diff --git a/data/03/C3/87/03C387E4FFF3FFBBFD83FE9B14A7FE6F.xml b/data/03/C3/87/03C387E4FFF3FFBBFD83FE9B14A7FE6F.xml new file mode 100644 index 00000000000..caf639fa219 --- /dev/null +++ b/data/03/C3/87/03C387E4FFF3FFBBFD83FE9B14A7FE6F.xml @@ -0,0 +1,3360 @@ + + + +A new species of Knodus (Characiformes: Characidae), with deep genetic divergence, from the Mearim and Munim river basins, Northeastern Brazil, and evidence for hidden diversity in adjacent river basins + + + +Author + +Aguiar, Rayane G. + + + +Author + +Guimarães, Erick C. +Universidade Federal do Oeste do Pará, Programa de Pós-Graduação Sociedade Natureza e Desenvolvimento, Instituto de Ciências da Educação, Av. Mendonça Furtado, 2946, Fátima, 68040 - 470 Santarém, PA, Brazil. erick. ictio @ yahoo. com. br (corresponding author). & Universidade Estadual do Maranhão, Laboratório de Ictiofauna e Piscicultura Integrada, Centro de Ciências Agrárias, Cidade Universitária Paulo VI, 65055 - 310 São LuÍs, MA, Brazil. (JPS) jadsonsantos @ professor. uema. br. + + + +Author + +Brito, Pâmella S. de + + + +Author + +Santos, Jadson P. +Universidade Estadual do Maranhão, Laboratório de Ictiofauna e Piscicultura Integrada, Centro de Ciências Agrárias, Cidade Universitária Paulo VI, 65055 - 310 São LuÍs, MA, Brazil. (JPS) jadsonsantos @ professor. uema. br. + + + +Author + +Katz, Axel M. +Universidade Federal do Rio de Janeiro, Laboratório de Sistemática e Evolução de Peixes Teleósteos, Departamento de Zoologia, Instituto de Biologia, Cidade Universitária, 21941 - 599 Rio de Janeiro, RJ, Brazil. axelmk @ gmail. com. & S / N, Boa Vista, 65500 - 000 Chapadinha, MA, Brazil. fpottoni @ gmail. com. +Brazil.axelmk@gmail.com + + + +Author + +Dias, Luiz Jorge B. da S. +Universidade Estadual do Maranhão, Centro de Educação, Ciências Exatas e Naturais. Cidade Universitária Paulo VI, Tirirical 65055 - 970 São LuÍs, MA, Brazil. luizjorgedias @ hotmail. com. +Brazil.luizjorgedias@hotmail.com + + + +Author + +Carvalho-Costa, Luis Fernando + + + +Author + +Ottoni, Felipe P. +Universidade Federal do Maranhão, Centro de Ciências Agrárias e Ambientais, Laboratório de Sistemática e Ecologia de Organismos Aquáticos, Campus de Chapadinha, BR- 222, km + +text + + +Neotropical Ichthyology + + +2022 + +e 210173 + + +2022-06-03 + + +20 + + +2 + + +1 +27 + + + + +http://dx.doi.org/10.1590/1982-0224-2021-0173 + +journal article +10.1590/1982-0224-2021-0173 +1982-0224 +13350457 +BE91D7F5-F56D-40B3-AA1D-A9C974083BC6 + + + + + + +Knodus guajajara +Aguiar, Brito, Ottoni & Guimarães + +, +new species + + +urn:lsid:zoobank.org:act: +4EEDDC6E-4088-4B60-828D-AF79EF84DC26 + + + + + +( +Figs. 1–3 +; +Tabs. 2–3 +) + + + +Knodus victoriae + +[non + +Knodus victoriae +(Steindachner, 1907) + +]. ―Guimarães +et al +. (2020a:90). ―Oliveira +et al +. (2020:6–7). + + + + + + +Holotype +. + +CICCAA 4883 +, +31.4 mm +SL, +Alto Alegre do Pindaré +municipality, +Igarapé Arapapá +, +Pindaré River +drainage, +Mearim River +basin, +03°42’26”S +46°00’25”W +, + +Nov 2016 + +. +E. C. Guimarães +& +P. S. Brito. + + + + + +FIGURE 1 | + +Knodus guajajara + + +, holotype, CICCAA 4883, 31.4 mm SL, Alto Alegre do Pindaré municipality, Igarapé Arapapá, Pindaré River drainage, Mearim River basin. + + + + + +FIGURE 2 | + +Knodus guajajara + + +, paratypes. +A. +CICCAA 1518, 22.8 mm SL, Brazil, Maranhão, Alto Alegre do Pindaré municipality, igarapé Jenipapo, Pindaré River drainage, Mearim River basin. +B. +CICCAA 2696, 40.2 mm SL, Brazil, Maranhão, Chapadinha municipality, stream in riparian forest on the road BR–222, Munim River basin. + + + + +TABLE 1 | +Specimens and DNA sequence information included in the study. Sequences made available by this study follow with asteristic. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
NºSpeciesDepositories of vouchersGenbank ID
