Research Article |
Corresponding author: Wilson J. E. M. Costa ( wcosta@acd.ufrj.br ) Academic editor: Nicolas Hubert
© 2020 Wilson J. E. M. Costa, Caio R. M. Feltrin, Axel M. Katz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Costa WJEM, Feltrin CRM, Katz AM (2020) A new species from subtropical Brazil and evidence of multiple pelvic fin losses in catfishes of the genus Cambeva (Siluriformes, Trichomycteridae). Zoosystematics and Evolution 96(2): 715-722. https://doi.org/10.3897/zse.96.56247
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A third pelvic-less species of Cambeva from river basins draining the Geral mountain range in southern Brazil is described. It is distinguished from other congeners lacking pelvic fin and girdle, C. pascuali and C. tropeiro, by having six pectoral-fin rays, 20–23 dorsal procurrent caudal-fin rays, 15–20 opercular and 25–30 interopercular odontodes and a different colour pattern consisting of flank dark brownish-grey with two irregular horizontal rows of small pale yellow grey marks. Whereas available molecular evidence indicates that C. pascuali is more closely related to C. zonata, a species with well-developed pelvic fin, and C. tropeiro is more closely related to C. balios, another species also with well-developed pelvic fin; osteological data strongly suggest that the new species herein described is more closely related to C. diatropoporos than to other congeners. Therefore, this study indicates that the pelvic fin and pelvic-fin support have been lost independently in each of these three species of Cambeva, which corresponds to 11% of all describe species. This result highly contrasts with the closely-related trichomycterine genera Trichomycterus, in which only one in 50 species lost pelvic fin and girdle (0.2%) and Scleronema with all the nine included species having well-developed pelvic fin. These data suggest a stronger tendency to losing pelvic fin in Cambeva, but factors favouring this evolutionary event are still unknown.
Mountain biodiversity, osteology, Rio Uruguai basin, Serra Geral, systematics
The presence or not of pelvic fin was considered an important morphological character for generic delimitation in teleost fishes by 19th century naturalists. Amongst the Trichomycterinae, one of the eight subfamilies of the Neotropical catfish Trichomycteridae, the two oldest genera were distinguished only on the basis of this character. Trichomycterus Valenciennes, 1832 was first diagnosed by its only included species, Trichomycterus nigricans Valenciennes, 1832, having pelvic fin (Valenciennes in Humboldt and Bonpland 1832), thus differing from the only other then-known trichomycterine Eremophilus mutisii Humboldt, 1805, in which pelvic fin is absent (
Recent taxonomical studies on the Trichomycterinae, which is a species-rich clade with over 220 species occurring in most river basins between southern Central America and Patagonia in southern South America (
Amongst the 26 species of Cambeva, only C. pascuali (Ochoa, Silva, Costa e Silva, Oliveira & Datovo, 2017) and C. tropeiro (Ferrer & Malabarba, 2011) lack pelvic fin and girdle (
Morphometric and meristic data were taken following
Holotype. UFRJ 12665, 69.2 mm SL; Brazil: Estado de Santa Catarina: Município de Campos Novos: Rio Inferno Grande, Rio Canoas drainage, upper Rio Uruguai basin, 27°21'29"S, 51°01'02"W, about 910 m a.s.l.; CRM Feltrin, 22 May 2019.
Paratypes. All from Brazil: Estado de Santa Catarina: Município de Campos Novos: Rio Canoas drainage, upper Rio Uruguai basin: UFRJ 12234, 61, 31.5–73.6 mm SL; UFRJ 12664, 5, 38.2–47.9 mm SL (C&S); CICCAA 04776, 10, 34.5–53.4 mm SL; UFRJ 12235, 8, 27.7–61.7 mm SL; collected with holotype. UFRJ 12662, 10, 35.8–62.2 mm SL; UFRJ 12663, 3 (C&S), 38.7–43.4 mm SL; 27°22'05"S, 51°00'34"W; CRM Feltrin, 12 January 2019.
