Zoosyst. Evol. 96 (2) 2020, 715-722 | DOI 10.3897/zse.96.56247 eee BERLIN A new species from subtropical Brazil and evidence of multiple pelvic fin losses in catfishes of the genus Cambeva (Siluriformes, Trichomycteridae) Wilson J. E. M. Costa’, Caio R. M. Feltrin?, Axel M. Katz! 1 Laboratory of Systematics and Evolution of Teleost Fishes, Institute of Biology, Federal University of Rio de Janeiro, Caixa Postal 68049, CEP 21941-971, Rio de Janeiro, Brazil 2 Av. Municipal, 45, Siderdpolis, CEP 88860-000, Santa Catarina, Brazil http://zoobank.org/DF 6448 D1-9AA4-48A2-912C-C7F7307BSEE2 Corresponding author: Wilson J. E. M. Costa (wcosta@acd.ufrj.br) Academic editor: Nicolas Hubert # Received 7 July 2020 # Accepted 30 September 2020 # Published 11 November 2020 Abstract 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 close- ly 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 7richomycterus, 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. Key Words Mountain biodiversity, osteology, Rio Uruguai basin, Serra Geral, systematics Introduction The presence or not of pelvic fin was considered an import- ant morphological character for generic delimitation in te- leost fishes by 19" century naturalists. Amongst the Tricho- mycterinae, one of the eight subfamilies of the Neotropical catfish Trichomycteridae, the two oldest genera were distin- guished only on the basis of this character. Trichomycterus Valenciennes, 1832 was first diagnosed by its only in- cluded species, 7richomycterus 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 (Humboldt 1805). This classificatory scheme was followed without criticism for a long time. Over 100 years after, two other trichomyc- terids were placed in the genus Eremophilus, E. candidus Miranda-Ribeiro, 1949 and E. camposi Miranda-Ribeiro, 1957, just by lacking pelvic fin (Miranda-Ribeiro 1949, 1957). However, morphological and molecular studies have consistently shown that E. candidus is a member of the genus 7richomycterus (Barbosa and Costa 2003; Katz et al. 2018) and E. camposi belongs to the microcambevine genus Listrura de Pinna, 1988 (de Pinna 1988; Costa et al. 2020a), therefore constituting independent evolutionary events of pelvic fin loss amongst trichomycterids. Copyright Wilson J. E. M. Costa et al. 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. 716 Recent taxonomical studies on the Trichomycterinae, which Is a species-rich clade with over 220 species occur- ring in most river basins between southern Central Amer- ica and Patagonia in southern South America (Katz et al. 2018), have revealed few other species also lacking pelvic fin. Costa and Bockmann (1993) have shown that some specimens of /tuglanis parahybae (Eigenmann, 1918) lack pelvic fin. More recently, five pelvic-less trichomyc- terines have been recorded from the Argentinean Andes (Fernandez and Liotta 2016), as well as two species lack- ing pelvic fin being described from south-eastern and southern Brazil (Ferrer and Malabarba 2011; Ochoa et al. 2017a). These Brazilian species are presently placed in Cambeva Katz, Barbosa, Mattos & Costa, 2018, a genus comprising 25 valid species endemic to south-eastern and southern Brazil (Katz et al. 2018; dos Reis et al. 2019), as well as a single species from north-eastern Argentina (Teran et al. 2017). 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 (Ferrer and Malabarba; 2011 Ochoa et al. 2017b), whereas, in all other 24 remaining species, these structures are well-developed, including an ossified pel- vic girdle and a pelvic fin with five rays. We herein de- scribe a third species of Cambeva lacking pelvic fin and girdle, exhibiting some morphological features indicating that it is not closely related to the other two congeners lacking pelvic fin. Material and methods Morphometric and meristic data were taken following Costa (1992), with modifications proposed by Costa et