015,1953,2005]" captionStart="Fig" captionStartId="15.[1349,1417,1105,1150]" captionTargetBox="[1346,2431,278,1044]" captionTargetPageId="15" captionText="Fig. 17. Spintherobolus papilliferus, MZUSP 51022, juvenile, 23.9 mm SL. Anterior and posterior pseudotympanums of left side, anterior is to the right. The skin and scales were removed, exposing the muscles of the lateral body wall, pleural ribs of the fifth and sixth vertebrae (first and second pleural ribs), and the black, pigmented wall of the swimbladder. The larger black area to the left represents thepseudotympanum present in all cheirodontines, while the black area to the right is the area of the anterior pseudotympanum character­ istic of the species of Spintherobolus." figureDoi="http://doi.org/10.5281/zenodo.10882032" httpUri="https://zenodo.org/record/10882032/files/figure.png" pageId="15" pageNumber="16">Fig. 17</figureCitation>
).
</paragraph>
<paragraph id="8BCC36A42A3CFFE4FA24F85D427FF57E" blockId="15.[1344,2426,1831,3450]" pageId="15" pageNumber="16">
A posterior pseudotympanum is situated between the first and second pleural ribs in all cheirodontines including the species of
<taxonomicName id="4C734D272A3CFFE4F6BAF7D64D19F75F" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4F6BAF7D64D19F75F" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
but all other cheirodontines lack the anterior pseudotympanum. The presence of a muscle hiatus anterior to first pleural rib has been also observed in
<taxonomicName id="4C734D272A3CFFE4F886F6C54E8AF62E" authority="Myers" authorityName="Myers" authorityYear="1927" family="Characidae" genus="Atopomesus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="pachyodus">
<emphasis id="B907EAB62A3CFFE4F886F6C54202F6E1" box="[1846,2315,2375,2426]" italics="true" pageId="15" pageNumber="16">Atopomesus pachyodus</emphasis>
Myers
</taxonomicName>
,
<taxonomicName id="4C734D272A3CFFE4FA12F6004346F62E" authority="(Schultz)" baseAuthorityName="Schultz" baseAuthorityYear="1956" box="[1442,2127,2434,2485]" family="Characidae" genus="Paracheirodon" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="axelrodi">
<emphasis id="B907EAB62A3CFFE4FA12F6004C7FF62E" box="[1442,1910,2434,2485]" italics="true" pageId="15" pageNumber="16">Paracheirodon axelrodi</emphasis>
(Schultz)
</taxonomicName>
, and an unidentified species of
<taxonomicName id="4C734D272A3CFFE4F896F6424262F668" authority="Gery. However" authorityName="Gery. However" box="[1830,2411,2496,2547]" family="Characidae" genus="Axelrodia" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4F896F6424CE2F668" box="[1830,2027,2496,2547]" italics="true" pageId="15" pageNumber="16">Axelrodia</emphasis>
Géry. However
</taxonomicName>
, these species do not have all of the synapomorphies shared by species of the
<taxonomicName id="4C734D272A3CFFE4F7A4F5B54263F5F7" box="[2068,2410,2615,2668]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="14" phylum="Chordata" rank="subFamily" subFamily="Cheirodontinae">Cheirodontinae</taxonomicName>
. Thus these three species are excluded from that subfamily and relationships with
<emphasis id="B907EAB62A3CFFE4F7F1F52C427FF57E" box="[2113,2422,2734,2789]" italics="true" pageId="15" pageNumber="16">
<taxonomicName id="4C734D272A3CFFE4F7F1F52C4260F57E" authorityName="Eigenmann" authorityYear="1911" box="[2113,2409,2734,2789]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
</paragraph>
<paragraph id="8BCC36A42A3CFFE4FA25F5684C50F2E1" blockId="15.[1344,2426,1831,3450]" pageId="15" pageNumber="16">
(22) In adult males the anterior ventral procurrent caudal-fin rays, those that have their proximal ends inserted anterior to the haemal spine of the antepenultimate vertebrae, are fused to one another proximally (
<figureCitation id="13482A212A3CFFE4F83EF45E430AF38A" box="[1934,2051,3036,3089]" captionStart="Fig" captionStartId="8.[62,111,812,845]" captionTargetBox="[62,874,224,770]" captionTargetPageId="8" captionText="Fig. 8. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Osteology of ventral procurrent caudal-fin rays and associated bony structures, anterior to right." figureDoi="http://doi.org/10.5281/zenodo.10882012" httpUri="https://zenodo.org/record/10882012/files/figure.png" pageId="15" pageNumber="16">Fig. 8</figureCitation>
). Fusion begins with sexual maturation only in males. Among characiforms this character is unique to species of
<emphasis id="B907EAB62A3CFFE4FA32F3134DB0F35D" box="[1410,1721,3217,3270]" italics="true" pageId="15" pageNumber="16">
<taxonomicName id="4C734D272A3CFFE4FA32F3134DBBF35D" authorityName="Eigenmann" authorityYear="1911" box="[1410,1714,3217,3270]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
although one species of undescribed cheirodontine not closely related to
<taxonomicName id="4C734D272A3CFFE4FAF4F28B4D79F2A5" authorityName="Eigenmann" authorityYear="1911" box="[1348,1648,3337,3390]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A3CFFE4FAF4F28B4D79F2A5" box="[1348,1648,3337,3390]" italics="true" pageId="15" pageNumber="16">Spintherobolus</emphasis>
</taxonomicName>
also has some fusion among its ventral procurrent rays.
</paragraph>
<paragraph id="8BCC36A42A23FFFBFF00FEA74F0EFB3E" blockId="16.[95,1177,293,3480]" pageId="16" pageNumber="17">
(23) The anterior ventral procurrent caudal-fin rays of males have reduced proximal portions, not rising above the area of fusion between the rays, while the posterior dorsal portions of these rays are fused into a flat compressed plate that inserts between the hemal spine of the preural vertebrae (
<figureCitation id="13482A212A23FFFBFEFDFD0D4ACAFD58" box="[333,451,655,707]" captionStart="Fig" captionStartId="8.[62,111,812,845]" captionTargetBox="[62,874,224,770]" captionTargetPageId="8" captionText="Fig. 8. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Osteology of ventral procurrent caudal-fin rays and associated bony structures, anterior to right." figureDoi="http://doi.org/10.5281/zenodo.10882012" httpUri="https://zenodo.org/record/10882012/files/figure.png" pageId="16" pageNumber="17">Fig. 8</figureCitation>
). The ventral procurrent caudal-fin rays posteriorly associated with the hemal spine of pu3, pu2, and pul (parhypural) are separate. The proximal portions of the anterior, ventral procurrent caudal-fin rays in males of
<taxonomicName id="4C734D272A23FFFBFFD3FC394A82FC74" box="[99,395,955,1007]" pageId="16" pageNumber="17">
<emphasis id="B907EAB62A23FFFBFFD3FC394A82FC74" box="[99,395,955,1007]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
species are reduced or lost during sexual maturation. In young specimens and females of
<taxonomicName id="4C734D272A23FFFBFEABFBB1494AFBFC" authorityName="Eigenmann" authorityYear="1911" box="[283,579,1075,1127]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBFEABFBB1494AFBFC" box="[283,579,1075,1127]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
and other
<taxonomicName id="4C734D272A23FFFBFC98FBB14F15FBFC" baseAuthorityName="Vari &amp; Siebert" baseAuthorityYear="1990" box="[808,1052,1075,1127]" family="Characidae" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="family">Characidae</taxonomicName>
these portions are ossified and well-developed.