CollectionTissue code +Voucher +
1 + +Bryconamericus exodon + +LBP261915118 +MH +002960 +
2 + +Bryconamericus iheringii + +LBP6064514473 +MH +002991 +
3 + +Knodus alpha + +STRI9564531 +MH +003217 +
4 + +Knodus alpha + +LBP9561663 +MH +003216 +
5 + +Knodus borki + +LBP5382012472 +MH +003218 +
6 + +Knodus borki + +LBP–53821KT248111
7 + +Knodus caquetae + + +MPUJ +63711073A +MH +003221 +
8 + +Knodus caquetae + + +MPUJ +63911073B +MH +003222 +
9 + +Knodus heteresthes + +LBP5737113870 +MH +003232 +
10 + +Knodus heteresthes + +LBP373177959 +MH +003233 +
11 + +Knodus megalops + +LBP5422312566B +MH +003234 +
12 + +Knodus meridae + +LBP158187569 +MH +003235 +
13 + +Knodus tiquiensis + +LBP–33217KT248096
14 + +Knodus tiquiensis + +LBP–33218KT248097
15 + +Knodus tiquiensis + +LBP–33216KT248098
16 + +Knodus victoriae + +LBP–27366KT248128
17 + +Knodus victoriae + +LBP27370KT248129
18 + +Knodus victoriae + +LBP273425607KT248130
19 + +Knodus victoriae + +LBP–27254KT248131
20 + +Knodus victoriae + +LBP–27368KT248132
21 + +Knodus victoriae + +LBP–27295KT248133
22 + +Knodus victoriae + +LBP27369KT248134
23 + +Knodus victoriae + +LBP–27253KT248135
24 + +Knodus victoriae + +LBP–27255KT248136
25 + +Knodus victoriae + +LBP–27367KT248137
26 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4817.14817 +MW +556675 +
27 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4817.24817 +MW +556676 +
28 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4817.34817 +MW +556677 +
29 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4818.14818 +MW +556678 +
30 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4818.24818 +MW +556679 +
31 + +Knodus +aff. +victoriae +(Balsas) + +* + +CICCAA +4818.34818 +MW +556680 +
32 + +Knodus +aff. +victoriae +(Itapecuru) + +* + +CICCAA +2064.12064 +MW +556681 +
33 + +Knodus +aff. +victoriae +(Itapecuru) + +* + +CICCAA +2064.22064 +MW +556682 +
34 + +Knodus +cf. +savannensis + +LBP–66360KT248199
+ + + +TABLE 1 | +(Continued) + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Nº + +Species + +Depositories of vouchers + +Genbank ID +
+Collection + +Tissue code + +Voucher +
35 + +Knodus +cf. +savannensis + +LBP–66339KT248200
36 + +Knodus +cf. +savannensis + +LBP–66340KT248201
37 + +Knodus +cf. +savannensis + +LBP–66356KT248202
38 + +Knodus +cf. +savannensis + +LBP–66362KT248203
39 + +Knodus +cf. +savannensis + +LBP–66363KT248204
40 + +Knodus +cf. +savannensis + +LBP–66363KT248218
41 + +Knodus +cf. +savannensis + +LBP–62500KT248219
42 + +Knodus +cf. +savannensis + +(Carolina-MA, Tocantins River basin)* +CICCAA3609.13609MW556695
43 + +Knodus +sp. + +“Guamá” +LBP430709141KT248223
44 + +Knodus +sp. + +“Guamá” +LBP–43718KT248225
45 + +Knodus +sp. + +“Guamá” +LBP–43067KT248226
46 + +Knodus +sp. + +“Guamá” +LBP–43068KT248227
47 + +Knodus +sp. + +“Tapajós” +LBP5704813750KT248123
48 + +Knodus +sp. + +“Tapajós” +LBP–57049KT248124
49 + +Knodus +sp. + +“Xingu” +LHGP–66314KT248184
50 + +Knodus +sp. + +“Xingu” +LHGP–66315KT248185
51 + +Knodus +sp. + +(Marabá)* +CICCAA2087.12087MW556692
52 + +Knodus +sp. + +(Marabá)* +CICCAA2087.22087MW556693
53 + +Knodus +sp. + +(Marabá)* +CICCAA2087.32087MW556694
54 + +Knodus +sp. + +(Maracaçumé)* +CICCAA2390.12390MW556683
55 + +Knodus +sp. + +(Maracaçumé)* +CICCAA2390.22390MW556684
56 + +Knodus +sp. + +(Maracaçumé)* +CICCAA2390.52390MW556685
57 + +Knodus guajajara +* + +CICCAA2052.22052MW556688
58 + +Knodus guajajara +* + +CICCAA2052.32052MW556689
59 + +Knodus guajajara +* + +CICCAA2052.42052MW556690
60 + +Knodus guajajara +* + +CICCAA2052.52052MW556691
61 + +Knodus guajajara +* + +CICCAA2391.12391MW556686
62 + +Knodus guajajara +* + +CICCAA2391.22391MW556687
