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<mods:title id="56619C1D0F07D244B0D46EBF103CC64A">Phylogenetic analysis and osteological comparison of the cave-dwelling spined loach, Bibarba parvoculus (Cypriniformes: Cobitidae), and its surface congener</mods:title>
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<paragraph id="14404F51383CFFC0FEB4F921FDF7F929" blockId="4.[314,592,1721,1746]" box="[314,592,1721,1746]" pageId="4" pageNumber="1063">
<heading id="4F08F83D383CFFC0FEB4F921FDF7F929" box="[314,592,1721,1746]" centered="true" fontSize="9" level="2" pageId="4" pageNumber="1063" reason="2">
OSTEOLOGY OF
<taxonomicName id="D3FF34D2383CFFC0FE63F921FDF7F929" ID-CoL="62GMJ" ID-ENA="1868606" authorityName="Chen &amp; Chen" authorityYear="2007" box="[493,592,1721,1744]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="genus">
<emphasis id="268B9343383CFFC0FE63F921FDF7F929" box="[493,592,1721,1744]" italics="true" pageId="4" pageNumber="1063">BIBARBA</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="5CE51CDA383CFFCCFF1FF979FE10FE29" lastPageId="8" lastPageNumber="1067" pageId="4" pageNumber="1063" type="description">
<paragraph id="14404F51383CFFC0FF1FF979FD4EF8AA" blockId="4.[145,761,1761,1875]" pageId="4" pageNumber="1063">
The cranial skeleton of
<taxonomicName id="D3FF34D2383CFFC0FE29F979FDA1F90F" authorityName="Chen &amp; Chen" authorityYear="2007" box="[423,518,1761,1782]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="genus">
<emphasis id="268B9343383CFFC0FE29F979FDA1F90F" box="[423,518,1761,1782]" italics="true" pageId="4" pageNumber="1063">Bibarba</emphasis>
</taxonomicName>
is typical of cobitids
<emphasis id="268B9343383CFFC0FF1FF899FF77F8EF" box="[145,208,1793,1814]" italics="true" pageId="4" pageNumber="1063">sensu</emphasis>
<bibRefCitation id="706E32A0383CFFC0FF59F898FED8F8EF" author="Sawada Y" box="[215,383,1792,1814]" pageId="4" pageNumber="1063" pagination="65 - 223" refId="ref9427" refString="Sawada Y. 1982. Phylogeny and zoogeography of the superfamily Cobitoidea (Cyprinoidei, Cypriniformes). Memoirs of the Faculty of Fisheries Hokkaido University 28: 65 - 223." type="journal article" year="1982">Sawada (1982)</bibRefCitation>
, with an evident metapterygoid quadrate fenestra in the suspensorium and an interorbital fenestra in the neurocranium (
<figureCitation id="8CC453D4383CFFC0FDF4F8A5FD18F8AB" box="[634,703,1853,1875]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Figs 4</figureCitation>
,
<figureCitation id="8CC453D4383CFFC0FD42F8A5FD7DF8AA" box="[716,730,1853,1875]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="4" pageNumber="1063">5</figureCitation>
).