</paragraph>
<paragraph id="8BCC36A42A23FFFBFF1FFB2F4815F95B" blockId="16.[95,1177,293,3480]" pageId="16" pageNumber="17">
(24) The symphyseal dentary joint surfaces are smooth oval articular surfaces, lacking the intercalated and folded bony surfaces found in other characiforms. The articulation is supported by tough ligamentous tissue. A similar condition was found in the cheirodontine
<taxonomicName id="4C734D272A23FFFBFCF5FA5B490CF9D1" authority="(Boulenger)" baseAuthorityName="Boulenger" baseAuthorityYear="1909" family="Characidae" genus="Pseudocheirodon" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="arnoldi">
<emphasis id="B907EAB62A23FFFBFCF5FA5B4F9DF990" box="[837,1172,1497,1547]" italics="true" pageId="16" pageNumber="17">Pseudocheirodon</emphasis>
<emphasis id="B907EAB62A23FFFBFFEFF9974BE2F9D1" box="[95,235,1557,1610]" italics="true" pageId="16" pageNumber="17">arnoldi</emphasis>
(Boulenger)
</taxonomicName>
, a species that otherwise displays none of the synapomorphies of
<taxonomicName id="4C734D272A23FFFBFC27F9D24B91F95B" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBFC27F9D24B91F95B" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
and clade B Cheirodontines.
</paragraph>
<paragraph id="8BCC36A42A23FFFBFF00F9484F72F603" blockId="16.[95,1177,293,3480]" pageId="16" pageNumber="17">
(25) Lateral line reduced to 2-6 perforated scales. Lateral line reduction or loss has been used for defining cheirodontine and other characid genera since Eigenmann (e.g. 1915, 1917).
<bibRefCitation id="EFE24B552A23FFFBFFD0F83F483AF86B" author="Weitzman, S. H. &amp; W. L. Fink" box="[96,819,1981,2032]" pageId="16" pageNumber="17" pagination="339 - 395" refId="ref31246" refString="Weitzman, S. H. &amp; W. L. Fink. 1983. Relationships of the neon tetras, a group of South American freshwater fishes (Teleostei, Characidae), with comments on the phylogeny of the New World characiforms. Bull. Mus. Comp. Zool., 150: 339 - 395." type="journal article" year="1983">Weitzman &amp; Fink (1983: 391-394)</bibRefCitation>
discussed laterosensory reduction in small characids and noted that its loss is often correlated with small size and that the condition has evolved repeatedly in small characids. They also suggested that such reductive characters are phylogenetically significant at some level, but in practice can sometimes prove problematic for use as synapomorphies.
</paragraph>
<caption id="DF0C662C2A23FFFBFB52F894439FF8E7" ID-DOI="http://doi.org/10.5281/zenodo.10882036" ID-Zenodo-Dep="10882036" httpUri="https://zenodo.org/record/10882036/files/figure.png" pageId="16" pageNumber="17" startId="16.[1250,1317,1814,1861]" targetBox="[1262,2324,287,1763]" targetPageId="16" targetType="figure">
<paragraph id="8BCC36A42A23FFFBFB52F894439FF8E7" blockId="16.[1250,2322,1814,1916]" pageId="16" pageNumber="17">
Fig. 18.
<taxonomicName id="4C734D272A23FFFBFA3EF8944CF5F8DE" authority=", USNM" authorityName="USNM" box="[1422,2044,1814,1861]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A23FFFBFA3EF8944C69F8DE" box="[1422,1888,1814,1861]" italics="true" pageId="16" pageNumber="17">Spintherobolus ankoseion,</emphasis>
USNM
</taxonomicName>
297935, adult male, 24.5 mm SL. Osteology of pectoral girdle.
</paragraph>
</caption>
<paragraph id="8BCC36A42A23FFFBFF1FF6224F22F203" blockId="16.[95,1177,293,3480]" pageId="16" pageNumber="17">
In
<taxonomicName id="4C734D272A23FFFBFF5BF622491EF64F" authorityName="Eigenmann" authorityYear="1911" box="[235,535,2464,2516]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBFF5BF622491EF64F" box="[235,535,2464,2516]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
we found lateral-line reduction to be a synapomorphy for the species of the genus according to the most parsimonious distribution of the 35 characters used in our analysis of cheirodontine and other characids. Lateral-line counts range from
<quantity id="4C8B9B412A23FFFBFDADF54F4994F49B" box="[541,669,2765,2816]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" metricValueMax="1.524" metricValueMin="0.508" pageId="16" pageNumber="17" unit="in" value="4.0" valueMax="6.0" valueMin="2.0">2-6 in</quantity>
<taxonomicName id="4C734D272A23FFFBFD08F54F48EAF49B" box="[696,995,2765,2816]" pageId="16" pageNumber="17">
<emphasis id="B907EAB62A23FFFBFD08F54F48EAF49B" box="[696,995,2765,2816]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
species (mean
<quantity id="4C8B9B412A23FFFBFEB7F48A4A85F4A1" box="[263,396,2824,2874]" metricMagnitude="-2" metricUnit="m" metricValue="9.398" pageId="16" pageNumber="17" unit="in" value="3.7">3.7 in</quantity>
<emphasis id="B907EAB62A23FFFBFE1BF48A498AF4A1" box="[427,643,2824,2874]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFE1BF48A4977F4A1" box="[427,638,2824,2874]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="broccae">S. broccae</taxonomicName>
,
</emphasis>
4.0 in
<taxonomicName id="4C734D272A23FFFBFCF5F48A4F2EF4A1" box="[837,1063,2824,2874]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A23FFFBFCF5F48A4862F4A1" box="[837,875,2824,2874]" italics="true" pageId="16" pageNumber="17">S.</emphasis>
<emphasis id="B907EAB62A23FFFBFCCBF48A4F2EF4A1" box="[891,1063,2824,2874]" italics="true" pageId="16" pageNumber="17">leptoura</emphasis>
</taxonomicName>
and
<emphasis id="B907EAB62A23FFFBFFD4F4C64A69F4E3" box="[100,352,2884,2936]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFFD4F4C64A52F4E3" box="[100,347,2884,2936]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="ankoseion">S. ankoseion</taxonomicName>
,
</emphasis>
<quantity id="4C8B9B412A23FFFBFEC8F4C64AFDF4E3" box="[376,500,2884,2936]" metricMagnitude="-1" metricUnit="m" metricValue="1.1683999999999999" pageId="16" pageNumber="17" unit="in" value="4.6">4.6 in</quantity>
<emphasis id="B907EAB62A23FFFBFDBDF4C6484FF4E3" box="[525,838,2884,2936]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFDBDF4C6483DF4E3" box="[525,820,2884,2936]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="papilliferus">S. papilliferus</taxonomicName>
).
</emphasis>
In the nearest clade to
<emphasis id="B907EAB62A23FFFBFE81F4FC4961F42F" box="[305,616,2942,2996]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFE81F4FC496AF42F" authorityName="Eigenmann" authorityYear="1911" box="[305,611,2942,2996]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
,
</emphasis>
the
<taxonomicName id="4C734D272A23FFFBFD57F4FC4885F42F" baseAuthorityName="Lutken" baseAuthorityYear="1875" box="[743,908,2942,2996]" family="Characidae" genus="Odontostilbe" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="piaba">
<emphasis id="B907EAB62A23FFFBFD57F4FC4885F42F" box="[743,908,2942,2996]" italics="true" pageId="16" pageNumber="17">O. piaba</emphasis>
</taxonomicName>
group, the means in lateral-line range from 6.5-9.2. In the next closest clade, the species of
<emphasis id="B907EAB62A23FFFBFCD3F47A4F32F3B6" box="[867,1083,3064,3117]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFCD3F47A4F3FF3B6" authorityName="Girard" authorityYear="1855" box="[867,1078,3064,3117]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Cheirodon</taxonomicName>
,
</emphasis>
the means range from 6.7-8.8. No other Cheirodontines have the reduced number of pored scales in
<emphasis id="B907EAB62A23FFFBFFD7F32C498AF379" box="[103,643,3246,3298]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBFFD7F32C4A9BF379" authorityName="Eigenmann" authorityYear="1911" box="[103,402,3246,3298]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
<taxonomicName id="4C734D272A23FFFBFE17F32C498AF379" box="[423,643,3246,3298]" family="Characidae" genus="Grundulus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Grundulus</taxonomicName>
</emphasis>
also has a small number of perforated lateral-line scales (5-6), but this appears independently derived according to the most parsimonious hypothesis of our data.
