Zoosyst. Evol. 97 (1) 2021, 147-159 | DOI 10.3897/zse.97.61006 Arte BERLIN Filling distribution gaps: Two new species of the catfish genus Cambeva from southern Brazilian Atlantic Forest (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/FCEDODBF-90AA-4299-B3C3-8B208971EBA6 Corresponding author: Wilson J. E. M. Costa (wcosta@acd.ufrj.br) Academic editor: Nicolas Hubert Received 21 November 2020 # Accepted 25 January 2021 # Published 2 March 2021 Abstract The fauna and flora of the Brazilian Atlantic Forest have been intensively inventoried since the 19" century, but some components of this rich biota are still poorly known, and some areas have been poorly sampled. Recent studies on a rich collection of mountain catfishes of the genus Cambeva have revealed a high diversity of species still undescribed in the region. Here we provide formal descriptions for two of these species, found in areas inserted in a broad gap of the presently known genus distribution. The first one is endemic to small coastal river basins of Santa Catarina state, southern Brazil; it is tentatively placed in an intrageneric clade, also including C. castroi, C. davisi, C. guareiensis and C. zonata, by all sharing the presence of a flat small process on the dorsal margin of the quadrate, laterally overlapping metapterygoid and situated just posterior to the syncondrial joint between the metapterygoid and the quadrate. Phylogenetic relationships of the second new species, endemic to the Rio Itajai-Mirim basin, are still obscure, but it shares a derived morphology of the mesethmoid with some species of the C. balios group. Although species of Cambeva have little external morphological variation when compared to other trichomycterine groups, the present study once more shows the importance of recording and using osteological characters to diagnose externally similar trichomycterine species. Key Words Comparative morphology, mountain biodiversity, osteology, systematics Introduction The fauna and flora of the Brazilian Atlantic Forest, con- sidered among the most important biodiversity hotspots in the world (Myers et al. 2000), have been intensively inventoried and described since the first half of the 19" century, when numerous European naturalists visited the region for the first time (e.g. Papavero 1971; Costa et al. 2020a). However, after about 200 years of regular stud- ies, some components of this rich biota are still poorly known. Among freshwater fishes, represented in streams of this region by over 270 species (e.g. Abilhoa et al. 2011), studies based on expeditions conducted in the last 30 years directed to sample fish in specialised biotopes have revealed a rich still unknown species diversity, first described in recent years (e.g. Costa 2009; Costa et al. 2020b). Among these specialised freshwater biotopes are mountain streams, with numerous endemic fishes be- ing first sampled and described in recent years, mainly belonging to the Trichomycterinae (Costa et al. 2020c, 2020d; Donin et al. 2020; dos Reis et al. 2020; Vilardo et al. 2020), a catfish subfamily (Siluriformes: Trichomyc- teridae) occurring between southern Central America and Patagonia, in southern South America (Katz et al. 2018). Trichomycterines are typically found in fast-flowing streams and each species is usually geographically re- 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. 148 stricted to small areas (e.g. Ferrer and Malabarba 2013; Costa et al. 2020a, c; Vilardo et al. 2020). In southern Brazil, the most species-rich trichomycterine genus is Cambeva Katz, Barbosa, Mattos & Costa, 2018, present- ly with 27 valid species (see below), among which only four were described before 1992 (Katz et al. 2018). The number of known species of Cambeva has been quickly increased in recent years as a result of field inventories in previously under-sampled areas and the increasingly interest in trichomycterid taxonomy after Costa (1992) and de Pinna (1992), but some areas remain unsampled. Species of Cambeva are present in both large river basins such as the Parana, Sao Francisco and Uruguai basins, and in small coastal river drainages. The greatest concen- tration of species occurs in the Rio Parana basin, with a total of 16 species (Katz and Costa 2020), including C. papillifera (Wosiacki & Garavello, 2005), new com- bination, which exhibits the typical combination of ex- ternal morphological features of Cambeva (i.e. pectoral fin lacking a filament at the tip of the first ray and hy- pertrophied cheek muscles), but equivocally omitted in Katz et al. (2018). Among the species not occurring in the Rio Parana basin, two occur in the Sao Francisco basin (Costa 1992), five in the Lagoa dos Patos system (Fer- rer and Malabarba 2011, 2013), two in the Rio Uruguai basin (Datovo et al. 2012; Costa et al. 2020d), and three in the Rio Ribeira de Iguape and smaller coastal basins (Eigenmann 1918; Bizerril 1994; Wosiacki and Oyakawa 2005). However, no species was described