</paragraph>
<paragraph id="14404F51383CFFC0FCCFFF5EFB67FBEE" blockId="4.[809,1426,197,1875]" pageId="4" pageNumber="1063">
In cave-dwelling
<taxonomicName id="D3FF34D2383CFFC0FB83FF5EFB0AFF23" box="[1037,1197,197,219]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="species" species="parvoculus">
<emphasis id="268B9343383CFFC0FB83FF5EFB0AFF23" box="[1037,1197,197,219]" italics="true" pageId="4" pageNumber="1063">B. parvoculus</emphasis>
</taxonomicName>
, the pterosphenoid forms the posterior rim of the interorbital fenestra, dorsoventrally separating the frontal and the parasphenoid, and posteriorly contacting only the sphenotic (
<figureCitation id="8CC453D4383CFFC0FC6DFED8FB9CFEAC" box="[995,1083,320,342]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Fig. 4C</figureCitation>
). The bifurcate pharyngeal process of the basioccipital is short and tapers to an end at a vertical approximately anterior to centrum 2 + 3 (
<figureCitation id="8CC453D4383CFFC0FCF6FE04FC4BFE48" box="[888,1004,412,434]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Fig. 4C, F</figureCitation>
). The entopterygoid is a slim bone with a blade-like posterior part and, anteriorly, articulates with the palatine and, posteriorly, with the metapterygoid (
<figureCitation id="8CC453D4383CFFC0FC53FE60FB88FDF4" box="[989,1071,504,526]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Fig. 4D</figureCitation>
). The posteroventral process of ceratobranchial 5 (pharyngeal bone) extends ventrally, forming a right angle with the ventral limb (
<figureCitation id="8CC453D4383CFFC0FABEFDADFA25FDB3" box="[1328,1410,565,587]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Fig. 4E</figureCitation>
). A single fontanelle enclosed by paired frontals, paired parietals and the supraoccipital is on the roof of the skull (
<figureCitation id="8CC453D4383CFFC0FCF7FD09FC76FD5E" box="[889,977,657,679]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6C</figureCitation>
). The anterior part of the frontal is narrow at the orbital region and about 36% as wide as the posterior part (
<figureCitation id="8CC453D4383CFFC0FBADFD56FB37FD1D" box="[1059,1168,718,740]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6C, F</figureCitation>
). The suborbital spine is relatively stout. A weakly ossified supraorbital with a grey value similar to that of the sclera ring in the 3D rendering (
<figureCitation id="8CC453D4383CFFC0FC5AFCB2FB8FFCB9" box="[980,1064,810,832]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6C</figureCitation>
) is at a vertical approximately anterior to the laterocaudal process of the suborbital spine. In the caudal skeleton, there are five hypurals, with hypurals 4 and 5 fused (in all specimens except for
<specimenCount id="02F984D8383CFFC0FCA7FC3DFC02FC42" box="[809,933,933,955]" pageId="4" pageNumber="1063" type="female">one female</specimenCount>
with hypurals 4 and 5 separate) (
<figureCitation id="8CC453D4383CFFC0FAA0FC3DFA24FC43" box="[1326,1411,933,955]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="4" pageNumber="1063">Fig. 4H</figureCitation>
). The vertebral count (including the first four centra forming the Weberian apparatus and the last pleural centrum) is 45 (
<emphasis id="268B9343383CFFC0FC52FB9AFC57FBEE" box="[988,1008,1026,1047]" italics="true" pageId="4" pageNumber="1063">N</emphasis>
= 2) or 46 (
<emphasis id="268B9343383CFFC0FBFCFB9AFB21FBEE" box="[1138,1158,1026,1047]" italics="true" pageId="4" pageNumber="1063">N</emphasis>
= 2).
</paragraph>
<paragraph id="14404F51383CFFC0FCCFFBB8FCD8FA91" blockId="4.[809,1426,197,1875]" pageId="4" pageNumber="1063">
In surface-dwelling
<taxonomicName id="D3FF34D2383CFFC0FBCFFBB8FB63FBCC" authorityName="bibarba Chen &amp; Chen" authorityYear="2007" box="[1089,1220,1056,1077]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="species" species="bibarba">
<emphasis id="268B9343383CFFC0FBCFFBB8FB63FBCC" box="[1089,1220,1056,1077]" italics="true" pageId="4" pageNumber="1063">B. bibarba</emphasis>
</taxonomicName>
, the skeleton is similar but has the following differences. The bifurcate pharyngeal process of the basioccipital is short and stout, ending at a vertical approximately anterior to centrum 1 (
<figureCitation id="8CC453D4383CFFC0FC58FB02FBEBFB49" box="[982,1100,1178,1200]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="4" pageNumber="1063">Fig. 5C, F</figureCitation>
). The posterior part of the entopterygoid is shallow (
<figureCitation id="8CC453D4383CFFC0FBC1FB21FB05FB36" box="[1103,1186,1209,1231]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="4" pageNumber="1063">Fig. 5D</figureCitation>
). The posteroventral process of the pharyngeal extends posteriorly, forming an obtuse angle with its ventral limb (
<figureCitation id="8CC453D4383CFFC0FB76FB6EFAEAFAF5" box="[1272,1357,1270,1292]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="4" pageNumber="1063">Fig. 5E</figureCitation>
). The anterior frontal is narrower, about 20% as wide as the posterior part (
<figureCitation id="8CC453D4383CFFC0FC51FAACFBEBFAB0" box="[991,1100,1332,1354]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6I, L</figureCitation>
). The vertebral count is 39 (
<emphasis id="268B9343383CFFC0FCBFFACBFCE2FA91" box="[817,837,1363,1384]" italics="true" pageId="4" pageNumber="1063">N</emphasis>
= 2).