</paragraph>
<paragraph id="8BCC36A42A23FFFBFA84F87A4D2FF597" blockId="16.[1250,2324,2040,3418]" pageId="16" pageNumber="17">
(26) The coracoid bone of the pectoral girdle of
<taxonomicName id="4C734D272A23FFFBFA91F7B64D43F7F3" authorityName="Eigenmann" authorityYear="1911" box="[1313,1610,2100,2152]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBFA91F7B64D43F7F3" box="[1313,1610,2100,2152]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
is reduced in length, and more or less discoid in shape (
<figureCitation id="13482A212A23FFFBF8F4F7F24CDBF73F" box="[1860,2002,2160,2212]" captionStart="Fig" captionStartId="16.[1250,1317,1814,1861]" captionTargetBox="[1262,2324,287,1763]" captionTargetPageId="16" captionText="Fig. 18. Spintherobolus ankoseion, USNM 297935, adult male, 24.5 mm SL. Osteology of pectoral girdle." figureDoi="http://doi.org/10.5281/zenodo.10882036" httpUri="https://zenodo.org/record/10882036/files/figure.png" pageId="16" pageNumber="17">Fig 18</figureCitation>
). In all other cheirodontines and characids examined by us the coracoid is either derived in other ways, for example in the gasteropelecids (
<bibRefCitation id="EFE24B552A23FFFBF81AF6A74E23F60D" author="Weitzman, S. H." pageId="16" pageNumber="17" pagination="213 - 263" refId="ref30974" refString="Weitzman, S. H. 1954. The osteology and the relation - ships of the South American characid fishes of the subfamily Gasteropelecinae. Stanford Ichthyol. Bull., 4: 213 - 263." type="journal article" year="1954">Weitzman, 1954: 225</bibRefCitation>
), or elongate and similar to that described by
<bibRefCitation id="EFE24B552A23FFFBFB52F6224DABF64A" author="Weitzman, S. H." box="[1250,1698,2464,2513]" journalOrPublisher="Stanford Ichthyol. Bull." pageId="16" pageNumber="17" pagination="1 - 77" part="8" refId="ref31045" refString="- 1962. The osteology of Brycon meeki, a generalized characoid fish, with an osteological definition of the family. Stanford Ichthyol. Bull., 8: 1 - 77." title="The osteology of Brycon meeki, a generalized characoid fish, with an osteological definition of the family" type="journal article" year="1962">Weitzman (1962: 41)</bibRefCitation>
for
<taxonomicName id="4C734D272A23FFFBF8B1F6224D2FF597" authority="Eigenmann &amp; Hildebrand." authorityName="Eigenmann &amp; Hildebrand." family="Bryconidae" genus="Brycon" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="meeki">
<emphasis id="B907EAB62A23FFFBF8B1F6224303F64A" box="[1793,2058,2464,2513]" italics="true" pageId="16" pageNumber="17">Brycon meeki</emphasis>
Eigenmann &amp; Hildebrand.
</taxonomicName>
</paragraph>
<paragraph id="8BCC36A42A23FFFBFA84F59A4CDDF2C1" blockId="16.[1250,2324,2040,3418]" pageId="16" pageNumber="17">
(27, state 1) The eyes of species of
<taxonomicName id="4C734D272A23FFFBF7A6F59A4E15F51D" authorityName="Eigenmann" authorityYear="1911" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBF7A6F59A4E15F51D" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
are small and derived compared to other Cheirodontines.
<taxonomicName id="4C734D272A23FFFBFA5EF5124CB1F55F" box="[1518,1976,2704,2756]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A23FFFBFA5EF5124C1FF55F" box="[1518,1814,2704,2756]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
<emphasis id="B907EAB62A23FFFBF899F5124CB1F55F" box="[1833,1976,2704,2756]" italics="true" pageId="16" pageNumber="17">broccae</emphasis>
</taxonomicName>
has a mean eye diameter of 28.7 %,
<taxonomicName id="4C734D272A23FFFBF915F54E4C73F565" box="[1701,1914,2764,2814]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A23FFFBF915F54E4DCCF565" box="[1701,1733,2764,2814]" italics="true" pageId="16" pageNumber="17">S.</emphasis>
<emphasis id="B907EAB62A23FFFBF968F54E4C73F565" box="[1752,1914,2764,2814]" italics="true" pageId="16" pageNumber="17">leptoura</emphasis>
</taxonomicName>
a mean of 27.2 %,
<taxonomicName id="4C734D272A23FFFBFB55F48A4ED0F4A2" box="[1253,1497,2824,2873]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A23FFFBFB55F48A4ED0F4A2" box="[1253,1497,2824,2873]" italics="true" pageId="16" pageNumber="17">S. ankoseion</emphasis>
</taxonomicName>
a mean of 27.6 %, and
<taxonomicName id="4C734D272A23FFFBF86CF48A43E5F4A2" box="[2012,2284,2824,2873]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A23FFFBF86CF48A43E5F4A2" box="[2012,2284,2824,2873]" italics="true" pageId="16" pageNumber="17">S. papilliferus</emphasis>
</taxonomicName>
a mean of 21.3 % compared to head length. Cheirodontine outgroups related to
<taxonomicName id="4C734D272A23FFFBF852F402421BF42F" authorityName="Eigenmann" authorityYear="1911" box="[2018,2322,2944,2996]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBF852F402421BF42F" box="[2018,2322,2944,2996]" italics="true" pageId="16" pageNumber="17">Spintherobolus</emphasis>
</taxonomicName>
have relatively large eyes, 33.9-40.3 % HL in clade C (
<taxonomicName id="4C734D272A23FFFBFA9AF47A4EFFF3B7" authorityName="Girard" authorityYear="1855" box="[1322,1526,3064,3116]" family="Characidae" genus="Cheirodon" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A23FFFBFA9AF47A4EFFF3B7" box="[1322,1526,3064,3116]" italics="true" pageId="16" pageNumber="17">Cheirodon</emphasis>
</taxonomicName>
) and 35.4-41.0 % of head length in clade E (
<taxonomicName id="4C734D272A23FFFBFA18F3B74D45F3F3" baseAuthorityName="Lutken" baseAuthorityYear="1875" box="[1448,1612,3125,3176]" family="Characidae" genus="Odontostilbe" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="40" phylum="Chordata" rank="species" species="piaba">
O.
<emphasis id="B907EAB62A23FFFBFA55F3B74D45F3F3" box="[1509,1612,3125,3176]" italics="true" pageId="16" pageNumber="17">piaba</emphasis>
</taxonomicName>
and related species) and many American characids have an eye size ranging from about 25 to 35 % HL (
<bibRefCitation id="EFE24B552A23FFFBF888F32C4E12F286" author="Weitzman, S. H. &amp; S. V. Fink &amp; A. Machado-Allison &amp; R. Royero" pageId="16" pageNumber="17" pagination="45 - 64" refId="ref31195" refString="Weitzman, S. H., S. V. Fink, A. Machado-Allison &amp; R. Royero. 1994. A new genus and species of Glandulocaudinae (Teleostei: Characidae) from southern Venezuela. Ichthyol. Explor. Freshwaters, 5: 45 - 64." type="journal article" year="1994">Weitzman et al., 1994: 55</bibRefCitation>
). Small eye size is apparently a reductive synapomorphy for
<emphasis id="B907EAB62A23FFFBF910F2A44CDDF2C1" box="[1696,2004,3366,3418]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="4C734D272A23FFFBF910F2A44CC1F2C1" authorityName="Eigenmann" authorityYear="1911" box="[1696,1992,3366,3418]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
</paragraph>
<caption id="DF0C662C2A22FFFAFF6FFD754819FCC7" ID-DOI="http://doi.org/10.5281/zenodo.10882038" ID-Zenodo-Dep="10882038" httpUri="https://zenodo.org/record/10882038/files/figure.png" pageId="17" pageNumber="18" startId="17.[223,289,759,807]" targetBox="[227,1277,309,717]" targetPageId="17" targetType="figure">
<paragraph id="8BCC36A42A22FFFAFF6FFD754819FCC7" blockId="17.[223,1300,759,860]" pageId="17" pageNumber="18">
Fig. 19. Alternate cladograms of the relationships of the species of
<taxonomicName id="4C734D272A22FFFAFE4DFCAF4805FCC7" authorityName="Eigenmann" authorityYear="1911" box="[509,780,813,860]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BCC36A42A22FFFAFE9DFB8348ACF983" blockId="17.[217,1297,1025,2584]" pageId="17" pageNumber="18">
(28) Relatively short pectoral-fin lengths are characteristic of males of
<taxonomicName id="4C734D272A22FFFAFCBFFBBD48DAFBEA" box="[783,979,1087,1137]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAFCBFFBBD482CFBEA" box="[783,805,1087,1137]" italics="true" pageId="17" pageNumber="18">S</emphasis>
.