+ + + + +Paratypes +. + +All from +Brazil +, +Maranhão +State: +CICCAA 1535 +, +1 +, +29.2 mm +SL, collected with holotype. + + +CICCAA 1585 +, +3 +, 28.0– +34.7 mm +SL, +Alto Alegre do Pindaré +municipality, igarapé +Arapapá +, +Pindaré River +drainage, +Mearim River +basin, +03°42’26”S +46°00’25”W +, + +Nov 2015 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4860 +, +2 +C&S, +29.9–31.6 mm +SL, +Alto Alegre do Pindaré +municipality, igarapé +Arapapá +, +Pindaré River +drainage, +Mearim River +basin, +03°42’26”S +46°00’25”W +, + +Nov 2015 + +, +E. C. Guimarães +& +P. S. Brito. + + +LBP 31041, 16, +24.5–31.4 mm +SL, +Alto Alegre do Pindaré +municipality, +Igarapé Igarapá +, +Pindaré River +drainage, +Mearim River +basin, +03°45’51”S +46°08’15”W +, + +Nov 2015 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4858 +, +5 +C&S, 22.9–27.0 mm SL, +Alto Alegre do Pindaré +municipality, +Igarapé Igarapá +, +Pindaré River +drainage, +Mearim River +basin, +03°45’51”S +46° 08’15”W +, + +Nov 2015 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 1517 +, +2 +, +21.8–24.4 mm +SL, +Alto Alegre do Pindaré +municipality, igarapé +Caititu +, +Pindaré River +drainage, +Mearim River +basin, +03°42’30”S +46°01’19”W +, + +Jul 2017 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4859 +, +2 +C&S, +23.5–27.6 mm +SL, +Alto Alegre do Pindaré +municipality, igarapé +Caititu +, +Pindaré River +drainage, +Mearim River +basin, +03°42’30”S +46°01’19”W +, + +Jul 2017 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 1536 +, +2 +, +28.8–29.8 mm +SL, +Buriticupu +municipality, +Buritizinho River +, +Pindaré River +drainage, +Mearim River +basin, +04°11’53”S +46°28’41”W +, + +Nov 2016 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4861 +, +1 +C&S, +31.9 mm +SL, +Buriticupu +municipality, +Buritizinho River +, +Pindaré River +drainage, +Mearim River +basin, +04°11’53”S +46°28’41”W +, + +Nov 2016 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 1227 +, +1 +, +29.6 mm +SL, +Buriticupu +municipality, +Buritizinho River +, +Pindaré River +drainage, +Mearim River +basin, +04°19’45”S +46°29’46”W +, + +27 Jan 2017 + +, +E. C. Guimarães +& +P. S. Brito. + + +UFRJ +7023, 10, +22.8–29.3 mm +SL, +Alto Alegre do Pindaré +municipality, igarapé +Jenipapo +, +Pindaré River +drainage, +Mearim River +basin, +03°51’20”S +46°11’09”W +, + +Jul 2017 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4862 +, +5 +C&S, +23.9–29.3 mm +SL, +Brazil +, +Maranhão State +, +Alto Alegre do Pindaré +municipality, igarapé +Jenipapo +, +Pindaré River +drainage, +Mearim River +basin, +03°51’20”S +46°11’09”W +, + +Jul 2017 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 1321 +, +39 +, +19.4–29.2 mm +SL, +Miranda do Norte +municipality, +Mearim River +basin, +04°19’45”S +46°29’46”W +, + +Nov 2016 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 4863 +, +11 +C&S, +19.7–24.4 mm +SL, +Miranda do Norte +municipality, +Mearim River +basin, +04°19’45”S +46°29’46”W +, + +Nov 2016 + +, +E. C. Guimarães +& +P. S. Brito. + + +CIUEMA 1021, 8, +21.2–26.9 mm +SL, +Alto Alegre do Pindaré +municipality, +Igarapé Timbira +, +Pindaré River +drainage, +Mearim River +basin, +03°32’57”S +44°39’38”W +, + +Nov 2015 + +, +E. C. Guimarães +& +P. S. Brito. + + +CICCAA 3423 +, +42 +, +19.5–30.21 mm +SL, + + +CICCAA 3424 +, +57 +, +22.35–26.9 mm +SL,, +Chapadinha +municipality, + +Riacho +da Raiz + +, +RESEX +Chapada Limpa, +Munim River +basin, +03°53’45”S +43°29’21”W +, + +11 Aug 2019 + +, +J. Reis +, +L. Oliveira +, +F. Ottoni +, +R +. Fernandes & + +A. Silva. + +Chapadinha municipality, +Bandeira River +, +Povoado Mata do Jeroca +, +RESEX +Chapada Limpa, +Munim River +basin, +03°59’40”S +43°29’24”W +, + +11 Aug 2019 + +, +J. Reis +, +L. Oliveira +, +F. Ottoni +, +R +. +Fernandes +& +A. Silva. + + + + +CICCAA 3425 +, +22 +, +20.22–26.71 mm +SL, + + +CICCAA 3426 +, +6 +, +28.87–32.6 mm +SL, +Chapadinha +municipality, stream at +Bairro Aldeia +, +Munim River +basin, +03°44’53”S +43°21’32”W +, + +28 Jan 2019 + +, +F. Carvalho +, +H. Silva +, +L. Oliveira +& +I. Gôuvea. + + +CICCAA 3467 +, +2 +, +21.22–21.43 mm +SL, +Anapurus +municipality, stream on the road at +Povoado de Paços +, +Munim River +basin, +03°33’44”S +43°03’52”W +, + +Sep 2019 + +, +D. Campos +, +J. Reis +& +F. Ottoni. + + +CICCAA 4808 +, +3 +, +25.19–34.2 mm +SL, + + +CICCAA 4822 +, +1 +, +36.89 mm +SL, +Chapadinha +municipality, stream at balneário +Repouso do Guerreiro +, +Bairro Independência +, +Munim River +basin, +03°44’57”S +43°20’26”W +, + +Nov 2019 + +, +B. Furtado +, +M. Paiva +, +A. Bezerra +, +M. Coelho +& +I. Gouvêa. + + +CICCAA 4490 +, +5 +, +24.73–29.11 mm +SL, +Chapadinha +municipality, +Riacho Fundo +, +Munim River +basin, +03°42’20”S +43°31’46”W +, + +23 Sep 2018 + +, +L. Sousa +, +L. Oliveira +& +I. Gouvêa +, + + +CICCAA 4491 +, +3 +C&S +23.26–28.19 mm +SL, +Chapadinha +municipality, +Riacho Fundo +, +Munim River +basin, +03°42’20”S +43°31’46”W +, + +23 Sep 2018 + +, +L. Sousa +, +L. Oliveira +& +I. Gôuvea +. + + + + + +FIGURE 3 | + +Knodus guajajara +, CICCAA + +4861, paratype, 31.9 mm SL, jaw suspensorium. +A. +Premaxillary. +B. +Maxilla. +C. +Dentary. Scale bar = 1 mm. + + + + +TABLE 2 | +Morphometric data (N = 110) of the holotype and paratypes of + +Knodus guajajara + +from the Mearim River basin. SD = Standard deviation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Holotype + +Paratypes + +Mean + +SD +
Standard length31.4 +19.4 +‒ +34.7 +25.3 +‒ +
+Percents of standard length +
Depth at dorsal-fin origin (body depth)29.8 +20.1 +‒ +30.2 +24.73.3
Snout to dorsal-fin origin55.3 +47.8 +‒ +57.1 +52.25.5
Snout to pectoral-fin origin27.1 +21.7 +‒ +29.32 +24.93.0
Snout to pelvic-fin origin45.7 +37.6 +‒ +49.3 +42.24.9
Snout to anal-fin origin60.2 +44.0 +‒ +61.6 +53.96.1
Caudal peduncle depth10.5 +7.6 +‒ +11.7 +9.31.2
Caudal peduncle length15.3 +7.1 +‒ +15.2 +11.11.9
Pectoral-fin length20.5 +14.8 +‒ +23.8 +18.72.6
Pelvic-fin length13.5 +9.8 +‒ +15.4 +12.61.7
Dorsal-fin base length9.1 +8.7 +‒ +15.2 +11.51.7
Dorsal-fin height26.9 +17.7 +‒ +27.0 +21.32.8
Anal-fin base length29.4 +25.5 +‒ +36.3 +30.53.8
Anal-fin lobe length13.0 +9.8 +‒ +20.3 +15.32.4
Eye to dorsal-fin origin42.5 +33.6 +‒ +43.8 +38.94.5
Dorsal-fin origin to caudal-fin base52.5 +39.4 +‒ +56.7 +46.95.8
+Percents of head length +
Head length23.9 +21.3 +‒ +26.5 +23.32.5
Horizontal eye diameter41.3 +32.6 +‒ +45.6 +39.54.8
Snout length24.9 +15.8 +‒ +29.6 +21.83.3
Least interorbital width35.1 +23.7 +‒ +36.8 +32.34.0
Upper jaw length47.5 +31.5 +‒ +48.8 +38.35.3
+ + + +TABLE 3 | +Morphometric data (N = 136) for the paratypes of +Knodus guajajara +from the Munim River basin. SD = Standard deviation. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Range + +Mean + +SD +
Standard length +19.5 +‒ +37.7 +25.3 +‒ +
+Percents of standard length +
Depth at dorsal-fin origin (body depth) +20.4 +‒ +37.2 +24.72.06
Snout to dorsal-fin origin +47.4 +‒ +53.4 +50.31.13
Snout to pectoral-fin origin +21.9 +‒ +27.1 +24.40.79
Snout to pelvic-fin origin +39.4 +‒ +45.7 +42.51.24
Snout to anal-fin origin +53.1 +‒ +59.8 +56.21.38
Caudal peduncle depth +8.3 +‒ +10.4 +9.30.41
Caudal peduncle length +9.2 +‒ +12.8 +10.40.75
Pectoral-fin length +18.5 +‒ +23.4 +20.90.74
Pelvic-fin length +12.0 +‒ +15.7 +13.80.68
Dorsal-fin base length +8.4 +‒ +12.6 +10.10.76
Dorsal-fin height +20.1 +‒ +24.6 +22.10.86
Anal-fin base length +22.8 +‒ +28.7 +26.41.17
Anal-fin lobe length +16.1 +‒ +20.2 +18.10.82
Eye to dorsal-fin origin +33.1 +‒ +39.2 +36.11.12
Dorsal-fin origin to caudal-fin base +47.4 +‒ +54.3 +49.51.14
+Percents of head length +
Head length +21.5 +‒ +25.7 +23.40.82
Horizontal eye diameter +33.8 +‒ +42.6 +37.41.56
Snout length +20.1 +‒ +25.9 +22.51.34
Least interorbital width +27.1 +‒ +35.3 +31.71.77