Cambeva flavopicta is distinguished from all other congeners, except C. pascuali and C. tropeiro, by the absence of pelvic fin and pelvic girdle (vs. pelvic fin and girdle present and well-developed). It is distinguished from both C. pascuali and C. tropeiro by having 6 pectoral-fin rays (vs. 5 in C. pascuali and 7 in C. tropeiro), more dorsal procurrent caudal-fin rays (20–23 vs. 17–18 in C. pascuali and 14–15 in C. tropeiro), more opercular odontodes (15–20 vs. 10–12 in C. pascuali and 12–14 in C. tropeiro), more interopercular odontodes (25–30 vs. 11–12 in C. pascuali and 18–24 in C. tropeiro) and colour pattern consisting of flank dark brownish-grey with two irregular horizontal rows of small pale yellow grey marks (vs. flank pale yellow with dark brown stripes or horizontal rows of spots in C. pascuali and flank yellowish-brown with dark brown spots irregularly arranged in C. tropeiro). Cambeva flavopicta also differs from C. pascuali by the presence of the anterior section of the infra-orbital canal (vs. absence), more vertebrae (38–39 vs. 37) and more branchiostegal rays (9 vs. 7); and from C. tropeiro by having more ventral procurrent caudal-fin rays (16–17 vs. 10–11), the first pectoral-fin ray terminating in a short filament (vs. without a filament) and caudal fin rounded (vs. subtruncate).
Morphometric data are presented in Table
Measurements | Holotype | Paratypes (n = 10) |
---|---|---|
Standard length (mm) | 69.2 | 41.2–73.6 |
Percent of standard length | ||
Body depth | 12.9 | 13.2–17.0 |
Caudal peduncle depth | 11.9 | 12.1–13.6 |
Body width | 10.4 | 9.8–11.3 |
Caudal peduncle width | 4.0 | 2.4–3.5 |
Pre-dorsal length | 64.5 | 62.9–66.7 |
Dorsal-fin base length | 12.6 | 10.5–12.4 |
Anal-fin base length | 9.4 | 8.5–10.3 |
Caudal-fin length | 15.7 | 14.2–18.4 |
Pectoral-fin length | 11.4 | 10.4–12.7 |
Head length | 19.0 | 18.9–22.6 |
Percent of head length | ||
Head depth | 48.5 | 45.5–56.0 |
Head width | 86.6 | 77.2–86.5 |
Snout length | 44.7 | 39.1–42.8 |
Interorbital length | 25.7 | 21.1–26.1 |
Preorbital length | 13.1 | 9.4–11.3 |
Eye diameter | 9.6 | 9.2–12.7 |
Dorsal and anal fins subtriangular; total dorsal-fin rays 11–12 (ii–iii + II–III + 6–7), total anal-fin rays 9 (ii + II + 5); anal-fin origin in vertical between middle and posterior half of dorsal-fin base, approximately between base of 3rd and 5th branched dorsal-fin ray. Dorsal-fin origin in vertical through centrum of 20th or 21st vertebra; anal-fin origin in vertical through centrum of 23rd or 24th vertebra. Pectoral fin subtriangular in dorsal view, posterior margin slightly convex, first pectoral-fin ray terminating in minute filament, about 10% or less of pectoral fin length without filament; total pectoral-fin rays 6 (I + 5). Pelvic fin and girdle absent. Posterior margin of caudal fin convex, upper and lower margins straight; total principal caudal-fin rays 13 (I + 11 + I), total dorsal procurrent rays 20–23 (ixx–xxii + I–II), total ventral procurrent rays 16–17 (xv–xvi + I). Vertebrae 38–39. Ribs 13 or 14. Two dorsal hypural plates, corresponding to hypurals 4 + 5 and 3, respectively; single ventral hypural plate corresponding to hypurals 1 and 2 and parhypural.
Supraorbital sensory canal continuous, connected to posterior section of infraorbital canal posteriorly. Supraorbital sensory canal with 3 pores: s1, adjacent to medial margin of anterior nostril; s3, adjacent and just posterior to medial margin of posterior nostril; and s6, in transverse line through posterior half of orbit; pore s6 slightly nearer orbit than its paired homologous pore. Infra-orbital sensory canal arranged in 2 segments, each with two pores; anterior segment with pore i1, in transverse line through anterior nostril, and pore i3, in transverse line just anterior to posterior nostril; posterior segment with pore i10, adjacent to ventral margin of orbit, and pore i11, posterior to orbit. Postorbital canal with 2 pores: po1, in vertical line above posterior portion of interopercular patch of odontodes, and po2, in vertical line above posterior portion of opercular patch of odontodes. Lateral line of body short, with 2 pores, posterior-most pore in vertical just posterior to pectoral-fin base.