from the wide area, about 250 km long, between the coastal river basins of northern Santa Catarina state (Bizerril 1994; Katz and Barbosa 2014) and the southern end of the Atlantic For- est, in the Lagoa dos Patos system. In the past two decades, the river basins of the south- ern portion of the Atlantic Forest have been inventoried by members of the Laboratory of Systematics and Evolu- tion of Teleost Fishes team (UFRJ), and more recently ex- haustively sampled by one of us (CRM), revealing a great species diversity comparable to those already recorded for neighbouring areas. The objective of this paper is to provide formal descriptions for two of the new species collected in this area. Material and methods Morphometric and meristic data were taken following Costa (1992), with modifications proposed by Costa et al. (2020a); 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 centrum was counted as a single element; counts of vertebrae 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 30 mm SL or zse.pensoft.net Wilson J.E.M. Costa et al.: Two new species of Cambeva less; counts of jaw teeth were approximate, due to their irregular arrangement, great number and frequent loss, making impossible accurate counts. Specimens were cleared and stained for bone and cartilage (C&S in lists of specimens) following Taylor and Van Dyke (1985). Oste- ological characters included in descriptions are those be- longing to structures with informative variability for diag- nosing species of Cambeva, including the mesethmoidal region, suspensorium and opercular apparatus, and hyoid skeleton. Terminology for bones followed Bockmann et al. (2004), except for “‘tendon-bone supraorbital’, here called ‘sesamoid supraorbital’ following Adriaens et al. (2010); ‘urohyal’, here called ‘parurohyal’ following Ar- ratia and Schultze (1990); and ‘pleural rib’, here called simply ‘rib’ following the morphological study by Britz and Bartsch (2003) supporting to exist 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 Maranhao, Campus Chapadinha (CICCAA). Comparative material is listed in Costa et al. (2020a). Throughout the text, geographical names follow Portuguese terms used in the region, thus avoiding com- mon errors or generalizations when tentatively translating them to English, besides making easier their identifica- tion in the field. Results Cambeva barbosae sp. nov. http://zoobank.org/AOC5B36A-9A31-4075-B5A 1-A3D9C6548105 Figs 1, 2, 3A-E, 4A-C, Table 1 Holotype. UFRJ 10000, 67.6 mm SL; Brazil: Santa Ca- tarina state: Aguas Mornas municipality: Rio Cubatao do Sul basin, 27°39'51"S, 48°51'26"W, about 130 m asl; A.M. Katz, F.R. Pereira & M.A. Barbosa, 31 May 2013. Paratypes. All from Brazil: Santa Catarina state. Rio Cu- batéo do Sul: UFRJ 9503, 10, 44.1-71.6 mm SL; UFRJ 9848, 5, 34.1-41.7 mm SL (C&S); collected with holo- type. Rio Maruim basin: — UFRJ 9505, 13.3-73.1 mm SL; 27°40'39"S, 48°50'53"W, about 90 m asl; same col- lectors and date as holotype. —- UFRJ 12629, 6, 38.7— 62.0 mm SL; stream tributary to Rio Forquilhas, Colénia Santana, Sao José municipality, 27°32'44"S, 48°42'21" W, about 40 m asl; C.R.M. Feltrin, 15 Nov. 2019. — UFRJ 6924, 10, 31.9-57.6 mm SL; small stream tributary to up- per Rio Forquilhas, village of Alto Forquilhas, Sao José municipality, 27°32'44"S, 48°42'21"W, about 40 m asl; C.R.M. Feltrin, 17 Jun. 2020. Rio Cubatao do Sul basin: UFRJ 6925, 25, 24.8-62.1 mm SL; unnamed stream, Zoosyst. Evol. 97 (1) 2021, 147-159 149 Figure 1. Cambeva barbosae sp. nov., UFRJ 10000, holotype, 67.6 mm SL: A. left lateral view; B. dorsal view; C. ventral view. Aguas Mornas municipality, 27°43'16"S, 48°53'23"W, about 190 m asl; C.R.M. Feltrin, 13 Jun. 2020. Rio Biguacu basin, Biguacu municipality: UFRJ 11717, 11, 31.6-67.9 mm SL; UFRJ 6921, 3 (C&S), 51.1-58.9 mm SL; Cachoeira Graciosa, 27°26'37"S, 48°40'59"W, about 60 m asl; C.R.M. Feltrin, Aug. 2017. — UFRJ 11872, 2, 54.1-54.9 mm SL; same locality and collector as UFRJ 11717, 27 Nov. 2018. —CICCAA 02617, 2, 53.5—62.3 mm SL; Riacho Canudos, 27°24'30"S, 48°45'17"W, about 70 m asl; C.R.M. Feltrin, 4 Dec. 2017. Additional material (non-types). Rio Biguac¢u basin, Bigua¢u municipality: UFRJ 12383, 4; UFRJ 12385, 11; Riacho Canudos, 27°25'20"S, 48°45'13"W, about 30 m asl; B. Mesquita & P.F. Amorim, 16 Aug. 2019. Floria- nopolis municipality: UFRJ 10603, 10; UFRJ 10669, 2 (C&S); Corrego Grande, Ilha de Santa Catarina, 27°36'10"S, 48°30'12"W, about 15 m asl; A.M. Katz, F. Pereira & PF. Amorim, 11 Jun. 2015. Diagnosis. Cambeva barbosae differs from all conge- ners, except C. castroi (de Pinna, 1992), C. concolor (Costa, 1992), C. crassicaudata (Wosiacki & de Pinna 2008), C. diabola (Bockmann, Casatti & de Pinna, 2004), C. guaraquessaba (Wosiacki, 2005), C. igobi (Wosia- cki & de Pinna, 2008), C. iheringi (Eigenmann, 1917), C. tupinamba (Wosiacki & Oyakawa, 2005), C. variegata (Costa, 1992), C. ytororo Teran, Ferrer, Benitez, Alonso, Aguilera & Mirande, 2017, and C. zonata (Eigenmann, 1918) by having eight pectoral-fin rays (vs. five to seven in all other species). Cambeva barbosae differs from all these species by the following combination of diagnos- tic features: nine principal dorsal-fin rays (vs. 12-13 in C. concolor, C. iheringi and C. variegata), 19-23 dorsal procurrent caudal-fin rays (vs. 15-16 in C. guarequess- aba and C. tupinamba;, 24—29 in C. crassicaudata and C. igobi; and 31-35 in C. ytororo), 8-12 ventral pro- current caudal-fin rays (vs. 17-19 in C. crassicaudata), 36-38 vertebrae (vs. 34-35 in C. concolor, C. iheringi and C. variegata); all jaw teeth incisiform (vs. conical in C. castroi and C. diabola, anterior teeth sub-incisiform, posterior teeth conical in C. zonata), and absence of a dark grey to black bar on the posterior portion of the cau- dal fin, contrasting with a white to pale yellow zone on the anterior portion of the fin (vs. presence in C. castroi and C. diabola). Also distinguished from C. concolor, C. crassicaudata, C. igobi, and C. variegata by having a distinctive process on the dorsal margin of the quadrate, zse.pensoft.net 150 Wilson J.E.M. Costa et al.: Two new species of Cambeva pot po2 j 4 il2 Figure 2. Head of Cambeva barbosae sp. nov., UFRJ 10000, holotype, 67.6 mm SL: A, left lateral view B. dorsal view; C, ventral view. Pores of the cephalic latero-sensory system are indicated in A and B. just posterior to the cartilage block joining quadrate and metapterygoid (Fig. 4B; vs. absence), and from C. cas- troi, C. concolor, C. crassicaudata, C. diabola, C. igobi, C. variegata (Costa, 1992), and C. zonata by the presence of a distinctive, anteriorly directed process on the lateral margin of the lateral ethmoid (Fig. 4A; vs. absence) and a deep notch between the lateral shell on the opercular zse.pensoft.net articular facet for the hyomandibula and the opercular ar- ticular facet for the preopercle in larger specimens (above about 50 mm SL, Fig. 4B; vs. absence). Description. Morphometric data appear in Table 1. Body moderately slender, subcylindrical and slightly depressed anteriorly, compressed posteriorly. Greatest Zoosyst. Evol. 97 (1) 2021, 147-159 151 Figure 3. Live specimens (not preserved) of: A-E. Cambeva barbosae sp. nov.: A. From the type locality; B—D. From Corrego Grande, Ilha de Santa Catarina; E. From Rio Bigua¢u basin; F. Cambeva botuvera sp. nov., from the type locality. Table 1. Morphometric data of Cambeva barbosae sp. nov. Holotype Paratypes (n = 12) Standard length (mm) 67.6 44.3-71.6 Percent of standard length Body depth 14.0 13.9-18.7 Caudal peduncle depth 12.0 11.3-13.0 Body width 10.2 10.2-14.1 Caudal peduncle width 3.1 2.9-4.7 Pre-dorsal length 60.2 61.5-66.6 Pre-pelvic length 54.5 54.9-61.6 Dorsal-fin base length 11.7 10.8-13.8 Anal-fin base length a7, 7.7-9.1 Caudal-fin length 17.3 14.4-18.5 Pectoral-fin length 14.9 12.1-15.1 Pelvic-fin length 10.3 8.7-11.2 Head length 20.4 19.7-22.8 Percent of head length Head depth 45.4 46.6-54.4 Head width 81.3 79.0-89.9 Snout length 40.1 41.2-45.5 Interorbital length 20.7 20.5-25.9 Preorbital length 15.3 13.6-16.7 Eye diameter 12.8 10.8-13.8 body depth in area just anterior to pelvic-fin base. Dorsal profile of head and trunk slightly convex, 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 trapezoidal in dorsal view. Anterior profile of snout convex in dorsal view. Eye small, dorsally positioned in head. Posterior nostril located nearer anterior nostril than orbital rim. Tip of maxillary and rictal barbels reaching posterior half of interopercular patch of odontodes; tip of nasal barbel surpassing posterior margin of orbit, reaching transverse line through middle of interopercular patch of odontodes. Mouth subterminal. Jaw teeth incisiform and slightly curved, 40-52 in premaxilla, 42-45 in dentary, arranged in three or four irregular rows. Branchial membrane attached to isthmus only at its anterior point. Branchiostegal rays 8 or 9. Dorsal and anal fins subtriangular; total dorsal-fin rays 11 Gi+ II + 7), total anal-fin rays 9 (11 + II + 5); anal-fin origin in vertical through posterior portion of dorsal-fin base, approximately at base of 5" branched dorsal-fin ray. Dorsal-fin origin in vertical between centrum of 19" and 20" vertebrae; anal-fin origin in vertical between centrum of 23" and 24" vertebra. Pectoral fin subtriangular in dorsal view, posterior margin slightly convex, tip of first pectoral-fin ray not forming filament; total pectoral-fin rays 8 (1+ 7). Pelvic fin subtruncate, its extremity in ver- tical through anterior portion of dorsal-fin base; pelvic-fin bases medially in close proximity; total pelvic-fin rays 5 (I + 4). Caudal fin truncate, postero-dorsal and poste- ro-ventral extremities rounded; total principal caudal-fin rays 13 (1+ 11 + J), total dorsal procurrent rays 19-23 (xvili-xxii + I), total ventral procurrent rays 10—12 (ix—xi + T). Vertebrae 36-38. Ribs 12—13. Two dorsal hypural plates, corresponding to hypurals 4 + 5 and 3, respective- ly; single