<emphasis id="B907EAB62A22FFFAFCF3FBBD48DAFBEA" box="[835,979,1087,1137]" italics="true" pageId="17" pageNumber="18">broccae</emphasis>
</taxonomicName>
(15.2-18.7, x = 16.9 %
<collectionCode id="ED62AE612A22FFFAFECCFBFE4AB0FB34" box="[380,441,1148,1199]" pageId="17" pageNumber="18">SL</collectionCode>
),
<taxonomicName id="4C734D272A22FFFAFE55FBFE49B0FB34" box="[485,697,1148,1199]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAFE55FBFE4AF2FB34" box="[485,507,1148,1199]" italics="true" pageId="17" pageNumber="18">S</emphasis>
.
<emphasis id="B907EAB62A22FFFAFDA9FBFE49B0FB34" box="[537,697,1148,1199]" italics="true" pageId="17" pageNumber="18">leptoura</emphasis>
</taxonomicName>
(15.6- 17.9, x = 16.9 %
<collectionCode id="ED62AE612A22FFFAFB0DFBFE4FF2FB34" box="[1213,1275,1148,1199]" pageId="17" pageNumber="18">SL</collectionCode>
), and
<taxonomicName id="4C734D272A22FFFAFEF4FB35495CFB77" box="[324,597,1207,1260]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A22FFFAFEF4FB354A52FB77" box="[324,347,1207,1260]" italics="true" pageId="17" pageNumber="18">S</emphasis>
.
<emphasis id="B907EAB62A22FFFAFEC5FB35495CFB77" box="[373,597,1207,1260]" italics="true" pageId="17" pageNumber="18">papilliferus</emphasis>
</taxonomicName>
(15.7-16.6, x = 16.1 %
<collectionCode id="ED62AE612A22FFFAFBE7FB354F9FFB77" box="[1111,1174,1207,1260]" pageId="17" pageNumber="18">SL</collectionCode>
). The males of
<taxonomicName id="4C734D272A22FFFAFE1BFB714996FABC" box="[427,671,1267,1319]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A22FFFAFE1BFB714996FABC" box="[427,671,1267,1319]" italics="true" pageId="17" pageNumber="18">S. ankoseion</emphasis>
</taxonomicName>
have relatively long pectoral fins (18.2-20.8, x = 19.0 %
<collectionCode id="ED62AE612A22FFFAFC21FAB248CEFAF8" box="[913,967,1328,1379]" pageId="17" pageNumber="18">SL</collectionCode>
) and the outgroup clades C and E also have relatively long pectoral fins, 17.0-20.9, x = 18.5 % SL and 19.6- 21.7, x = 20.2 % SL respectively.
</paragraph>
<caption id="DF0C662C2A22FFFAFAEDFB374DD9F95F" ID-DOI="http://doi.org/10.5281/zenodo.10882042" ID-Zenodo-Dep="10882042" httpUri="https://zenodo.org/record/10882042/files/figure.png" pageId="17" pageNumber="18" startId="17.[1373,1441,1205,1252]" targetBox="[1374,2455,311,1138]" targetPageId="17" targetType="figure">
<paragraph id="8BCC36A42A22FFFAFAEDFB374DD9F95F" blockId="17.[1372,2450,1205,1732]" pageId="17" pageNumber="18">
Fig. 20. Tukey box plots of the number of branched anal-fin rays in species of
<emphasis id="B907EAB62A22FFFAF8CBFB6E4384FA82" box="[1915,2189,1260,1305]" italics="true" pageId="17" pageNumber="18">
<taxonomicName id="4C734D272A22FFFAF8CBFB6E438DFA82" authorityName="Eigenmann" authorityYear="1911" box="[1915,2180,1260,1305]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
Visual comparison of these box plots shows that the anal-fin ray counts of
<taxonomicName id="4C734D272A22FFFAF991FAD54C11FA1F" box="[1569,1816,1367,1412]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A22FFFAF991FAD54D36FA1F" box="[1569,1599,1367,1412]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAF9E0FAD54C11FA1F" box="[1616,1816,1367,1412]" italics="true" pageId="17" pageNumber="18">papilliferus</emphasis>
</taxonomicName>
are obviously different from all other species. Also, although there is overlap, the counts of
<taxonomicName id="4C734D272A22FFFAFA25FA434D41FA74" box="[1429,1608,1473,1519]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAFA25FA434EBDFA74" box="[1429,1460,1473,1519]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAFA79FA434D41FA74" box="[1481,1608,1473,1519]" italics="true" pageId="17" pageNumber="18">broccae</emphasis>
</taxonomicName>
are obviously significantly different from those of
<taxonomicName id="4C734D272A22FFFAF9A7FA774DD6F9B8" box="[1559,1759,1525,1571]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAF9A7FA774D3DF9B8" box="[1559,1588,1525,1571]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAF9FDFA774DD6F9B8" box="[1613,1759,1525,1571]" italics="true" pageId="17" pageNumber="18">leptoura</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A22FFFAF8E5FA77433CF9B8" box="[1877,2101,1525,1571]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A22FFFAF8E5FA774C7DF9B8" box="[1877,1908,1525,1571]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAF83AFA77433CF9B8" box="[1930,2101,1525,1571]" italics="true" pageId="17" pageNumber="18">ankoseion</emphasis>
</taxonomicName>
. However, there were no significant differences between the counts of
<taxonomicName id="4C734D272A22FFFAF6C1F9A94EE4F914" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAF6C1F9A94EE4F914" italics="true" pageId="17" pageNumber="18">S. leptoura</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A22FFFAF9E5F9E24C26F914" box="[1621,1839,1632,1679]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A22FFFAF9E5F9E24C26F914" box="[1621,1839,1632,1679]" italics="true" pageId="17" pageNumber="18">S. ankoseion</emphasis>
</taxonomicName>
when subject to a Mann-Whitney rank-sum test.