Upper jaw length +34.5 +‒ +41.2 +37.91.47
+ + + +Morphological diagnosis. + +Knodus guajajara + +differs from + +K. borki +Zarske, 2008 + +and + +K. delta +Géry, 1972 + +by having a complete lateral line ( +vs +. incomplete lateral line) and from + +K. cupariensis +de Sousa, Silva-Oliveira, Canto & Ribeiro, 2020 + +and + +K. geryi + +Lima +, Britski & Machado, 2004 by having caudal fin lobes with sparse chromatophores and lacking basal blotches ( +vs. +a dark basal blotch on each caudal fin lobe, Sousa +et al. +, 2020; fig. 1); from + +Knodus borki + +, + +K. diaphanus +(Cope, 1878) + +, + +K. victoriae + +, + +K. heteresthes +(Eigenmann, 1908) + +, + +K. deuterodonoides +(Eigenmann, 1914) + +, + +K. longus +Zarske & Géry, 2006 + +, + +K. septentrionalis +Géry, 1972 + +, + +K. figueiredoi +Esguícero & Castro, 2014 + +, + +K. gery + +i, + +K. meridae +Eigenmann, 1911 + +, + +K. nuptialis +Menezes & Marinho, 2019 + +, + +K. orteguasae +(Fowler, 1943) + +, + +K. tiquiensis +Ferreira + +& +Lima +, 2006, + +K. angustus +Menezes, Ferreira & Netto-Ferreira, 2020 + +, + +K. rufford +Deprá, Ota, Vitorino-Júnior & Ferreira, 2021 + +and + +K. obolus +Deprá, Ota, Vitorino-Júnior & Ferreira, 2021 + +by having 20–25 branched rays in the anal-fin (mode 23) ( +vs. +12–19, combined); and from + +K. tiquiensis + +by having a single humeral spot ( +vs. +two). + +Knodus guajajara + +is distinguished from + +K. breviceps +(Eigenmann, 1908) + +and + +K. savannensis +Géry, 1961 + +by having a conspicuous round humeral blotch ( +vs. +inconspicuous and vertically elongate); from + +K. dorsomaculatus +Ferreira & Netto-Ferreira, 2010 + +by having a hyaline dorsal-fin ( +vs. +dark blotch on the base of the first five branched dorsal fin rays); from + +K. alpha +(Eigenmann, 1914) + +, + +K. chapadae +(Fowler, 1906) + +, + +K. geryi + +, + +K. hypopterus +(Fowler, 1943) + +, + +K. mizquae +(Fowler, 1943) + +and + +K. shinahota +Ferreira & Carvajal, 2007 + +by having 4 or 5 rows of scales between the lateral line and the dorsal-fin origin ( +vs. +6 rows of scales); from + +K. cinarucoensis +(Román-Valencia +et al. +, 2008) + +, + +K. gamma +Géry, 1972 + +and + +K. longus +Zarske & Géry, 2006 + +by having 12 or 13 scales in the median series between the tip of the supraoccipital spine and the dorsal-fin origin ( +vs. +10 or 11 rows of scales in + +K. gamma + +and +17 to 18 in + +K. longus + +); from + +K. jacunda +(Fowler, 1913) + +, + +K. moenkhausii +(Eigenmann & Kennedy, 1903) + +, + +Knodus cismontanus +(Eigenmann, 1914) + +, + +Knodus caquetae +Fowler, 1945 + +, + +K. tanaothoros +(Weitzman, Menezes, Evers & Burns, 2005) + +, + +K. weitzmani +(Menezes, Netto-Ferreira & Ferreira, 2009) + +and by having 3 to 5 maxillary teeth ( +vs. +absence in + +K. jacunda + +, one in + +K. caquetae + +, and +2 in + +K. moenkhausii + +, + +K. tanaothoros +, +K. weitzmani + +and + +K. cismontanus + +); from + +K. megalops +Myers, 1929 + +by having 3 or 4 tricuspid teeth in the premaxillary outer row ( +vs. +5); from + +K. jacunda + +by having 3 to 5 maxillary teeth ( +vs +. absence); from + +K. smithi +(Fowler, 1913) + +by having 3 to 5 cusps on the teeth of the inner row of the premaxilla ( +vs. +7); and from + +K. figueredoi + +, + +K. heteresthes + +, + +K. meridae + +, + +K. mizquae + +, + +K. moenkhausii + +, + +K. victoriae + +, + +K. pasco +Zarske, 2007 + +by having 12 circumpeduncular scales ( +vs. +13–14 combined). Furthermore, + +Knodus guajajara + +differs from + +K. victoriae + +by having more total vertebrae 33–35, mode 34 ( +vs. +30–33, mode 32). + + + + +Description. +Morphometric data presented in +Tabs. 2–3 +. Body comparatively small, with largest specimen examined measuring +37.7 mm +SL. Greatest body depth at dorsal-fin origin. Dorsal profile of head convex from upper lip to vertical through middle