ventral hypural plate corresponding to hypurals 1 and 2 and parhypural. Laterosensory system (Fig. 2A—B). Supraorbital sen- sory canal continuous, connected to posterior section of infraorbital 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 poste- rior half of orbit; pore s6 nearer orbit than its paired ho- mologous pore. Anterior segment of infraorbital sensory canal absent; posterior segment with two pores, pore 110, adjacent to ventral margin of orbit, and pore 111, posterior to orbit. Postorbital canal with 2 pores: pol, in vertical line above posterior portion of interopercular patch of odontodes, and po2, in vertical line above posterior por- tion of opercular patch of odontodes. Lateral line of body short, with 2 pores, posterior-most pore in vertical just posterior to pectoral-fin base. zse.pensoft.net A premaxilla B ete eile metapterygoid quadrate ee f | mesethmoid 2 ine preopercle lep interopercle Hy : sesamoid supraorbital f 4 1mm m premaxilla E maxilla rah cp SEL EE, el * metapterygoid ie antorbital ~ \- autopalatine 1mm hyomandibula : IRA ed eo Le a SNAN ete eee EN interopercle \\\ WEES: aria Wilson J.E.M. Costa et al.: Two new species of Cambeva ventral hypohyal anterior ceratohyal aa dees > branchiostegal rays ventral hypohyal branchiostegal rays Figure 4. Osteological structures of: A-C. Cambeva barbosae sp. nov.; D-F. Cambeva botuvera sp. nov.: A, D. Mesethmoidal region and adjacent structures, left and middle portions, dorsal view; B, E. Left suspensorium and opercular series, lateral view; C, F. Hyoid arch, ventral view. Abbreviations of structures indicated by arrows are: lep, lateral ethmoid lateral process; mlf, mesethmoidal lateral flap; oig, opercular interarticular gap; ppp parurohyal posterior process. Larger stippling represents cartilaginous areas. Mesethmoidal region (Fig. 4A). Mesethmoid robust, its anterior margin slightly concave; mesethmoid cornu narrow, extremity rounded. Lateral ethmoid connected to autopalatine by weak articular facet, which in specimens larger than about 50 mm SL 1s latero-posteriorly edged by deep notch posteriorly followed by prominent anteriorly directed process. Antorbital thin, drop-shaped; sesamoid supraorbital slender, without processes, its length about three times antorbital length. Premaxilla sub-rectangular in dorsal view, laterally narrowing, moderate in length, slightly longer than maxilla. Maxilla boomerang-shaped, slender, slightly curved. Autopalatine sub-rectangular in dorsal view, medial margin sinuous, lateral margin slight- ly concave; autopalatine posterolateral process well-de- veloped, narrow, its length about two thirds autopalatine length excluding posterolateral. Cheek region (Fig. 4B). Metapterygoid thin, subtrian- gular, large, its largest length about equal horizontal length of quadrate excluding dorsal process. Quadrate slender, dorsal process with constricted base, dorsoposterior mar- gin separated from hyomandibula outgrowth by small in- terspace; small process on dorsal margin, just posterior to cartilage block joining quadrate and metapterygoid, laterally overlapping ventral part of metapterygoid. Hyo- mandibula long, with well-developed anterior outgrowth; middle portion of dorsal margin of hyomandibula out- growth with shallow concavity. Opercle relatively robust, opercular odontode patch depth about four fifths of dor- sal hyomandibula articular facet, with 15—18 odontodes; zse.pensoft.net odontodes pointed, slightly curved, arranged in irregu- lar transverse rows; dorsal process of opercle short and pointed; opercular articular face for hyomandibula with prominent trapezoidal lateral flap, separated from small articular facet for preopercle by deep gap. Interopercle moderate, about two thirds hyomandibula length, with 30-36 odontodes; odontodes pointed, arranged in irreg- ular longitudinal rows. Preopercle compact, with short ventral flap. Hyoid region (Fig. 4C). Parurohyal robust, lateral process triangular, straight, laterally directed, tip pointed; parurohyal head well-developed, with indistinct antero- lateral paired process; middle foramen small and ellip- tical; posterior process short, about two thirds distance between anterior margin of parurohyal and anterior inser- tion of lateral process. Ceratohyals slender. Colouration in alcohol (Fig. 1). Flank, dorsum and head side light brown. Dorsum, flank and head with pale dark brown spots, variable in size and shape, often incon- spicuous in specimens from Rio Biguacu basin. Venter yellowish grey. Darker pigment concentrated between anterior and posterior nostrils, around opercular patch of odontodes and posterior portion of caudal peduncle; na- sal and maxillary barbels dark brown, rictal barbel pale brown with dark brown base. Ventral surface of head yellowish white. Unpaired fins dark yellowish grey, dark brown spots on basal portion of dorsal and caudal fins. Paired fins pale yellow, basal portion of pectoral fin dark yellowish grey. Zoosyst. Evol. 97 (1) 2021, 147-159 Colouration in life (Figs. 2A—E). Similar to coloura- tion in alcohol, but yellow pigmentation more intense on trunk and fins orangish yellow in unspotted specimens from Rio Biguacu basin. Distribution and habitat. Cambeva barbosae occurs in fast-flowing low-altitude streams (about 15—190 m asl), of coastal river basins, between the Biguac¢u and the Cu- batéo do Sul river basins, as well as in smaller drainages in the Santa Catarina island (Fig. 5). Etymology. Cambeva barbosae is named in honour of the Brazilian ichthyologist Maria Anais Barbosa, for her efforts to collect and study trichomycterines from Santa Catarina. Remarks. The species collected in the Santa Catarina island and identified as 7richomycterus sp. by Bertaco (2009), probably is Cambeva barbosae, since both were collected at the same locality in Corrego Grande. Cambeva botuvera sp. nov. http://zoobank.org/145BAE1A-008B-4D80-BDFD-C814347B73DB Figs 3F, 4D-F, 6, 7, Table 1 Holotype. UFRJ 6911, 66.9 mm SL; Brazil: Santa Catarina state: Botuvera municipality: village of Ou- rinhos: Ribeiréo Ourinhos, Rio Itajai-Mirim basin, 27°14'22"S, 49°10'22"W, about 160 m asl; C.R.M. Fel- trin, 9 Apr. 2018. Paratypes. All from Brazil: Santa Catarina state: de Botuvera municipality: Rio Itajai-Mirim basin. UFRJ 11920, 6, 36.4—70.9 mm SL; collected with holotype. — UFRJ 12196, 14, 42.6-80.0 mm SL; UFRJ 6912, 3, 39.2— 57.3 mm SL (C&S); stream belonging to the Rio Ourinhos subdrainage, Ourinhos, 27°14'22"S, 49°10'22"W, about 160 m asl; C.R.M. Feltrin, 26 Aug. 2018. — UFRJ 12200, 8, 29.9-72.5 mm SL; stream belonging to the Rio Ourin- hos subdrainage, Ourinhos, 27°14'6"S, 49°10'10"W, about 170 m asl; C.R.M. Feltrin, 26 Aug. 2018. — CIC- CAA 12618, 5, 48.1-66.4 mm SL; stream belonging to the Rio Ourinhos subdrainage, Ourinhos, 27°14'34"S, 49°10'39"W, about 170 m asl; C.R.M. Feltrin, 23 Oct. 2018. — UFRJ 12202, 9, 43.3-71.6 mm SL; stream tributary of Rio Itajai-Mirim, 27°12'18"S, 49°8'14"W, about 170 m asl; C.R.M. Feltrin, 24 Oct. 2018. — UFRJ 11918, 15, 31.4-81.1 mm SL; stream tributary of Rio Ita- jai-Mirim near Lajeado Baixo, 27°12'18"S, 49°8'14"W, about 170 m asl; C.R.M. Feltrin, 8 Apr. 2018. — UFRJ 12195, 7, 31.0-40.0 mm SL; UFRJ 12201, 9, 32.8- 64.4 mm SL; stream tributary of Rio Itajai-Mirim near Lajeado Baixo, 27°12'18"S, 49°8'14"W, about 170 m asl; C.R.M. Feltrin, 25 Aug. 2018. Diagnosis. Cambeva botuvera is distinguished from all other species of the genus, except C. balios (Ferrer & Malabarba, 2013), C. cubataonis (Bizerril, 1994), 153 C. davisi (Haseman, 1911), C. diatropoporos (Ferrer & Malabarba, 2013), C. guareiensis Katz & Costa, 2020, C. horacioi Reis, Frota, Fabrin & da Graca, 2020, C. papillifera, , C. perkos (Datovo, Carvalho & Ferrer, 2012), C. plumbea (Wosiacki, 2005), C. stawiarski (Miranda Ribeiro, 1968), and C. tropeira (Ferrer & Malabarba, 2011), by having seven pectoral-fin rays (vs. five, six or eight). Cambeva botuvera differs from these congeners by the following combination of character states: 16-20 dorsal procurrent caudal-fin rays (vs.14—15 in C. tropeira, 21—22 in C. cubataonis and C. plumbea, and 27-29 in C. stawiarski), 14-16 ventral procurrent caudal-fin rays (vs. 9-13 in C. balios, C. cubataonis, C. davisi, C. diatropoporos, and C. guareiensis);, 39—40 vertebrae (vs. 35-38 in C. cubataonis, C. diatropoporos, C. guareiensis, C. horacioi, and C. stawiarski); eight or nine branchiostegal rays (vs. ten in C. perkos and C. stawiarski); jaw teeth conical (incisiform in C. davisi, C. guareiensis and C. stawiarski),; minute papillae on the ventral surface of the head (vs. hypertrophied in C. papillifera), relatively long maxillary and rictal barbels, reaching between the interopercular patch of odontodes and the pectoral-fin base (vs. rudimentary in C. papillifera); pelvic fin and girdle well-developed (vs. absent in C. tropeira),; anterior segment of the latero-sensory infraorbital series absent (vs. present in C. diatropoporos and C. tropeira); and colouration consisting of dorsum and dorsal portion of flank with rounded brown blotches, without a distinctive yellow longitudinal zone on the dorsal portion of the flank (vs. minutes dots or no distinctive marks in C. papillifera and C. plumbea; presence of a distinctive yellow longitudinal zone on the dorsal portion of the flank in C. perkos). Cambeva botuvera is also distinguished from C. balios, C. cubataonis, C. davisi, C. diatropoporos, C. guareiensis, C. plumbea, and C. tropeira by having a long posterior process of the parurohyal, slightly longer than the length between the anterior-most point of parurohyal head and lateral process insertion (Fig. 4F; vs. shorter). Description. Morphometric data appear in Table 2. Body moderately slender, subcylindrical and slightly depressed anteriorly, compressed posteriorly. Greatest body depth in area just anterior to pelvic-fin base. Dorsal profile of head and trunk slightly convex, approximately straight on caudal peduncle; ventral