al. (2020b); measurements are presented as percent of standard length (SL), except for those related to head morphology, which are expressed as percent of head length. Fin-ray counts include all elements; vertebra counts include all vertebrae except those participating in the Weberian apparatus; the compound caudal cen- trum was counted as a single element; counts of verte- brae and procurrent fin rays were made only in cleared and stained specimens; counts of principal-fin rays were made in all available specimens, except juveniles about 25 mm SL or less; counts of jaw teeth were approxi- mate, due to their irregular arrangement, great number and frequent loss, making impossible accurate counts. Specimens were cleared and stained for bone and car- tilage (C&S in lists of specimens) following Taylor and Van Dyke (1985); osteological characters included in the description are those belonging to structures that have informative variability for positioning the new species amongst congeners, including the mesethmoidal region, suspensorium and opercular apparatus and the paruro- hyal. Terminology for bones followed Bockmann et al. (2004), except for: the ‘antorbital’ and the ‘tendon-bone supraorbital’ that are here called ‘lacrimal-antorbital’ and ‘fronto-lacrimal tendon bone’, respectively, following zse.pensoft.net Wilson J. E. M. Costa et al.: New Cambeva from subtropical Brazil Baskin (1973); ‘epibranchial 5 cartilage’, here replaced by ‘ceratobranchial 4 accessory element’ following Car- valho et al. (2013); ‘urohyal’, here substituted by ‘paru- rohyal’ following Arratia and Schultze (1990); ‘pleural rib’, here substituted by ‘rib’ following Britz and Bartsch (2003) that have shown to exist only a single rib type in teleosts. Osteological illustrations were made using a stereomicroscope Zeiss Stemi SV 6 with camera lucida. Cephalic laterosensory system terminology follows Arra- tia and Huaquin (1995), with modifications proposed by Bockmann et al. (2004). Specimens are deposited in the ichthyological collection of the Institute of Biology of the Federal University of Rio de Janeiro, Rio de Janeiro city and in the Centre of Agrarian and Environmental Scienc- es, Federal University of Maranhaéo, Campus Chapadinha (CICCAA). Comparative material is listed in Costa et al. (2020b), with the addition of Cambeva diatropopo- ros UFRJ 6913, 1 (C&S). Within the text, geographical names follow Portuguese terms used in the region, thus avoiding common errors or generalisations when tenta- tively translating them to English, besides making easier their identification in the field. Results Cambeva flavopicta sp. nov. http://zoobank. org/375CE4CA-F 1 ED-4E2D-B379-A F63A | F4E5D8 Figs 1-2, Table 1 Type material. Holotype. UFRJ 12665, 69.2 mm SL; Brazil: Estado de Santa Catarina: Municipio de Campos Novos: Rio Inferno Grande, Rio Canoas drainage, up- per Rio Uruguai basin, 27°21'29"S, 51°01'02"W, about 910 ma.s.l.; CRM Feltrin, 22 May 2019. Paratypes. All from Brazil: Estado de Santa Catarina: Municipio 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. Diagnosis. 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 distin- guished from both C. pascuali and C. tropeiro by hav- ing 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 yel- lowish-brown with dark brown spots irregularly arranged Zoosyst. Evol. 96 (2) 2020, 715-722 Figure 1. Cambeva flavopicta sp. nov., UFRJ 12665, holotype, 69.2 mm SL: A. Left lateral view; B. Dorsal view; C. Ventral view. 