</paragraph>
<paragraph id="14404F51383CFFCCFCCFFAE9FE10FE29" blockId="4.[809,1426,197,1875]" lastBlockId="8.[145,761,197,464]" lastPageId="8" lastPageNumber="1067" pageId="4" pageNumber="1063">
Sexual dimorphism is demonstrated in the pectoral fin and in the pectoral girdle, nasal bones and anterior intermuscular bones based on the
<specimenCount id="02F984D8383CFFC0FB52FA37FAFEFA3D" box="[1244,1369,1454,1476]" pageId="4" pageNumber="1063" type="male">two males</specimenCount>
and
<specimenCount id="02F984D8383CFFC0FCA7FA55FC1FFA1A" box="[809,952,1485,1507]" pageId="4" pageNumber="1063" type="female">four females</specimenCount>
(
<figureCitation id="8CC453D4383CFFC0FC46FA55FB98FA1B" box="[968,1087,1485,1507]" captionStart="Figure 1" captionStartId="1.[163,242,688,710]" captionTargetBox="[164,1446,197,647]" captionTargetId="figure-559@1.[163,1446,195,648]" captionTargetPageId="1" captionText="Figure 1. Distribution map and photographs of Bibarba species. A, type locality of B. bibarba (circle) and B. parvoculus (triangle). The inserted map shows the worldwide distribution of the Northern Clade and the Southern Clade of the family Cobitidae with the enlarged area framed. B, Bibarba species, upper: surface-dwelling B. bibarba (HBU20170801, male); lower: cave-dwelling B. parvoculus (LJH 2016120051, male). C, left pectoral fin of B. bibarba (HBU20170801, male) in dorsal view. D, left pectoral fin of B. parvoculus (LJH 2016120051, male) in dorsal view. Arrowheads indicate the lamina circularis on second and third pectoral rays. Scale bar: 10 mm in B, 1.5 mm in C, D." figureDoi="http://doi.org/10.5281/zenodo.5300449" httpUri="https://zenodo.org/record/5300449/files/figure.png" pageId="4" pageNumber="1063">Figs 1C, D</figureCitation>
,
<figureCitation id="8CC453D4383CFFC0FBC3FA55FB2FFA1B" box="[1101,1160,1485,1507]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="4" pageNumber="1063">5C, F</figureCitation>
,
<figureCitation id="8CC453D4383CFFC0FB1BFA55FB04FA1A" box="[1173,1187,1485,1507]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">6</figureCitation>
). In the pectoral fin, a duplication of the lamina circularis on the second and third pectoral rays occurs in males (
<figureCitation id="8CC453D4383CFFC0FA82F992FA26F9D9" box="[1292,1409,1546,1568]" captionStart="Figure 1" captionStartId="1.[163,242,688,710]" captionTargetBox="[164,1446,197,647]" captionTargetId="figure-559@1.[163,1446,195,648]" captionTargetPageId="1" captionText="Figure 1. Distribution map and photographs of Bibarba species. A, type locality of B. bibarba (circle) and B. parvoculus (triangle). The inserted map shows the worldwide distribution of the Northern Clade and the Southern Clade of the family Cobitidae with the enlarged area framed. B, Bibarba species, upper: surface-dwelling B. bibarba (HBU20170801, male); lower: cave-dwelling B. parvoculus (LJH 2016120051, male). C, left pectoral fin of B. bibarba (HBU20170801, male) in dorsal view. D, left pectoral fin of B. parvoculus (LJH 2016120051, male) in dorsal view. Arrowheads indicate the lamina circularis on second and third pectoral rays. Scale bar: 10 mm in B, 1.5 mm in C, D." figureDoi="http://doi.org/10.5281/zenodo.5300449" httpUri="https://zenodo.org/record/5300449/files/figure.png" pageId="4" pageNumber="1063">Fig. 1C, D</figureCitation>
). In the pectoral girdle, the coracoids, mesocoracoid and scapula are much stouter in the male (
<figureCitation id="8CC453D4383CFFC0FB77F9D0FC9BF985" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6D, E, J, K</figureCitation>
) and all are fused with the cleithrum in surfacedwelling