profile straight to slightly con- vex 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 trapezoidal in dorsal view. Anterior profile of snout convex in dorsal view. Eye small, dor- sally positioned in head. Posterior nostril located nearer anterior nostril than orbital rim. Tip of maxillary and ric- tal barbels reaching area between interopercular patch of odontodes and pectoral-fin base; tip of nasal barbel reach- ing area between eye and opercular patch of odontodes. Mouth subterminal. Jaw teeth 40-42 in both premaxilla zse.pensoft.net 154 Wilson J.E.M. Costa et al.: Two new species of Cambeva Figure 5. Map of geographical distribution of Cambeva in isolated coastal basins in the southern end of the Atlantic Forest, southern Brazil: C. barbosae sp. nov. (dots), C. botuvera sp. nov. (triangles), C. cubataonis (squares); black symbols are type localities, red symbols are paratypes of the new species, and white symbols are additional material, non-types. Table 2. Morphometric data of Cambeva botuvera sp. nov. Holotype Paratypes (n = 12) Standard length (mm) 66.9 39.2-81.1 Percent of standard length Body depth 14.7 13.2-17.9 Caudal peduncle depth U6: 10.1-12.6 Body width o.5 9.1-12.7 Caudal peduncle width Se 2.8-4.5 Pre-dorsal length 62:2 61.0-66.8 Pre-pelvic length 5756. 55.1-61.3 Dorsal-fin base length Ate 5 10.1-11.9 Anal-fin base length o.9 8.5-10.9 Caudal-fin length 15:8 13.7-17.4 Pectoral-fin length iia 11.7-13.8 Pelvic-fin length 8.4 7.9-10.1 Head length 20.9 19.1-21.8 Percent of head length Head depth 46.9 44.7-50.8 Head width 80.7 77.1-85.0 Snout length 42.3 41.9-44.7 Interorbital length 22.0 21.4-25.0 Preorbital length 14.4 14.2-15.8 Eye diameter Os7 9.7-13.2 and dentary, irregularly arranged, pointed and slightly curved. Branchial membrane attached to isthmus only at its anterior point. Branchiostegal rays 8 or 9. Dorsal and anal fins subtriangular; total dorsal-fin rays 12 (i + II + 7), total anal-fin rays 9 (11 + II + 5); zse.pensoft.net anal-fin origin in vertical through middle of dorsal-fin base or slightly posterior to it, approximately at base of 5" branched dorsal-fin ray. Dorsal-fin origin in vertical through centrum of 21‘ or 22" vertebra; anal-fin origin in vertical through centrum of 25" vertebra. Pectoral fin subtriangular in dorsal view, posterior margin slightly convex, first pectoral-fin ray not terminating in filament; total pectoral-fin rays 7 (I + 6). Pelvic fin subtruncate, its extremity in vertical through anterior portion of dor- sal-fin base; pelvic-fin bases medially in close proximity; total pelvic-fin rays 5 (I + 4). Caudal fin truncate, poste- ro-dorsal and postero-ventral extremities rounded; total principal caudal-fin rays 13 (1 + 11 + J), total dorsal pro- current rays 16-19 (xv—xix + I), total ventral procurrent rays 12-16 (xi-xv + I). Vertebrae 39-40. Ribs 12 or 13. Two dorsal hypural plates, corresponding to hypurals 4 + 5 and 3, respectively, often coalesced to form single plate; single ventral hypural plate corresponding to hypurals 1 and 2 and parhypural. Laterosensory system (Fig. 7A, B). Supraorbital sen- sory canal continuous, connected to posterior section of infraorbital 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 nearer orbit than its paired Zoosyst. Evol. 97 (1) 2021, 147-159 155 Figure 6. Cambeva botuvera sp. nov., UFRJ 6911, holotype, 66.9 mm SL: A. left lateral view; B. dorsal view; C, ventral view. homologous pore. Single infraorbital sensory canal seg- ment, with two pores, corresponding to pore 110, adja- cent to ventral margin of orbit, and pore i111, posterior to orbit; anterior segment of infraorbital canal absent. Postorbital canal with 2 pores: pol, in vertical line above posterior portion of interopercular patch of odontodes, and po2, in vertical line above posterior portion of oper- cular patch of odontodes. Lateral line of body short, with 2 pores, posterior-most pore in vertical just posterior to pectoral-fin base. Mesethmoidal region (Fig. 4D). Mesethmoid robust, its anterior margin slightly concave; mesethmoid cornu nar- row, extremity rounded; narrow lateral flap on intersection between cornu and main bone axis, posteriorly extending parallel to lateral bone margin. Lateral ethmoid connect- ed to autopalatine by weak articular facet; minute lateral projection on lateral ethmoid margin close to middle por- tion of sesamoid supraorbital, often absent. Antorbital thin, drop-shaped; sesamoid supraorbital slender, without pro- cesses, its length about three times antorbital length. Pre- maxilla sub-trapezoidal in dorsal view, laterally narrowing, moderate in length, slightly longer than maxilla. Maxilla boomerang-shaped, slender, slightly curved. Autopalatine sub-rectangular in dorsal view, medial margin sinuous, lat- eral margin slightly concave; autopalatine posterolateral process well-developed, narrow, its length about two thirds autopalatine length excluding posterolateral. Cheek region (Fig. 4E). Metapterygoid