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 terminat- ing in a short filament (vs. without a filament) and caudal fin rounded (vs. subtruncate). Description. Morphometric data are presented in Table 1. Body moderately slender, subcylindrical and slightly depressed anteriorly, compressed posteriorly. Greatest body depth at midway between opercle and anal- fin origin. Dorsal profile of head and trunk slightly con- vex, approximately straight on caudal peduncle; ventral profile straight to slightly convex between lower jaw and end of anal-fin base, straight on caudal peduncle. Skin papillae minute. Anus and urogenital papilla in vertical through anterior portion of dorsal-fin base. Head trape- zoidal in dorsal view. Anterior profile of snout convex in dorsal view. Eye small, dorsally positioned in head. Pos- terior nostril located slightly nearer anterior nostril than orbital rim. Tip of maxillary barbel reaching area between Table 1. Morphometric data of Cambeva flavopicta. Measurements Holotype Paratypes (n = 10) Standard length (mm) 69:2 41.2-73.6 Percent of standard length Body depth re 13.2-17.0 Caudal peduncle depth 139 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 Toe 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 115 all 9.4-11.3 Eye diameter 9.6 9.2-12.7 interopercular patch of odontodes and pectoral-fin base; rictal barbel reaching posterior part of interopercular patch of odontodes; tip of nasal barbel reaching area just zse.pensoft.net 718 premaxilla arr A maxilla \---.]mesethmoid lacrimal-antorbital/- autopalatine/" >: . | 0,5 mm fronto-lacrimal tendon bone} B metapterygoid Wilson J. E. M. Costa et al.: New Cambeva from subtropical Brazil hyomandibula opercle quadrate a interopercle Figure 2. 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. anterior to opercular patch of odontodes. Mouth subter- minal. Jaw teeth slightly pointed, few external-most teeth incisiform in larger specimens (above about 65 mm SL); premaxillary teeth about 45, slightly curved, arranged in 3 irregular rows; dentary teeth about 40-45, curved inside mouth, arranged in 3 irregular rows, more concentrated near dentary symphysis. Branchial membrane attached to isthmus only at its anterior point. Branchiostegal rays 9. Dorsal and anal fins subtriangular; total dorsal-fin (i+ II +5); anal-fin origin in vertical between middle and posterior half of dorsal-fin base, approximately between base of 3 and 5" branched dorsal-fin ray. Dorsal-fin or- igin in vertical through centrum of 20" or 21% vertebra; anal-fin origin in vertical through centrum of 23" or 24" vertebra. Pectoral fin subtriangular in dorsal view, poste- rior margin slightly convex, first pectoral-fin ray termi- nating in minute filament, about 10% or less of pecto- ral 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 pro- current rays 20—23 (ixx—xxui + I-II), total ventral procur- rent 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 cor- responding to hypurals | and 2 and parhypural. Latero-sensory system (Fig. 1). Supraorbital sensory canal continuous, connected to posterior section of infra- orbital canal posteriorly. Supraorbital sensory canal with 3 pores: sl, 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 homol- ogous pore. Infra-orbital sensory canal arranged in 2 seg- ments, each with two pores; anterior segment with pore 11, in transverse line through anterior nostril, and pore 13, in transverse line just anterior to posterior nostril; posterior segment with pore 110, adjacent to ventral margin of orbit, and pore i111, posterior to orbit. Postorbital canal with 2 zse.pensoft.net pores: pol, in vertical line above posterior portion of in- teropercular 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. Osteology (Fig. 2). 