</paragraph>
</caption>
<paragraph id="8BCC36A42A22FFFAFE9BF9A24A5CF583" blockId="17.[217,1297,1025,2584]" pageId="17" pageNumber="18">
The distribution of this feature is ambiguous in three equally most parsimonious cladograms based on other characters (Figs. 3,19a-b). First, in all three cladograms this feature may be considered a synapomorphy for hypothesizing the monophyly of
<taxonomicName id="4C734D272A22FFFAFD94F8CD4844F81F" authorityName="Eigenmann" authorityYear="1911" box="[548,845,1871,1924]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A22FFFAFD94F8CD4844F81F" box="[548,845,1871,1924]" italics="true" pageId="17" pageNumber="18">Spintherobolus</emphasis>
</taxonomicName>
, but with a reversal to a longer pectoral fin in
<taxonomicName id="4C734D272A22FFFAFC9BF80F4F2CF85B" box="[811,1061,1933,1984]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A22FFFAFC9BF80F4845F85B" box="[811,844,1933,1984]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAFCEFF80F4F25F85B" box="[863,1068,1933,1984]" italics="true" pageId="17" pageNumber="18">ankoseion,</emphasis>
</taxonomicName>
requiring two steps in each. However, in the cladogram having
<taxonomicName id="4C734D272A22FFFAFE38F7864942F7A2" box="[392,587,2052,2105]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAFE38F7864942F7A2" box="[392,587,2052,2105]" italics="true" pageId="17" pageNumber="18">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A22FFFAFD79F78648A9F7A2" box="[713,928,2052,2105]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAFD79F78648A9F7A2" box="[713,928,2052,2105]" italics="true" pageId="17" pageNumber="18">S. leptoura</emphasis>
</taxonomicName>
united as sister species (
<figureCitation id="13482A212A22FFFAFE26F7C24935F7EF" box="[406,572,2112,2164]" captionStart="Fig" captionStartId="17.[223,289,759,807]" captionTargetBox="[227,1277,309,717]" captionTargetPageId="17" captionText="Fig. 19. Alternate cladograms of the relationships of the species of Spintherobolus." figureDoi="http://doi.org/10.5281/zenodo.10882038" httpUri="https://zenodo.org/record/10882038/files/figure.png" pageId="17" pageNumber="18">Fig. 19a</figureCitation>
) another, two step, equally par ­ simonious hypothesis is supported by the data. This hypothesis indicates that a short pectoral fin was independently acquired in
<taxonomicName id="4C734D272A22FFFAFC28F7774FA2F6B3" box="[920,1195,2293,2344]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A22FFFAFC28F77748B1F6B3" box="[920,952,2293,2344]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAFC79F7774FA2F6B3" box="[969,1195,2293,2344]" italics="true" pageId="17" pageNumber="18">papilliferus</emphasis>
</taxonomicName>
and in the clade uniting
<taxonomicName id="4C734D272A22FFFAFD7DF6B348ADF6FE" box="[717,932,2353,2405]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAFD7DF6B348ADF6FE" box="[717,932,2353,2405]" italics="true" pageId="17" pageNumber="18">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A22FFFAFB81F6B34E04F6FE" box="[1073,1293,2353,2405]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAFB81F6B34F5AF6FE" box="[1073,1107,2353,2405]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAFBD8F6B34E04F6FE" box="[1128,1293,2353,2405]" italics="true" pageId="17" pageNumber="18">leptoura</emphasis>
</taxonomicName>
(
<figureCitation id="13482A212A22FFFAFF5BF6EE4A9FF639" box="[235,406,2412,2466]" captionStart="Fig" captionStartId="17.[223,289,759,807]" captionTargetBox="[227,1277,309,717]" captionTargetPageId="17" captionText="Fig. 19. Alternate cladograms of the relationships of the species of Spintherobolus." figureDoi="http://doi.org/10.5281/zenodo.10882038" httpUri="https://zenodo.org/record/10882038/files/figure.png" pageId="17" pageNumber="18">Fig. 19a</figureCitation>
). In this hypothesis the character of short pectoral fin does not support
<taxonomicName id="4C734D272A22FFFAFCC9F62E4FADF645" authorityName="Eigenmann" authorityYear="1911" box="[889,1188,2476,2526]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A22FFFAFCC9F62E4FADF645" box="[889,1188,2476,2526]" italics="true" pageId="17" pageNumber="18">Spintherobolus</emphasis>
</taxonomicName>
as a clade.
</paragraph>
</subSubSection>
<subSubSection id="C369652F2A22FFFAFF69F5DC490BF41B" pageId="17" pageNumber="18" type="etymology">
<paragraph id="8BCC36A42A22FFFAFF69F5DC490BF41B" blockId="17.[217,1293,2654,2944]" pageId="17" pageNumber="18">
Etymology.
<taxonomicName id="4C734D272A22FFFAFE45F5DC4816F509" authorityName="Eigenmann" authorityYear="1911" box="[501,799,2654,2706]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A22FFFAFE45F5DC4816F509" box="[501,799,2654,2706]" italics="true" pageId="17" pageNumber="18">Spintherobolus</emphasis>
</taxonomicName>
comes from the Greek spinther meaning spark and obelos, to spit.
<bibRefCitation id="EFE24B552A22FFFAFB63F5184970F491" author="Eigenmann, C. H." bookContentInfo="pls. 1 - 17" journalOrPublisher="Mem. Carnegie Mus." pageId="17" pageNumber="18" pagination="1 - 99" part="7" refId="ref29042" refString="- 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Mem. Carnegie Mus., 7: 1 - 99, pls. 1 - 17." title="The Cheirodontinae, a subfamily of minute characid fishes of South America" type="journal article" year="1915">Eigenmann (1915:19)</bibRefCitation>
stated “emitting small sparks, referring to the appearance of yellow neuromasts on the head”.
</paragraph>
</subSubSection>
<caption id="DF0C662C2A22FFFAFAEBF51E4C2EF337" ID-DOI="http://doi.org/10.5281/zenodo.10882045" ID-Zenodo-Dep="10882045" httpUri="https://zenodo.org/record/10882045/files/figure.png" pageId="17" pageNumber="18" startId="17.[1371,1437,2716,2764]" targetBox="[1371,2413,1844,2665]" targetPageId="17" targetType="figure">
<paragraph id="8BCC36A42A22FFFAFAEBF51E4C2EF337" blockId="17.[1369,2448,2716,3244]" pageId="17" pageNumber="18">
Fig. 21. Tukey box plots of the number of vertebrae in species of
<emphasis id="B907EAB62A22FFFAF987F5534C42F49B" box="[1591,1867,2769,2816]" italics="true" pageId="17" pageNumber="18">
<taxonomicName id="4C734D272A22FFFAF987F5534C48F49B" authorityName="Eigenmann" authorityYear="1911" box="[1591,1857,2769,2816]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
Visual comparison of these box plots shows that the vertebral counts of
<taxonomicName id="4C734D272A22FFFAF705F4844E89F4F1" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A22FFFAF705F4844E89F4F1" italics="true" pageId="17" pageNumber="18">S. papilliferus</emphasis>
</taxonomicName>
are obviously different from all other species. Those of
<taxonomicName id="4C734D272A22FFFAFA3FF4F04D49F43A" box="[1423,1600,2930,2977]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAFA3FF4F04D49F43A" box="[1423,1600,2930,2977]" italics="true" pageId="17" pageNumber="18">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A22FFFAF91BF4F04C65F43A" box="[1707,1900,2930,2977]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAF91BF4F04DC1F43A" box="[1707,1736,2930,2977]" italics="true" pageId="17" pageNumber="18">S.</emphasis>
<emphasis id="B907EAB62A22FFFAF96CF4F04C65F43A" box="[1756,1900,2930,2977]" italics="true" pageId="17" pageNumber="18">leptoura</emphasis>
</taxonomicName>
are not significantly different and those of
<taxonomicName id="4C734D272A22FFFAF960F4244CBDF44E" box="[1744,1972,2982,3029]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A22FFFAF960F4244CBDF44E" box="[1744,1972,2982,3029]" italics="true" pageId="17" pageNumber="18">S. ankoseion</emphasis>
</taxonomicName>
, although overlapping with those of
<taxonomicName id="4C734D272A22FFFAF9DDF45E4C1CF397" box="[1645,1813,3036,3084]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A22FFFAF9DDF45E4D8CF397" box="[1645,1669,3036,3084]" italics="true" pageId="17" pageNumber="18">S</emphasis>
.
<emphasis id="B907EAB62A22FFFAF92CF45E4C1CF397" box="[1692,1813,3036,3084]" italics="true" pageId="17" pageNumber="18">broccae</emphasis>
</taxonomicName>
and 5.
<taxonomicName id="4C734D272A22FFFAF804F45E434CF397" box="[1972,2117,3036,3084]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A22FFFAF804F45E434CF397" box="[1972,2117,3036,3084]" italics="true" pageId="17" pageNumber="18">leptoura</emphasis>
</taxonomicName>
are significantly different from these two species when subject to Mann-Whitney rank-sum tests (T = 398, P &lt;0.001 and T = 246, P &lt;0.001 respectively).