thin, sub- triangular, large, its largest length slightly shorter than horizontal length of quadrate excluding dorsal process. Quadrate slender, dorsal process with constricted base, dorsoposterior margin separated from hyomandibula out- growth by small interspace. Hyomandibula long, with well-developed anterior outgrowth; middle portion of dorsal margin of hyomandibula slightly concave. Opercle relatively slender, opercular odontode patch depth about half length of dorsal hyomandibula articular facet, with 15-18 odontodes; odontodes pointed, nearly straight, arranged in irregular transverse rows; dorsal process of opercle short and pointed; opercular articular faces for hyomandibula and preopercle rounded and in close prox- imity. Interopercle moderate, about two thirds hyoman- dibula length, with 25—30 odontodes; odontodes pointed, arranged in irregular longitudinal rows. Preopercle com- pact, with short ventral flap. Hyoid region (Fig. 4F). Parurohyal robust, lateral process sub-triangular, slightly curved, latero-poste- riorly directed, tip pointed; parurohyal head well-de- veloped, with indistinct anterolateral paired process; middle foramen broad, oval; posterior process long, zse.pensoft.net 156 Wilson J.E.M. Costa et al.: Two new species of Cambeva Figure 7. Head of Cambeva botuvera sp. nov., UFRJ 6911, holotype, 66.9 mm SL: A. left lateral view; B. dorsal view; C. ventral view. Pores of the cephalic latero-sensory system are indicated in A and B. slightly longer than distance between anterior margin of parurohyal and anterior insertion of lateral process. Ceratohyals slender. Colouration in alcohol (Fig. 6). Flank, dorsum and head side pale yellowish brown. Dorsum and dorsal por- tion of flank with rounded brown blotches, darker and of- ten horizontally coalesced along lateral midline, sometimes zse.pensoft.net forming interrupted or complete darker stripe. Ventral part of flank with pale grey spots, often inconspicuous or ab- sent. Small dark brown spots on lateral and dorsal surfaces of head, darker pigment concentrated between anterior and posterior nostrils and around opercular patch of odontodes; nasal barbel dark brown, maxillary and rictal barbels pale brown with dark brown bases. Venter and ventral surface of Zoosyst. Evol. 97 (1) 2021, 147-159 head yellowish white. Unpaired fins hyaline with yellowish brown bases. Pectoral fin hyaline with dark brown spots on basal portion. Pelvic fin whitish hyaline. Colouration in life (Fig. 3F). Similar to colouration in alcohol, but yellow pigmentation slightly more intense on trunk and fins. Distribution and habitat. Cambeva botuvera occurs in fast-flowing low-altitude streams (about 160—170 m asl), of the Rio Itajai-Mirim basin (Fig. 5). Etymology. The name botuvera is an allusion to the occur- rence of the species in the municipality of Botuvera, Santa Catarina, southern Brazil. This name is derived from the Tupi-Guarani, possibly meaning brilliant mountain. Discussion Cambeva is a morphologically homogeneous genus, with relatively little external morphological variation when compared to the closely related genus 7richomycterus (sensu Katz et al. 2018). However, as here and elsewhere observed (Costa et al. 2020d), osteological characters may be useful to distinguish closely species and to ten- tatively support relationship hypotheses. Unfortunately, due to the absence of osteological data in descriptions of species that are rare in ichthyological collections, present diagnoses are still directed to combinations of character states of the external morphology that may be checked in original descriptions (see diagnoses above). Anyway, the present osteological comparative analysis indicates some osteological characters potentially diagnosing intragener- ic clades that will help to support species allocation in future studies. Molecular phylogenies including species of Cambe- va have been directed to more inclusive trichomycterid groups (Ochoa et al. 2017, 2020; Katz et al., 2018; Costa et al., 2020c), consequently comprising a limited sample of taxa belonging to this genus. Therefore, phylogenetic relationships among species of Cambeva are still poor- ly known (Costa et al., 2020d). However, the available molecular analyses, including a recent unilocus analysis (Donin et al. 2020), have corroborated similar intragener- ic clades, including a clade named as the C. balios group by Costa et al. (2020d), comprising all species from the Lagoa dos Patos system and part of the species from the Parana-Uruguay system, and another clade, here named as C. davisi group, including species from the Rio Parana basin and smaller coastal basins. Costa et al. (2020d) de- scribed some osteological character states presumably supporting the C. balios group and a subclade, but no morphological feature supporting the C. davisi group or its subclades has been described. The present comparative analysis suggests that the new species herein described do not belong to a single intrage- neric