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-an- torbital 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 medi- al margins slightly concave, with small notch on middle part of medial margin; autopalatine posterolateral pro- cess 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 ar- ticulatory cartilaginous block. Quadrate slender, dorsal process with constricted base and antero-dorsal projec- tion, dorsoposterior margin separated from hyomandib- ula 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; odon- todes pointed, nearly straight, arranged tn irregular trans- verse rows; odontode patch depth about half interoper- cular 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. Paruro- Zoosyst. Evol. 96 (2) 2020, 715-722 92°0'W FAllbe, 48°0'W Figure 3. Map of geographical distribution of Cambeva flavopicta sp. nov. (triangle), and type localities of C. diatropoporos (dia- mond), C. pascuali (pentagon), C. poikilos (star) and C. tropeiro (dot). hyal robust, lateral process latero-posteriorly directed, abruptly narrowing distally, tip sharply pointed; paruro- hyal head well-developed, with distinctive anterolateral paired process; middle foramen small and rounded; pos- terior process well-developed. Colouration in alcohol (Fig. 1). Flank dark brown- ish-grey, with two irregular horizontal rows of small pale yellow grey marks with varied shapes, mostly horizon- tally 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 lon- gitudinal row of rounded, longitudinally-elongated dark brown to black blotches and small pale brown dots. Ven- ter 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 ante- riorly, 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. Distribution and habitat. Cambeva flavopicta 1s only known from the upper Rio Inferno Grande and tributaries, Rio Canoas drainage, upper Rio Uruguai basin, southern Brazil (Fig. 3). The collecting sites were typical mountain rivers draining the Serra Geral, with fast flowing waters (Fig. 4). The species was found close to the riverbank, buried below marginal vegetation. Etymology. From the Latin, the name flavopicta (painted with yellow) refers to the characteristic coloura- tion of this new species, with yellow marks over dark brown ground. Discussion Phylogenetic relationships amongst species of Cambeva are still poorly known. With rare exceptions (Bockmann et al. 2004; Ferrer and Malabarba 2013), osteological data that have been broadly used to diagnose species and es- tablish morphological phylogenetic hypotheses in tricho- mycterid groups (e.g. Baskin 1973), have been broadly omitted even in recent taxonomical studies on Cambeva (e.g. Wosiacki and Garavello 2004; Ferrer and Malabarba 2011; Ochoa et al. 2017b). Similarly, trichomycterid mo- lecular phylogenies have included only a few species of this genus (Ochoa et al. 2017a, 2020; Katz et al. 2018). Therefore, the phylogenetic position of the new species herein described is still uncertain, but some derived os- teological character states, shared by C. flavopicta and other congeners here examined or with available osteo- logical data from literature, highly suggest possible rela- tionships as below discussed. zse.pensoft.net 720 Wilson J. E. M. Costa et al.: New Cambeva from subtropical Brazil Figure 4. Rio Inferno Grande (upper Rio Uruguai basin, Santa Catarina, Brazil), at the exact point where the holotype and paratypes of Cambeva flavopicta sp. nov. were collected. At first glance, the absence of pelvic fin and girdle in all specimens of C. flavopicta could be considered as ev- idence of close relationships with the only two congeners sharing this condition, C. pascuali from the Rio Parana- panema drainage, Rio Parana basin and C. tropeiro from the Lagoa dos Patos system. However, a recent unilocus phylogeny (Donin et al. 2020) indicates that C. pascuali and C. tropeiro are not closely-related species. The for- mer species appears as being closer to C. zonata, a spe- cies from the Rio Ribeira do Iguape basin in south-east- ern Brazil, whereas the latter is sister to C. balios (Ferrer & Malabarba, 2013), a species also endemic to the Lagoa dos Patos system and sharing with C. tropeiro a similar colour pattern consisting of rounded dark brown spots on the flank (Ferrer and Malabarba 2011: fig. 1, 2013: fig. 1). On the other hand, osteological evidence described below supports C. flavopicta as being closely related to species having a well-developed pelvic fin. The long premaxilla