</paragraph>
</caption>
<subSubSection id="C369652F2A22FFFAFEC8F3874FB0F23B" pageId="17" pageNumber="18" type="key">
<paragraph id="8BCC36A42A22FFFAFEC8F3874F60F3A3" blockId="17.[376,1129,3077,3128]" box="[376,1129,3077,3128]" pageId="17" pageNumber="18">
Key to species of
<taxonomicName id="4C734D272A22FFFAFCABF3874F60F3A3" authorityName="Eigenmann" authorityYear="1911" box="[795,1129,3077,3128]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="17" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="B907EAB62A22FFFAFCABF3874F60F3A3" box="[795,1129,3077,3128]" italics="true" pageId="17" pageNumber="18">Spintherobolus</emphasis>
</taxonomicName>
</paragraph>
<key id="FFE261952A22FFF8FF69F30248C1FB7A" lastPageId="19" lastPageNumber="20" pageId="17" pageNumber="18">
<keyStep id="30872E012A22FFFAFF69F3024284F239" pageId="17" pageNumber="18">
<keyLead id="308295912A22FFFAFF69F3024E00F23B" pageId="17" pageNumber="18">
<paragraph id="8BCC36A42A22FFFAFF69F3024FB0F23B" blockId="17.[217,1292,3196,3488]" pageId="17" pageNumber="18">
1 -Branchedanal-finrays 11-16 (
<figureCitation id="13482A212A22FFFAFC4FF3FE4F86F32B" box="[1023,1167,3196,3248]" captionStart="Fig" captionStartId="17.[1373,1441,1205,1252]" captionTargetBox="[1374,2455,311,1138]" captionTargetPageId="17" captionText="Fig. 20. Tukey box plots of the number of branched anal-fin rays in species of Spintherobolus. Visual comparison of these box plots shows that the anal-fin ray counts of S. papilliferus are obviously different from all other species. Also, although there is overlap, the counts of S. broccae are obviously significantly different from those of S. leptoura and S. ankoseion. However, there were no significant differences between the counts of S. leptoura and S. ankoseion when subject to a Mann-Whitney rank-sum test." figureDoi="http://doi.org/10.5281/zenodo.10882042" httpUri="https://zenodo.org/record/10882042/files/figure.png" pageId="17" pageNumber="18">Fig. 20</figureCitation>
); vertebrae32-34(
<figureCitation id="13482A212A22FFFAFD2CF338482FF376" box="[668,806,3258,3309]" captionStart="Fig" captionStartId="17.[1371,1437,2716,2764]" captionTargetBox="[1371,2413,1844,2665]" captionTargetPageId="17" captionText="Fig. 21. Tukey box plots of the number of vertebrae in species of Spintherobolus. Visual comparison of these box plots shows that the vertebral counts of S. papilliferus are obviously different from all other species. Those of S. broccae and S. leptoura are not significantly different and those of S. ankoseion, although overlapping with those of S. broccae and 5. leptoura are significantly different from these two species when subject to Mann- Whitney rank-sum tests (T = 398, P &lt;0.001 and T = 246, P &lt;0.001 respectively)." figureDoi="http://doi.org/10.5281/zenodo.10882045" httpUri="https://zenodo.org/record/10882045/files/figure.png" pageId="17" pageNumber="18">Fig. 21</figureCitation>
);scales in a longitudinalseries31-33(
<figureCitation id="13482A212A22FFFAFD6CF3764862F2B3" box="[732,875,3316,3368]" captionStart="Fig" captionStartId="18.[83,131,881,917]" captionTargetBox="[83,892,214,840]" captionTargetPageId="18" captionText="Fig. 22. Tukey box plots of the number of longitudinal series scales in species of Spintherobolus. Visual comparison of these box plots shows that these scale counts of S. papilliferus are obviously different from all the other species. The counts of. S. leptoura may be different from those of S. broccae and S. ankoseion, but the available counts for S. leptoura are too few (3) for useful statistical comparisons. The counts of S. broccae and S. ankoseion are not significantly different." figureDoi="http://doi.org/10.5281/zenodo.10882047" httpUri="https://zenodo.org/record/10882047/files/figure.png" pageId="17" pageNumber="18">Fig. 22</figureCitation>
);horizontal eye diameterlonger than snout length; coastal Atlanticdrainages,east of Serra Geral.
</paragraph>
<subSubSection id="C369652F2A22FFF8FB7EF2EC48C1FB7A" lastPageId="19" lastPageNumber="20" pageId="17" pageNumber="18" type="description">
<paragraph id="8BCC36A42A22FFF9FB7EF2EC4873F5BF" blockId="17.[217,1292,3196,3488]" lastBlockId="18.[177,891,2466,2596]" lastPageId="18" lastPageNumber="19" pageId="17" pageNumber="18">
.. 2
<keyLead id="308295912A22FFFAFA68F2B34284F239" pageId="17" pageNumber="18">- Branchedanal-finrays 9, rarely 10; vertebrae35-36;scales in a longitudinal series</keyLead>
<keyLead id="308295912A21FFF9FF01F6204873F5BF" pageId="18" pageNumber="19">
<keyStep id="30872E012A21FFF9FF01F6204873F5BF" pageId="18" pageNumber="19">
34-36; horizontal eye diameter shorter than snout length; headwaters of rio Tietê, upper rio Parana drainage.........
<taxonomicName id="4C734D272A21FFF9FD17F67F4873F5BF" box="[679,890,2557,2596]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A21FFF9FD17F67F49B0F5BF" box="[679,697,2557,2596]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FD0CF67F4873F5BF" box="[700,890,2557,2596]" italics="true" pageId="18" pageNumber="19">papilliferus</emphasis>
</taxonomicName>
</keyStep>
</keyLead>
</paragraph>
<caption id="DF0C662C2A21FFF9FFE3FCF349CAFB4E" ID-DOI="http://doi.org/10.5281/zenodo.10882047" ID-Zenodo-Dep="10882047" httpUri="https://zenodo.org/record/10882047/files/figure.png" pageId="18" pageNumber="19" startId="18.[83,131,881,917]" targetBox="[83,892,214,840]" targetPageId="18" targetType="figure">
<paragraph id="8BCC36A42A21FFF9FFE3FCF349CAFB4E" blockId="18.[82,892,881,1237]" pageId="18" pageNumber="19">
Fig. 22. Tukey box plots of the number of longitudinal series scales in species of
<emphasis id="B907EAB62A21FFF9FE56FC1949BFFC26" box="[486,694,923,957]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="4C734D272A21FFF9FE56FC1949B8FC26" authorityName="Eigenmann" authorityYear="1911" box="[486,689,923,957]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
Visual comparison of these box plots shows that these scale counts of
<taxonomicName id="4C734D272A21FFF9FFCFFC6B4A36FB96" box="[127,319,1001,1037]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A21FFF9FFCFFC6B4B87FB96" box="[127,142,1001,1037]" italics="true" pageId="18" pageNumber="19">S</emphasis>
.
<emphasis id="B907EAB62A21FFF9FF18FC6B4A36FB96" box="[168,319,1001,1037]" italics="true" pageId="18" pageNumber="19">papilliferus</emphasis>
</taxonomicName>
are obviously different from all the other species. The counts of.
<taxonomicName id="4C734D272A21FFF9FDB7FB904993FBAE" box="[519,666,1042,1077]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FDB7FB904917FBAE" box="[519,542,1042,1077]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FD9DFB904993FBAE" box="[557,666,1042,1077]" italics="true" pageId="18" pageNumber="19">leptoura</emphasis>
</taxonomicName>
may be different from those of
<taxonomicName id="4C734D272A21FFF9FEDCFBB84AFCFBC7" box="[364,501,1082,1116]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A21FFF9FEDCFBB84A72FBC7" box="[364,379,1082,1116]" italics="true" pageId="18" pageNumber="19">S</emphasis>
.