<taxonomicName id="D3FF34D2383CFFC0FC17F91EFBB1F963" authorityName="bibarba Chen &amp; Chen" authorityYear="2007" box="[921,1046,1669,1691]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="species" species="bibarba">
<emphasis id="268B9343383CFFC0FC17F91EFBB1F963" box="[921,1046,1669,1691]" italics="true" pageId="4" pageNumber="1063">B. bibarba</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D4383CFFC0FBA7F91DFB3BF963" box="[1065,1180,1669,1691]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6J, K</figureCitation>
), but autogenous in cave-dwelling
<taxonomicName id="D3FF34D2383CFFC0FC5FF93CFBD6F941" box="[977,1137,1699,1721]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="species" species="parvoculus">
<emphasis id="268B9343383CFFC0FC5FF93CFBD6F941" box="[977,1137,1699,1721]" italics="true" pageId="4" pageNumber="1063">B. parvoculus</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D4383CFFC0FB0CF93BFB50F940" box="[1154,1271,1699,1721]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6D, E</figureCitation>
). The lateral lamina at the ventral limb of the cleithrum is smaller in males than in females of
<taxonomicName id="D3FF34D2383CFFC0FBE9F97AFB79F90F" authorityName="bibarba Chen &amp; Chen" authorityYear="2007" box="[1127,1246,1761,1783]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="1063" phylum="Chordata" rank="species" species="bibarba">
<emphasis id="268B9343383CFFC0FBE9F97AFB79F90F" box="[1127,1246,1761,1783]" italics="true" pageId="4" pageNumber="1063">B. bibarba</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D4383CFFC0FB63F979FAC5F90F" box="[1261,1378,1761,1783]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6H, K</figureCitation>
). In addition, a vertically positioned lamina derived from the second pectoral radial contacting the mesocoracoid (
<figureCitation id="8CC453D4383CFFC0FCBFF8A5FC46F8AB" box="[817,993,1853,1875]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="4" pageNumber="1063">Fig. 6A, D, G, J</figureCitation>
), is much stronger in males (
<figureCitation id="8CC453D4383CFFCCFAB9F8A5FF38FF22" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" lastPageId="8" lastPageNumber="1067" pageId="4" pageNumber="1063">Fig. 6D, J</figureCitation>
). In the neurocranium, a slim laterally convex nasal is lateral to the supraethmoidethmoid complex, which is stouter in males than in females of
<taxonomicName id="D3FF34D23830FFCCFF1FFEBAFEAFFECF" authorityName="bibarba Chen &amp; Chen" authorityYear="2007" box="[145,264,289,311]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="8" pageNumber="1067" phylum="Chordata" rank="species" species="bibarba">
<emphasis id="268B93433830FFCCFF1FFEBAFEAFFECF" box="[145,264,289,311]" italics="true" pageId="8" pageNumber="1067">B. bibarba</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D43830FFCCFE99FEB9FED8FECE" box="[279,383,289,311]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="8" pageNumber="1067">Fig. 6I, L</figureCitation>
), but similar in male and female
<taxonomicName id="D3FF34D23830FFCCFF1FFED8FE8BFEAC" box="[145,300,320,341]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="8" pageNumber="1067" phylum="Chordata" rank="species" species="parvoculus">