recorded for C. flavopicta, with its longest length greater than the distance between the extremities of the mesethmoid cornua (Fig. 2A), is sim- ilar to the long premaxilla shared only by some conge- ners (Ferrer and Malabarba 2013: fig. 2) belonging to a molecularly well-supported clade from rivers of the Lagoa dos Patos system draining the Serra Geral (e.g. Katz et al. 2018; hereafter the C. balios group). Amongst species of the C. balios group, C. flavopicta shares with C. diatropoporos (Ferrer & Malabarba, 2013) and C. poikilos (Ferrer & Malabarba, 2013), a derived com- zse.pensoft.net pact autopalatine with an almost indistinct posterolateral process (Fig. 2A; Ferrer and Malabarba 2013: fig. 2b, c), not present in any congener here examined. Amongst spe- cies herein examined, the presence of a short process on the anteroventral portion of the metapterygoid, just ante- rior to the articulatory cartilaginous block, is shared only by C. flavopicta and C. poikilos, but the metapterygoid of C. diatropoporos, which 1s possibly more closely related to C. flavopicta than to C. poikilos, was not illustrated in previous studies and specimens were not available to this study. According to Ferrer and Malabarba (2013: fig. 2c), C. diatropoporos has a long antorbital, about half the length of the fronto-lacrimal tendon bone length, a derived condition here recorded only for C. flavopicta (Fig. 2A). In addition, C. diatropoporos possesses a co- lour pattern similar to that of C. flavopicta, including flank dark brownish-grey with small pale yellow grey marks with varied shapes (Ferrer and Malabarba 2013: fig. 10), corroborating a hypothesis of close relationships between C. flavopicta and C. diatropoporos. Besides lacking pelvic fin and girdle, C. flavopicta also differs from C. diatropoporos and C. poikilos in having more dorsal procurrent rays in the caudal fin (21—23 vs. 15-19), more ventral procurrent rays in the caudal fin (16-17 vs. 10-15), the posterior margin of the caudal fin convex (vs. straight) and presence of a small filament on the tip of the first pectoral-fin ray (vs. absence); from C. diatropoporos by having the extremity of the parurohyal lateral process sharply pointed (Fig. 2C; vs. broad and rounded, Ferrer Zoosyst. Evol. 96 (2) 2020, 715-722 and Malabarba 2013: fig. 5b) and fewer pectoral-fin rays (6 vs. 7); and from C. poikilos by the presence of the ante- rior section of the infra-orbital canal (vs. absence). 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 sis- ter group of Cambeva (Katz et al., 2018), with all the nine included species having well-developed pelvic fin and girdle (Ferrer and Malabarba 2020). These data sug- gest a stronger tendency to losing pelvic fin in Cambeva than in these closely-related genera, but factors favour- ing this evolutionary event are still unknown. Amongst trichomycterids of the TSVSGM-clade (Costa and Bock- mann 1994: Costa et al. 2020a), parallel loss of pelvic fin and reduction or loss of other fins in the subfamilies Glanapteryginae and Microcambevinae have been re- lated to their interstitial habits (Costa et al. 2020a). On the other hand, conflicting field reports about the habitat of pelvic-less species of Cambeva do not indicate that they share similar habits. Specimens of C. flavopicta were found buried in terrestrial marginal vegetation of a fast flowing stream (see above), whereas specimens of C. pascuali were found under rocks and aquatic vegeta- tion of a muddy bottom area bordered by riparian vegeta- tion (Ochoa et al. 2017b), but no data are available about the specific habitat of C. tropeiro. Therefore, little is still known about the ecology and behaviour in species of the genus Cambeva, presently not allowing explanations in- volving habits for the evolutionary meaning of multiple pelvic fin losses. Acknowledgements We are grateful to Jodo Anténio de Bittencourt Vitto for assistance