<emphasis id="B907EAB62A21FFF9FE24FBB84AFCFBC7" box="[404,501,1082,1116]" italics="true" pageId="18" pageNumber="19">broccae</emphasis>
</taxonomicName>
and
<emphasis id="B907EAB62A21FFF9FDFDFBB849F6FBC7" box="[589,767,1082,1116]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="4C734D272A21FFF9FDFDFBB849F3FBC7" box="[589,762,1082,1116]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">S. ankoseion</taxonomicName>
,
</emphasis>
but the available counts for
<taxonomicName id="4C734D272A21FFF9FECBFBE14903FB1E" box="[379,522,1123,1157]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FECBFBE14903FB1E" box="[379,522,1123,1157]" italics="true" pageId="18" pageNumber="19">S. leptoura</emphasis>
</taxonomicName>
are too few (3) for useful statistical comparisons. The counts of
<taxonomicName id="4C734D272A21FFF9FD19FB09483AFB36" box="[681,819,1163,1197]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A21FFF9FD19FB09483AFB36" box="[681,819,1163,1197]" italics="true" pageId="18" pageNumber="19">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A21FFF9FFE3FB314BFDFB4E" box="[83,244,1203,1237]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A21FFF9FFE3FB314B60FB4E" box="[83,105,1203,1237]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FFC4FB314BFDFB4E" box="[116,244,1203,1237]" italics="true" pageId="18" pageNumber="19">ankoseion</emphasis>
</taxonomicName>
are not significantly different.
</paragraph>
</caption>
<caption id="DF0C662C2A21FFF9FC05FCED4DDFFB30" ID-DOI="http://doi.org/10.5281/zenodo.10882050" ID-Zenodo-Dep="10882050" httpUri="https://zenodo.org/record/10882050/files/figure.png" pageId="18" pageNumber="19" startId="18.[949,999,879,915]" targetBox="[948,1754,211,843]" targetPageId="18" targetType="figure">
<paragraph id="8BCC36A42A21FFF9FC05FCED4DDFFB30" blockId="18.[948,1753,879,1195]" pageId="18" pageNumber="19">
Fig. 24. Graph of body depth as a function of standard length for
<taxonomicName id="4C734D272A21FFF9FBE9FC1A4E92FC20" box="[1113,1435,920,955]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FBE9FC1A4E16FC20" box="[1113,1311,920,955]" italics="true" pageId="18" pageNumber="19">Spintherobolus</emphasis>
<emphasis id="B907EAB62A21FFF9FA80FC1A4E92FC20" box="[1328,1435,920,955]" italics="true" pageId="18" pageNumber="19">leptoura</emphasis>
</taxonomicName>
(black symbols) and
<taxonomicName id="4C734D272A21FFF9FC05FC424F5FFC78" box="[949,1110,960,995]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A21FFF9FC05FC4248C5FC78" box="[949,972,960,995]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FC67FC424F5FFC78" box="[983,1110,960,995]" italics="true" pageId="18" pageNumber="19">ankoseion</emphasis>
</taxonomicName>
(empty symbols). Axes are logarithmic. Note that the 95 % confidence intervals in lengths of about 21 to 28 mm SL do not overlap with the means of the of the compared species indicating a significant difference in these size ranges. These growth pattern differences are used to separate these species in the key and text.
</paragraph>
</caption>
<caption id="DF0C662C2A21FFF9FC04F8134FA7F76D" ID-DOI="http://doi.org/10.5281/zenodo.10882052" ID-Zenodo-Dep="10882052" httpUri="https://zenodo.org/record/10882052/files/figure.png" pageId="18" pageNumber="19" startId="18.[948,998,1937,1973]" targetBox="[947,1750,1260,1892]" targetPageId="18" targetType="figure">
<paragraph id="8BCC36A42A21FFF9FC04F8134FA7F76D" blockId="18.[945,1753,1937,2294]" pageId="18" pageNumber="19">
Fig. 25. Graph of caudal peduncle depth as a function of standard length for
<taxonomicName id="4C734D272A21FFF9FA96F8384D67F847" box="[1318,1646,1978,2012]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FA96F8384EE7F847" box="[1318,1518,1978,2012]" italics="true" pageId="18" pageNumber="19">Spintherobolus</emphasis>
<emphasis id="B907EAB62A21FFF9F9B2F8384D67F847" box="[1538,1646,1978,2012]" italics="true" pageId="18" pageNumber="19">leptoura</emphasis>
</taxonomicName>
(black symbols) and
<taxonomicName id="4C734D272A21FFF9FB3EF8614E3CF79E" box="[1166,1333,2019,2053]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A21FFF9FB3EF8614FACF79E" box="[1166,1189,2019,2053]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FB04F8614E3CF79E" box="[1204,1333,2019,2053]" italics="true" pageId="18" pageNumber="19">ankoseion</emphasis>
</taxonomicName>
(empty symbols). Axes are logarithmic. Note that the 95 % confidence intervals in lengths of about 21 to 27 mm SL do not overlap with the means of the compared species indicating a significant difference in these size ranges. These growth pattern differences are used to separate these species in the key and text.
</paragraph>
</caption>
<caption id="DF0C662C2A21FFF9FFE3F82C4AE3F6A3" ID-DOI="http://doi.org/10.5281/zenodo.10882054" ID-Zenodo-Dep="10882054" httpUri="https://zenodo.org/record/10882054/files/figure.png" pageId="18" pageNumber="19" startId="18.[83,131,1966,2001]" targetBox="[83,892,1305,1932]" targetPageId="18" targetType="figure">
<paragraph id="8BCC36A42A21FFF9FFE3F82C4AE3F6A3" blockId="18.[82,892,1966,2360]" pageId="18" pageNumber="19">
Fig. 23. Tukey box plots of the number of procurrent caudal-fin rays in species of
<emphasis id="B907EAB62A21FFF9FDB1F85549D8F862" box="[513,721,2007,2041]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="4C734D272A21FFF9FDB1F85549C4F862" authorityName="Eigenmann" authorityYear="1911" box="[513,717,2007,2041]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">Spintherobolus</taxonomicName>
.
</emphasis>
Significant differences were not found between
<taxonomicName id="4C734D272A21FFF9FDCBF87D483FF7BA" box="[635,822,2047,2081]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A21FFF9FDCBF87D483FF7BA" box="[635,822,2047,2081]" italics="true" pageId="18" pageNumber="19">S. papilliferus</emphasis>
</taxonomicName>
and
<emphasis id="B907EAB62A21FFF9FFE5F7A54BE6F7D2" box="[85,239,2087,2121]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="4C734D272A21FFF9FFE5F7A54BE2F7D2" box="[85,235,2087,2121]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">S. leptoura</taxonomicName>
,
</emphasis>
between
<taxonomicName id="4C734D272A21FFF9FE3EF7A5492CF7D2" box="[398,549,2087,2121]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FE3EF7A5492CF7D2" box="[398,549,2087,2121]" italics="true" pageId="18" pageNumber="19">S. leptoura</emphasis>
</taxonomicName>
and
<emphasis id="B907EAB62A21FFF9FD30F7A5483AF7D2" box="[640,819,2087,2121]" italics="true" pageId="18" pageNumber="19">
<taxonomicName id="4C734D272A21FFF9FD30F7A54827F7D2" box="[640,814,2087,2121]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">S. ankoseion</taxonomicName>
,
</emphasis>
and between
<taxonomicName id="4C734D272A21FFF9FF6AF7CD4A9CF7EB" box="[218,405,2127,2160]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A21FFF9FF6AF7CD4A9CF7EB" box="[218,405,2127,2160]" italics="true" pageId="18" pageNumber="19">S. papilliferus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A21FFF9FE5DF7CD499FF7EB" box="[493,662,2127,2160]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A21FFF9FE5DF7CD490BF7EB" box="[493,514,2127,2160]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FDA0F7CD4993F7EB" box="[528,666,2127,2160]" italics="true" pageId="18" pageNumber="19">ankoseion.</emphasis>