portion of eye; slightly concave from this point to tip of supraoccipital spine; straight to slightly convex from posterior tip of supraoccipital spine to dorsal-fin origin; dorsal-fin base straight; slightly convex to straight from end of dorsal-fin base to adipose fin and concave from latter point to anterior dorsal-procurrent ray. Ventral profile of body convex from lower lip to anal-fin origin; straight, posterodorsally inclined along anal-fin base. Dorsal and ventral profile of caudal peduncle slightly concave. + + +Mouth sub-terminal; jaws isognathous. Posterior terminus of maxilla extending beyond anterior margin of orbit. Premaxillary teeth in two rows; outer row with 3(18) and 4(8) tricuspid teeth, inner row with 4(26) tri-to pentacuspid teeth. Maxilla with 3(11), 4(13), 5(2) tri-to pentacuspid teeth. Dentary with three anteriormost teeth large, tri and pentacuspid teeth, followed by 4(2), 5(2), 6(4), 7(13) and 8(5) smaller, uni, tricuspid or pentacuspid teeth ( +Fig. 3 +). + +Scales cycloid, moderately large, with 6–8 well-marked radii; circuli only present proximally. Lateral line complete, slightly curved ventrally along its anterior third, with 36(8), 37(61), 38*(137), 39(13), or 40(7) perforated scales. Longitudinal series of scales between dorsal-fin origin and lateral line 4(30) or 5*(202). Longitudinal series of scales between lateral line and pelvic-fin origin 4*(63), or 5(184). Predorsal scales 11(3), 12(106) or 13*(122). Circumpeduncular scales 12*(220). Single series of scales on anal-fin base. +Dorsal fin rays ii,7(45) or 8*(191). Dorsal-fin origin slightly posterior to middle of body, and slightly posterior to vertical through pelvic-fin origin. First unbranched ray approximately half length of second unbranched ray. Pectoral-fin rays i,9(9), 10(142) or 11*(94); tip of pectoral fin reaching or exceeds pelvic fin origin when adpressed. Pelvic-fin rays i,5,i(41) or i,6,i*(189); tip of pelvic fin usually reaching first anal fin rays when adpressed. Anal-fin rays iv, 20(2), 21(8), 22(40), 23*(140), 24(37) or 25(15). Anal-fin origin located at vertical through middle of dorsal fin. Anal-fin rays decreasing gradually in length. Caudal fin forked, lobes equal in size. Principal caudal-fin rays i,9,8,i(219). Dorsal procurrent caudal-fin rays 8(1), 9(1), 10(18), 11(3) or 12(3) and ventral procurrent caudal-fin rays 9(3), 10(2), 11(15), 12(3) or 13(3). Adipose-fin origin approximately at vertical through base of 19th to 21st branched anal-fin rays. Gill rakers on first gill arch 15(2), 16(4) or 17(2): hypobranchial 2(4) or 3 (4), ceratobranchial 7(8), cartilage between epibranchial and ceratobranchial 1(8), epibranchial 5(4) or 6(4). Gill rakers setiform. Branchiostegal rays 4(8), 3(8) on anterior ceratohyal and 1(8) on posterior ceratohyal. Total vertebrae 33(3), 34(18) or 35(8). First dorsal fin pterygiophore inserted between 10th and 12th vertebrae. First anal fin pterygiophore inserted between 16th and 17th vertebrae. Supraneurals 4(2), 5(12) or 6(9). + +Color in alcohol. +Ground of body and head light brown, with dorsal region of body (especially dorsal profile), dorsal portion of head (just above eyes), and upper snout area (upper jaw and adjacent areas) darker. Ventral region of trunk and head with light brown coloration, especially on scales adjacent to anal-fin base. Dark colored chromatophores concentrated on scales of dorsal region (from both body and head), on opercle, and lateral band of flank (especially on posterior region, near caudal peduncle and caudal-fin base). Conspicuous vertically elongated humeral spot, with some dark scattered chromatophores above and below this blotch. Lateral band formed by chromatophores located above lateral line, extending gradually from humeral blotch to middle rays of caudal fin, where they become