during field expeditions. Instituto do Meio Am- biente, Santa Catarina and Instituto Chico Mendes de Con- servacao da Biodiversidade provided collecting permits. This work was partially supported by Conselho Nacion- al de Desenvolvimento Cientifico e Tecnolégico (CNPq; grant 307349/2015-2 to WJEMC). The manuscript bene- fited from a detailed review provided by F. Ottoni. References Arratia G, Huaquin L (1995) Morphology of the lateral line system and of the skin of diplomystid and certain primitive loricarioid catfishes and systematic and ecological considerations. Bonner Zoologische Monographien 36: 1-110. Arratia G, Schultze H-P (1990) The urohyal: development and homol- ogy within Osteichthyans. Journal of Morphology 203: 247-282. https://do1.org/10.1002/jmor. 1052030302 72k Barbosa MA, Costa WJEM (2003) Validade, relagdes filogenéticas e re- descrig¢ao de Eremophilus candidus Ribeiro, 1949 (Teleostei, Siluri- formes, Trichomycteridae). Arquivos do Museu Nacional 61: 179-188. Baskin JN (1973) Structure and relationships of the Trichomycte- ridae. Supplementary material of de Pinna (2016). Neotropical Ichthyology 14: $1—S62. Bockmann FA, Casatti L, de Pinna MCC (2004) A new species of trichomycterid catfish from the Rio Paranapanema, southeastern Brazil (Teleostei; Siluriformes), with comments on the phylogeny of the family. Ichthyological Exploration of Freshwaters 15: 225-242. Britz R, Bartsch P (2003) The myth of dorsal ribs in gnathostome ver- tebrates. Proceedings of the Royal Society of London, Biological Sciences Series B 270: S1—S4. https://doi.org/10.1098/rsbl.2003.0035 Carvalho M, Bockmann FA, de Carvalho MR (2013) Homology of the fifth epibranchial and accessory elements of the ceratobranchials among Gnathostomes: insights from the development of Ostariophysans. PLoS ONE 8(4): e62389. https://doi.org/10.1371/journal.pone.0062389 Costa WJEM (1992) Description de huit nouvelles espéces du genre Trichomycterus (Siluriformes: Trichomycteridae), du Brésil orien- tal. Revue frangaise d’Aquariologie et Herpetologie 18: 101-110. Costa WJEM, Bockmann FA (1993) Un nouveau genre néotropical de la famille des Trichomycteridae (Siluriformes: Loricarioidei). Revue francaise d’ Aquariologie et Herpetologie 20: 43-46. Costa WJEM, Bockmann FA (1994) A new genus and species of Sarcoglan- idinae (Siluriformes: Trichomycteridae) from southeastern Brazil, with a re-examination of subfamilial phylogeny. Journal of Natural History 28: 715—730. https://doi.org/10.1080/0022293940077033 1 Costa WJEM, Henschel E, Katz AM (2020a). Multigene phylogeny reveals convergent evolution in small interstitial catfishes from the Amazon and Atlantic forests (Siluriformes: Trichomycteridae). Zoo- logica Scripta 49: 159-173. https://doi.org/10.1111/zsc.12403 Costa WJEM, Katz AM, Mattos JLO, Amorim PF, Mesquita BO, Vilar- do PJ, Barbosa MA (2020b) Historical review and redescription of three poorly known species of the catfish genus Trichomycterus from south-eastern Brazil (Siluriformes: Trichomycteridae). Journal of Natural History 53(47—48): 2905-2928. https://doi.org/10.1080/0 0222933.2020.1752406 de Pinna MCC (1988) A new genus of trichomycterid catfish (Siluroidei, Glanapteryginae), with comments on its phylogenetic relationships. Revue Suisse de Zoologie 95: 113-128. https://doi.org/10.5962/bhl. part.79642 Donin LM, Ferrer J, Carvalho TP (2020) Taxonomical study of Tricho- mycterus (Siluriformes: Trichomycteridae) from the Ribeira de Iguape River basin reveals a new species recorded in the early 20" century. Journal of Fish Biology 96(4): 886-904. https://doi. org/10.1111/jfb.14278 dos Reis RB, Frota A, Fabrin TMC, da Graga WJ (2019) A new species of Cambeva (Siluriformes, Trichomycteridae) from the Rio Ivai ba- sin, Upper Rio Parana basin, Parana State, Brazil. Journal of Fish Biology 96(2): 350-363. https://doi.org/10.1111/jfb.14204 Fernandez L, Liotta J (2016) Silvinichthys