</taxonomicName>
However, significant differences using Mann-Whitney rank sum tests were found between
<taxonomicName id="4C734D272A21FFF9FE26F71D495BF75B" box="[406,594,2207,2240]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="papilliferus">
<emphasis id="B907EAB62A21FFF9FE26F71D495BF75B" box="[406,594,2207,2240]" italics="true" pageId="18" pageNumber="19">S. papilliferus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A21FFF9FD17F71D4822F75B" box="[679,811,2207,2240]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A21FFF9FD17F71D49B7F75B" box="[679,702,2207,2240]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FD7EF71D4822F75B" box="[718,811,2207,2240]" italics="true" pageId="18" pageNumber="19">broccae</emphasis>
</taxonomicName>
(
<emphasis id="B907EAB62A21FFF9FCF5F71D4871F75B" box="[837,888,2207,2240]" italics="true" pageId="18" pageNumber="19">T =</emphasis>
984,
<emphasis id="B907EAB62A21FFF9FF27F7444BA1F773" box="[151,168,2246,2280]" italics="true" pageId="18" pageNumber="19">P</emphasis>
&lt;0.001), between
<taxonomicName id="4C734D272A21FFF9FE78F7444944F773" box="[456,589,2246,2280]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A21FFF9FE78F7444944F773" box="[456,589,2246,2280]" italics="true" pageId="18" pageNumber="19">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A21FFF9FD2FF7444838F773" box="[671,817,2246,2280]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FD2FF74449BCF773" box="[671,693,2246,2280]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FD73F7444838F773" box="[707,817,2246,2280]" italics="true" pageId="18" pageNumber="19">leptoura</emphasis>
</taxonomicName>
(
<emphasis id="B907EAB62A21FFF9FCFBF7444872F773" box="[843,891,2246,2280]" italics="true" pageId="18" pageNumber="19">T =</emphasis>
195,
<emphasis id="B907EAB62A21FFF9FF29F7724BC2F694" box="[153,203,2288,2319]" italics="true" pageId="18" pageNumber="19">P =</emphasis>
<emphasis id="B907EAB62A21FFF9FF68F7724BE0F694" box="[216,233,2288,2319]" italics="true" pageId="18" pageNumber="19">&lt;</emphasis>
0.001), and between
<taxonomicName id="4C734D272A21FFF9FD82F77249B0F694" box="[562,697,2288,2319]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="broccae">
<emphasis id="B907EAB62A21FFF9FD82F77249B0F694" box="[562,697,2288,2319]" italics="true" pageId="18" pageNumber="19">S. broccae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D272A21FFF9FCBCF7724B9BF6A3" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="ankoseion">
<emphasis id="B907EAB62A21FFF9FCBCF772482DF694" box="[780,804,2288,2319]" italics="true" pageId="18" pageNumber="19">S.</emphasis>
<emphasis id="B907EAB62A21FFF9FC82F7724872F694" box="[818,891,2288,2319]" italics="true" pageId="18" pageNumber="19">
<emphasis id="B907EAB62A21FFF9FFE3F69B4B9BF6A3" box="[83,146,2329,2360]" italics="true" pageId="18" pageNumber="19">ankoseion</emphasis>
</emphasis>
</taxonomicName>
(
<emphasis id="B907EAB62A21FFF9FF1EF69B4BC9F6A3" box="[174,192,2329,2360]" italics="true" pageId="18" pageNumber="19">T</emphasis>
= 604,
<emphasis id="B907EAB62A21FFF9FE9FF69B4A4BF6A3" box="[303,322,2329,2360]" italics="true" pageId="18" pageNumber="19">P</emphasis>
= &lt;0.001).
</paragraph>
</caption>
<paragraph id="8BCC36A42A21FFF8FC04F6C848C5FF6D" blockId="18.[948,1752,2374,2594]" lastBlockId="19.[259,973,217,435]" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
<keyLead id="308295912A21FFF9FC04F6C84DD1F582" pageId="18" pageNumber="19">
<keyStep id="30872E012A21FFF9FC04F6C84DD1F582" pageId="18" pageNumber="19">
2. Branched anal-fin rays 11 -14, usually 12- 13, ›`‹ = 12.6 for both species; ventral procurrent caudal-fin rays 12-16, ›`‹ = 14.0 for S. ankoseıoıı and 14.1 for
<taxonomicName id="4C734D272A21FFF9FAFEF6534EE5F66E" box="[1358,1516,2512,2549]" family="Characidae" genus="Spintherobolus" kingdom="Animalia" order="Characiformes" pageId="18" pageNumber="37" phylum="Chordata" rank="species" species="leptoura">
<emphasis id="B907EAB62A21FFF9FAFEF6534EE5F66E" box="[1358,1516,2512,2549]" pageId="18" pageNumber="19">S. leptoura</emphasis>
</taxonomicName>
(Fig. 23); third infraorbital bone present. 3
</keyStep>
</keyLead>
<keyLead id="308295912A20FFF8FEB4FF5B48C5FF6D" box="[260,972,217,257]" pageId="19" pageNumber="20">
<keyStep id="30872E012A20FFF8FEB4FF5B48C5FF6D" box="[260,972,217,257]" pageId="19" pageNumber="20">Branched anal-fin rays 13-16, usually 14-</keyStep>
</keyLead>
</paragraph>
<paragraph id="8BCC36A42A20FFF8FEB5FE8848C2FE37" blockId="19.[259,973,217,435]" pageId="19" pageNumber="20">
<keyLead id="308295912A20FFF8FEB5FE8848C2FE37" pageId="19" pageNumber="20">
<keyStep id="30872E012A20FFF8FEB5FE8848C2FE37" pageId="19" pageNumber="20">
15, i = 14.1; ventral procurrent caudal-fin rays 14-17, i = 15.4; third infraorbital absent; Rio de Janeiro region and area near Santos, São Paulo).
<taxonomicName id="4C734D272A20FFF8FC86FE1348C2FE37" box="[822,971,401,428]" pageId="19" pageNumber="20">
<emphasis id="B907EAB62A20FFF8FC86FE1348C2FE37" box="[822,971,401,428]" pageId="19" pageNumber="20">S. broccne</emphasis>
</taxonomicName>
</keyStep>
</keyLead>
</paragraph>
<keyLead id="308295912A20FFF8FF14FE6948C1FB7A" pageId="19" pageNumber="20">
<keyStep id="30872E012A20FFF8FF14FE6948C1FB7A" pageId="19" pageNumber="20">
<paragraph id="8BCC36A42A20FFF8FF14FE694874FCDA" blockId="19.[164,971,488,1249]" pageId="19" pageNumber="20">3. Body depth of specimens of both sexes between about 21.0 and 28.0 mm SL 32.4- 385:Á SL, i = 35.8 % (Fig. 24); male body depth regardless of body length 29.3-37.9 &quot;78 SL, i = 34.2 %; male caudal-peduncle depth 12.8- 15.6 % SL, i = 14.8 % (Fig. 25); freshwaters around Baia de Paranaguá, Paraná south to Barra do Sai, Santa Catarina.</paragraph>
<paragraph id="8BCC36A42A20FFF8FCBEFCD148C0FCF5" blockId="19.[164,971,488,1249]" box="[782,969,851,878]" pageId="19" pageNumber="20">
<taxonomicName id="4C734D272A20FFF8FCBEFCD148C0FCF5" box="[782,969,851,878]" pageId="19" pageNumber="20">
<emphasis id="B907EAB62A20FFF8FCBEFCD148C0FCF5" box="[782,969,851,878]" pageId="19" pageNumber="20">S. aııkoseíoıı</emphasis>
</taxonomicName>
</paragraph>
<paragraph id="8BCC36A42A20FFF8FEB1FC0348C1FB7A" blockId="19.[164,971,488,1249]" pageId="19" pageNumber="20">
Body depth of specimens of both sexes between about 21.0 and 28.0 mm SL 27.5- 33.7 3/11 SL, i = 31.0 3/1) (Fig. 24); male body depth regardless of body length 26.6-29.7 %, i = 28.3; male caudal peduncle depth 11.0- 13.1 % SL, i = 12.3
<date id="FFCD10642A20FFF8FDF1FBDF495FFBE7" box="[577,598,1117,1150]" pageId="19" pageNumber="20">9i</date>
, (Fig. 25); rio Quilombo, rio Ribeira de Iguape drainage, São Paulo.
<taxonomicName id="4C734D272A20FFF8FC94FB3E48C1FB7A" box="[804,968,1212,1249]" pageId="19" pageNumber="20">
<emphasis id="B907EAB62A20FFF8FC94FB3E48C1FB7A" box="[804,968,1212,1249]" pageId="19" pageNumber="20">S. Ieptoura</emphasis>
</taxonomicName>
</paragraph>
</keyStep>
</keyLead>
</subSubSection>
</keyLead>
</keyStep>
</key>
</subSubSection>
</treatment>
</document>
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