thicker and more concentrated. All fins including adipose hyaline, with chromatophores on inter-radial membranes of dorsal fin. Anal and caudal fins with posterior margins dark grey. + + +Color in life. +Yellowish coloration in dorsolateral region and silver coloration in ventrolateral region of body ( +Fig. 2 +). Gray lateral band located in middle portion of trunk, with bright pigment extending from operculum to caudal-fin base. Dorsal portion of eye yellowish-orange and lower portion silver; mouth and dorsolateral portion of head yellowish with concentrated chromatophores; operculum with silver coloration. Fins hyaline with chromatophores present and especially concentrated in medial region of dorsal fin, distal region of anal and caudal fins, on first pectoral-fin ray, and just posterior to medial lateral band on caudal fin. Adipose fin with yellowish-orange pigmentation; yellowish-orange pigments also present in middle region of caudal fin. + + +Sexual dimorphism. +Hooks present on pelvic fins and anal fin of sexually mature males. Pelvic fins with 5 to 14 hooks per branched ray; anal fin with 2 to 8 hooks per ray. Hooks concentrated on anterior anal fin rays ( +Fig. 4 +). + + + + +Geographical distribution. + +Knodus guajajara + +occurs in the rivers, streams, and lagoons of the Mearim and Munim river basins, in the State of +Maranhão +, Northeast +Brazil +( +Fig. 5 +). + + + + +Etymology. +The specific epithet honors the Guajajara indigenous tribe, which is one of the most numerous indigenous peoples in +Brazil +. They inhabit more than 10 Indigenous Lands on the eastern margin of the Amazon, all located in +Maranhão +. A noun in apposition the language spoken by them is the Teneteara, from the Tupi-Guarani linguistic family. + + + + +Conservation status. + +Knodus guajajara + +occurs in two distinct river basins of +Maranhão +, the Mearim and Munim river basins. It was found in several collecting sites along these basins, and it is always abundant. Therefore, according to the guidelines given in the International Union for Conservation of Nature (IUCN) categories and criteria (IUCN, 2019), we recommend that + +Knodus guajajara + +should be categorized as Least Concern (LC). + + + + +Molecular species delimitation + + +Molecular diagnosis. + +Knodus guajajara + +is diagnosed molecularly by a combination of 19 nucleotide substitutions, four of them unique* (listed below): + +COI 279 (C→A), COI 282 (C→T), COI 306 (C→T), COI 327 (C→T), COI 357 (A→G*), COI 372 (A→G*), COI 378 (A→G), COI 414 (T→C), COI 450 (T→C), COI 459 (C→T), COI 510 (T→C), COI 528 (C→T), COI 537 (C→T), COI 565 (C→T), COI 585 (T→C), COI 589 (C→T), COI 597 (C→T), COI 627 (A→G*), COI 645 (T→A*). + +Traditional DNA Barcoding. +COI sequences support the existence of a new species of + +Knodus + +inhabiting the Mearim and Munim River basins. After trimming, the final alignment yielded 401 base pairs with 132 variable sites and 62 haplotypes. + +Knodus guajajara + +is 14% divergent, on average, from the other taxa, with a minimum genetic distance of 11% to + +K. borki + +, + +K. caquetae + +and + +Knodus +sp. + +“Maracaçumé”, and a maximum genetic distance of 16% to + +Bryconamericus exodon +Eigenmann,1907 + +and + +Knodus tiquiensis +. + +The genetic distance between the new species and + +K +. +victoriae + +is 13% ( +Tab. 4 +). + + + +FIGURE 4 | + +Knodus guajajara +, + +CICCAA 4861, paratype, male, 31.9 mm SL Maranhão, Mearim River basin. +A. +Hooks on pelvic fin. +B. +Hooks on anal fin. (Photographed by F. P. Ottoni). + + + + +FIGURE 5 | +Geographical distribution of + +Knodus guajajara + +. + + + + +FIGURE 6 | +Bayesian Inference phylogenetic tree generated by this study. The results of GMYC, bPTP and ABGD are included. Numbers above and below branches are posterior probability values. The