pachonensis, a new catfish from high altitude, with a key to the species of the genus (Siluri- formes: Trichomycteridae). Ichthyological Exploration of Freshwa- ters 27: 375-383. Ferrer J, Malabarba LR (2011) A new Trichomycterus lacking pelvic fins and pelvic girdle with a very restricted range in Southern Brazil (Siluriformes: Trichomycteridae). Zootaxa 2912: 59-67. https://doi. org/10.11646/zootaxa.2912.1.5 zse.pensoft.net 722 Ferrer J, Malabarba LR (2013) Taxonomic review of the genus 7richo- mycterus Valenciennes (Siluriformes: Trichomycteridae) from the laguna dos Patos system, Southern Brazil. Neotropical Ichthyology 11: 217-246. https://doi.org/10.1590/S1679-62252013000200001 Ferrer J, Malabarba LR (2020) Systematic revision of the Neotropical cat- fish genus Scleronema (Siluriformes: Trichomycteridae), with descrip- tions of six new species from Pampa grasslands. Neotropical Ichthy- ology 18(2): e190081. https://doi.org/10.1590/1982-0224-2019-008 1 Humboldt A von (1805) Mémoire sur |’Eremophilus et Astroblepus, deux nouveaux genres de |’ordre des apodes. In: Voyage de Hum- boldt et Bonpland, Deuxieme partie. Observations de Zoologie et d’ Anatomie compare, 17—20. [67 pls] Katz AM, Barbosa MA, Mattos JLO, Costa WJEM (2018) Multigene analysis of the catfish genus 7richomycterus and description of a new South American trichomycterine genus (Siluriformes, Tricho- mycteridae). Zoosystematics and Evolution 94: 557-566. https:// doi.org/10.3897/zse.94.29872 Miranda Ribeiro P (1949) Notas para 0 estudo dos Pygidiidae Brasile- iros. (Pisces — Pygidiidae — Pygidiinae) III. Boletim do Museu Na- cional, Zoologia, Nova Série 88: 1-3. [2 pls] Miranda Ribeiro P (1957) Notas para o estudo dos Pygidiidae Brasile- iros (Pisces — Pygidiidae). VI. Publicagdes Avulsas do Departamen- to de Zoologia, Sao Paulo 13(5): 71-73. Ochoa LE, Roxo FF, DoNascimiento C, Sabaj MH, Datovo A, Alfa- ro M, Oliveira C (2017a) Multilocus analysis of the catfish family Trichomycteridae (Teleostei: Ostariophysi: Siluriformes) support- ing a monophyletic Trichomycterinae. Molecular Phylogenetics and Evolution 115: 71-81. https://doi.org/10.1016/j.ympev.2017.07.007 zse.pensoft.net Wilson J. E. M. Costa et al.: New Cambeva from subtropical Brazil Ochoa LE, Silva GSC, Costa e Silva GJ, Oliveira C, Datovo A (2017b) New species of Trichomycterus (Siluriformes: Trichomycteridae) lacking pelvic fins from Paranapanema basin, southeastern Brazil. Zootaxa 4319: 550-560. https://doi.org/10.11646/zootaxa.4319.3.7 Ochoa LE, Datovo A, DoNascimiento C, Roxo FF, Sabaj MH, Chang J, Melo BEF, Silva Gabriel SC, Foresti F, Alfaro M, Oliveira C (2020) Phylogenomic analysis of trichomycterid catfishes (Teleostei: Silu- riformes) inferred from ultraconserved elements. Scientific Reports 10: e2697. https://doi.org/10.1038/s41598-020-595 19-w Taylor WR, Van Dyke GC (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone and carti- lage study. Cybium 9: 107-109. http://sfi.mnhn.fr/cybium/numer- 0s/1985/92/01-Taylor%5b92%5d107-119.pdf Teran GE, Ferrer J, Benitez M, Alonso F, Aguilera G, Mirande JM (2017) Living in the waterfalls: A new species of 7richomycterus (Siluriformes: Trichomycteridae) from Tabay stream, Misiones, Argentina. PLoS ONE 12(6): e0179594. https://doi.org/10.1371/journal.pone.0179594 Valenciennes A (1832) Nouvelles observations sur le Capitan de Bogo- ta, Eremophilus mutisti. In. Humboldt A, Bonpland A (1799-1803) Recueil d’ observations de Zoologie et d’ Anatomie Comparée, faites dans Il’Ocean Atlantique, dans |’interieur du Nouveau Continent et dans la Mer du Sud pendant les années 1799, 1800, 1801, 1802 et 1803, deuxieme volume. Observations de Zoologie et d’Anatomie comparée, 341-348. Wosiacki WB, Garavello JC (2004) Five new species of Trichomyc- terus from the Iguacu (rio Parana Basin), southern Brazil (Siluri- formes: Trichomycteridae). Ichthyological Exploration of Freshwa- ters 15: 1-16.