Mesethmoid robust, its anterior margin nearly straight; mesethmoid cornu subtriangular in dorsal view, basal portion wide, abruptly narrowing distally, extremity pointed. Lacrimal-antorbital thin, drop-shaped; fronto-lacrimal tendon bone slender and with lateral process, sometimes membranous expansion, its length about one and half times or twice lacrimal-antorbital length. Premaxilla sub-rectangular in dorsal view, long, longer than distance between extremities of mesethmoid cornua. Maxilla boomerang-shaped, slender, about 80% maxilla length, slightly curved. Autopalatine subrectangular in dorsal view, compact, lateral and medial margins slightly concave, with small notch on middle part of medial margin; autopalatine posterolateral process almost indistinct. Metapterygoid thin, subtriangular, large, its largest length about equal horizontal length of quadrate excluding dorsal process; anteroventral portion of metapterygoid with short process just anterior to articulatory cartilaginous block. Quadrate slender, dorsal process with constricted base and antero-dorsal projection, dorsoposterior margin separated from hyomandibula outgrowth by interspace. Hyomandibula long, with well-developed anterior outgrowth; middle portion of dorsal margin of hyomandibula outgrowth with shallow concavity. Opercle slender, with 15–20 odontodes; odontodes pointed, nearly straight, arranged in irregular transverse rows; odontode patch depth about half interopercular odontode patch length; dorsal process of opercle short and pointed; ventral process of opercle moderate, about half opercle length. Interopercle moderate, about two thirds hyomandibula length, with 25–30 odontodes; odontodes pointed, arranged in irregular longitudinal rows; anterior margin of interopercle truncate; dorsal interopercular process with deep anterior concavity. Preopercle compact, with minute ventral flap. Parurohyal robust, lateral process latero-posteriorly directed, abruptly narrowing distally, tip sharply pointed; parurohyal head well-developed, with distinctive anterolateral paired process; middle foramen small and rounded; posterior process well-developed.
Osteological structures of Cambeva flavopicta sp. nov., UFRJ 12664, paratype, 47.9 mm SL: A. Mesethmoidal region and adjacent structures, left and middle portions, dorsal view; B. Left suspensorium and opercular series, lateral view; C. Parurohyal, ventral view. Arrow indicates the ventral process on the metapterygoid. Larger stippling represents cartilaginous areas.
Flank dark brownish-grey, with two irregular horizontal rows of small pale yellow grey marks with varied shapes, mostly horizontally orientated, often horizontally coalesced on flank longitudinal midline, sometimes vertically coalesced on caudal peduncle; in some specimens, pale yellow marks restricted to small spots. Dorsum pale yellow with longitudinal row of rounded, longitudinally-elongated dark brown to black blotches and small pale brown dots. Venter yellowish-white, with brown chromatophores slightly concentrated on area just anterior to urogenital region and close to pectoral-fin base. Side and dorsal surface of head light yellowish-grey with irregularly-shaped dark brown to black spots; ventral surface of head yellowish-white, with brown chromatophores slightly concentrated anteriorly, to brown on lower jaw. Barbels brown. Opercle dark brown to black, interopercle yellowish-white with minute brown dots. Fins grey with small dark brown to black spots.
Cambeva flavopicta is only known from the upper Rio Inferno Grande and tributaries, Rio Canoas drainage, upper Rio Uruguai basin, southern Brazil (Fig.
From the Latin, the name flavopicta (painted with yellow) refers to the characteristic colouration of this new species, with yellow marks over dark brown ground.
Phylogenetic relationships amongst species of Cambeva are still poorly known. With rare exceptions (
At first glance, the absence of pelvic fin and girdle in all specimens of C. flavopicta could be considered as evidence of close relationships with the only two congeners sharing this condition, C. pascuali from the Rio Paranapanema drainage, Rio Paraná basin and C. tropeiro from the Lagoa dos Patos system. However, a recent unilocus phylogeny (
The long premaxilla recorded for C. flavopicta, with its longest length greater than the distance between the extremities of the mesethmoid cornua (Fig.
As discussed above, this study indicates that the pelvic fin and pelvic-fin support have been lost independently in three lineages of the genus Cambeva, corresponding to three species in a total of 27 (11.1%). This result highly contrasts with the closely-related trichomycterine genus Trichomycterus, in which only one species, T. candidus, does not have pelvic fin and girdle, amongst a total of 50 valid species (0.2%), as well as Scleronema, the sister group of Cambeva (
We are grateful to João Antônio de Bittencourt Vitto for assistance during field expeditions. Instituto do Meio Ambiente, Santa Catarina and Instituto Chico Mendes de Conservação da Biodiversidade provided collecting permits. This work was partially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; grant 307349/2015-2 to WJEMC). The manuscript benefited from a detailed review provided by F. Ottoni.