gray bars correspond to the congruent results of the different species delimitation methods (Con.). + + + +Phylogenetic analysis. +The BI phylogenetic analysis delimited 19 lineages: two of them are outgroups ( + +Bryconamericus +species + +) and the remaining correspond to + +Knodus +species + +( +Fig. 6 +). One of the 17 lineages of + +Knodus + +represents the new species described herein ( + +Knodus guajajara + +) with haplotypes from the Mearim and Munim river basins forming an exclusive clade supported by the highest possible support value (PP = 1) ( +Fig. 6 +). The BI phylogenetic analysis also delimited new lineages of + +Knodus + +inhabiting the +Tocantins +River ( + +Knodus +sp. + +“Marabá”) and the Maracaçumé River ( + +Knodus +sp. + +“Maracaçumé”). In addition, two well supported clades were recovered within + +Knodus +cf. +savannensis + +: the first one containing + +Knodus +cf. +savannensis + +( +Tocantins +River, Carolina, MA), + +Knodus + +“Guamá” and + +Knodus +cf. +savannensis + +(KT248218 and KT248219) (posterior probability = 0.96). The second clade of + +Knodus +cf. +savannensis + +corresponds to six haplotypes (KT248199, KT248200, KT248201, KT248202, KT248203, and KT248204) of + +Knodus +cf. +savannensis + +(PP = 1). Two other lineages closely related to + +K. victoriae + +were detected: + +Knodus +cf. +victoriae + +(Balsas River drainage) and + +Knodus +cf. +victoriae + +(Itapecuru River basin) (PP = 1). + + + +TABLE 4 | +Kimura-2-parameters pairwise genetic distances among species. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Species + +1 + +2 + +3 + +4 + +5 + +6 + +7 + +8 + +9 + +10 + +11 + +12 + +13 + +14 + +15 + +16 + +17 + +18 +
+1 + + +Bryconamericus exodon + +
+2 + + +Bryconamericus iheringii + +0.12
+3 + + +Knodus heteresthes + +0.150.16
+4 + + +Knodus + +cf. s +avanensis +(Clade 1) +0.130.170.09
+5 + + +Knodus + +cf. s +avanensis +(Clade 2) +0.130.160.080.06
+6 + + +Knodus +sp. + +“Tapajós” +0.130.180.080.100.09
+7 + + +Knodus +sp. + +“Xingu” +0.150.160.050.090.100.09
+8 + + +Knodus tiquiensis + +0.140.180.130.110.110.120.14
+9 + + +Knodus borki + +0.110.150.120.120.110.110.120.15
+10 + + +Knodus megalops + +0.130.140.140.140.140.120.130.150.08
+11 + + +Knodus caquetae + +0.110.140.140.140.130.120.140.150.070.04
+12 + + +Knodus meridae + +0.160.160.120.110.100.120.130.150.110.130.13
+13 + + +Knodus alpha + +0.140.180.100.060.090.130.090.150.140.140.150.15
+14 + + +Knodus +sp. + +“Marabá” +0.140.160.160.130.120.140.150.140.150.150.150.170.15
+15 + + +Knodus +sp. + +“Maracaçumé” +0.120.140.120.130.110.120.130.110.110.120.100.120.150.14
+16 + + +Knodus victoriae + +0.150.180.070.080.080.100.080.130.110.150.140.120.100.160.12
+17 + + +Knodus +aff. +victoriae +(Balsas) + +0.130.190.090.090.090.100.100.130.120.130.150.100.100.140.140.05
+18 + + +Knodus +aff. +victoriae +(Itapecuru) + +0.140.150.100.090.090.100.110.130.120.110.140.120.090.140.120.080.06
19 + +Knodus guajajara + +0.160.130.150.130.130.140.140.160.110.130.110.130.130.130.110.130.150.13
+ + +GMYC, bPTP and ABGD. +The results of the three species delimitation methods (GMYC, bPTP, and ABGD) were congruent ( +Fig. 6 +), and all three delimited the same 19 lineages. Two of them are outgroups ( + +Bryconamericus +species + +), and 17 are + +Knodus + +lineages (species) ( +Fig. 6 +). The new species herein described ( + +K. guajajara + +) was recovered in all three species delimitation methods. Furthermore, some potential undescribed species were detected: + +Knodus + +“Marabá”, + +Knodus + +“Maracaçumé”, + +Knodus +cf. +victoriae + +(Itapecuru River basin) and + +Knodus +cf. +victoriae + +(Balsas River drainage). + +Knodus victoriae + +(upper Parnaiba River basin) was also recovered, as well as two + +K. +cf. +savannensis + +clades. + + + + \ No newline at end of file