<emphasis id="268B93433830FFCCFF1FFED8FE8BFEAC" box="[145,300,320,341]" italics="true" pageId="8" pageNumber="1067">B. parvoculus</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D43830FFCCFEB5FED8FE0EFEAC" box="[315,425,320,342]" captionStart="Figure 6" captionStartId="10.[145,223,1780,1802]" captionTargetBox="[240,1330,197,1740]" captionTargetId="figure-25@10.[239,1330,195,1741]" captionTargetPageId="10" captionText="Figure 6. Pecotral girdle and skull roof of Bibarba species and Cobitis microcephala. AC, B. parvoculus (GTEU20150382, female). DF, B. parvoculus (LJH 2016120051, male). GI, B. bibarba (HBU20170802, female). JL, B. bibarba (HBU20170801, male). MO, Co. microcephala (HBU20170890, male). A, D, G, J, M, pectoral girdle in medial view. B, E, H, K, N, pectoral girdle in lateral view. C, F, I, L, O, skull roof in dorsal view, the bottom of neurocranium removed for" figureDoi="http://doi.org/10.5281/zenodo.5300459" httpUri="https://zenodo.org/record/5300459/files/figure.png" pageId="8" pageNumber="1067">Fig. 6C, F</figureCitation>
). The anterior intermuscular bone associated with vertebrae 4 and 5 is ramified and hatchet-like and stouter in male than in female
<taxonomicName id="D3FF34D23830FFCCFF1FFE04FEA0FE48" authorityName="bibarba Chen &amp; Chen" authorityYear="2007" box="[145,263,412,433]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="8" pageNumber="1067" phylum="Chordata" rank="species" species="bibarba">
<emphasis id="268B93433830FFCCFF1FFE04FEA0FE48" box="[145,263,412,433]" italics="true" pageId="8" pageNumber="1067">B. bibarba</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D43830FFCCFE98FE04FE24FE48" box="[278,387,412,434]" captionStart="Figure 5" captionStartId="9.[163,245,1748,1770]" captionTargetBox="[226,1378,196,1707]" captionTargetId="figure-25@9.[225,1380,195,1709]" captionTargetPageId="9" captionText="Figure 5. Skeleton of surface-dwelling Bibarba bibarba (HBU20170801, male) (except F from the HBU20170802, female). A, radiograph of the whole skeleton. B, cranial skeleton and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone in lateral view. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Scale bar: 5000 µm in A, 1500 µm in BD, F, H, 900 µm in E, G." figureDoi="http://doi.org/10.5281/zenodo.5300457" httpUri="https://zenodo.org/record/5300457/files/figure.png" pageId="8" pageNumber="1067">Fig. 5C, F</figureCitation>
), but similar in male and female
<taxonomicName id="D3FF34D23830FFCCFF1FFE23FE8CFE36" box="[145,299,442,464]" class="Actinopterygii" family="Cobitidae" genus="Bibarba" kingdom="Animalia" order="Cypriniformes" pageId="8" pageNumber="1067" phylum="Chordata" rank="species" species="parvoculus">
<emphasis id="268B93433830FFCCFF1FFE23FE8CFE36" box="[145,299,442,464]" italics="true" pageId="8" pageNumber="1067">B. parvoculus</emphasis>
</taxonomicName>
(
<figureCitation id="8CC453D43830FFCCFEB4FE22FE00FE29" box="[314,423,442,464]" captionStart="Figure 4" captionStartId="7.[163,241,1744,1766]" captionTargetBox="[228,1377,196,1703]" captionTargetId="figure-25@7.[227,1379,195,1705]" captionTargetPageId="7" captionText="Figure 4. Skeleton of cave-dwelling Bibarba parvoculus (LJH 2016120051, male) (except F from a female GTEU20150382). A, radiograph of the whole skeleton. B, cranial skeleton, pectoral girdle and associated anterior vertebrae. C, neurocranium. D, suspensorium plus branchiostegals. E, G, pharyngeal bone. F, neurocranium of a female. H, radiograph of the caudal skeleton. All pictures are in lateral view except G in dorsal view. Abbreviations: ach, anterior ceratohyal; bc, bony capsule; boc, basioccipital; boc pp, pharyngeal process of basioccipital; br, branchiostegal ray; den, dentary; ect, ectopterygoid; ent," figureDoi="http://doi.org/10.5281/zenodo.5300455" httpUri="https://zenodo.org/record/5300455/files/figure.png" pageId="8" pageNumber="1067">Fig. 4C, F</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>