clade. According to Costa et al. (2020d), species of the C. balios group have a comparatively long premaxilla, 157 distinctively longer than the maxilla (Costa et al. 2020d: fig. 2A), although this condition is not so evident in C. balios (e.g. Ferrer and Malabarba 2013: fig. 2A). Costa et al. (2020d: fig. 2B) also assigned the presence of a short process on the anteroventral portion of the metapterygoid for a subclade of the C. balios group, including at least C. flavopicta Costa, Feltrin & Katz, 2020 and C. poiki- los Ferrer & Malabarba, 2013, but possibly this unique derived character state is also present in other species of the C. balios group, since a small projection at the same place is present in C. balios. Cambeva flavopicta and C. poikilos share a series of unique derived features in the mesethmoidal region, including a robust autopalatine with a short postero-lateral process (Costa et al. 2020d: fig. 2A). In the species here described, the premaxilla is not so long, there is no similar process on the metaptery- goid, and the autopalatine is relatively slender, with a well-developed postero-lateral process (Fig. 4A, D), as primitively occurring in other trichomycterine taxa. Bockmann et al (2004: fig. 6) described an osseus con- tact area reinforcing metapterygoid-quadrate connection as a unique apomorphic condition in C. diabola. In fact, although variable in shape and size, a flat small process on the dorsal margin of quadrate, laterally overlapping metapterygoid and situated just posterior to the syncon- drial joint between the metapterygoid and the quadrate, 1s also present in species of the C. davisi group, including at least C. castroi, C. davisi, C. guareiensis and C. zona- ta. Among species herein described, this process is small but evident in C. barbosae (Fig. 4B), highly suggesting its inclusion in the C. davisi group. In C. botuvera, the metapterygoid and quadrate are posteriorly separate and there is no process (Fig. 4E). On the other hand, the lat- eral ethmoid of C. barbosae bears a deep notch just pos- terior to the articular face for the autopalatine, posteri- orly followed by a prominent anteriorly directed process (Fig. 4A). A similar condition occurs at least in C. diat- ropoporos and C. poikilos (Ferrer and Malabarba 2013: fig. 2B), two species of the C. balios group. However, both C. flavopicta and species of the C. davisi group often have a small notch followed by a minute process at the same position, suggesting that this morphological feature may be a basal condition for a more inclusive clade in- cluding both the C. balios and C. davisi groups. Phylogenetic relationships of C. botuvera are still ob- scure, since it does not exhibit derived osteological char- acter states here described for species of the C. davisi group and those described by Costa et al. (2020d) for the C. balios group. However, the presence of a lateral flap on the intersection between the cornu and main axis of the mesethmoid, posteriorly extending parallel to the lateral bone margin (Fig. 4D) suggests that C. botuvera may be more closely related to the C. balios group, since a simi- lar condition was observed in some species of this group, including C. balios, C. poikilos, and C. tropeira. Presently, three species are known from the coastal river basins of the Atlantic Forest of southern Brazil, comprising the two new species herein described and C. cubataonis zse.pensoft.net 158 (Bizerril, 1994) (Fig. 5). All these species have similar colour patterns (1.e. dark brown spots over flank, Figs 1, 6; Katz and Barbosa 2014: fig. 1) and distinction is not easy using only characters of the external morphology. However, as occurring in other congeners and most trichomycterines, bone morphology highly differs in these species, showing important diagnostic character states, mainly concentrated in the mesethmoidal region and cheek bones (see discussion above and Fig. 4). Therefore, the regular use of such structures in taxonomical studies will substantially improve trichomycterine taxonomy, especially in groups for which molecular data are not widely available. Acknowledgements We are grateful to Alexandre Bianco, Beatrizz Mesqui- ta, Caroline Freitas, Filipe Pereira, Georg Beckmann, Joao de Bittencourt Vitto, José Leonardo Mattos, Luiz Fernando Ugioni, Maria Anais Barbosa, Pedro Amorim, and Ronaldo dos Santos Jr, for collecting specimens of the new species or assistance during field expeditions in southern Brazil; and to Morevy Cheffe, Roger Dalcin and Vinicius Abilhoa for sending important comparative ma- terial. The final version of the manuscript benefitted from criticisms provided by Walter Azevedo-Santos. Instituto do Meio Ambiente, Santa Catarina, and Instituto Chico Mendes de Conservacéo da Biodiversidade provided collecting permits. This work was partially supported by Conselho Nacional de Desenvolvimento Cientifico e Tec- nologico (CNPq; grant 307349/2015-2 to WJEMC). References Adriaens D, Baskin JN, Coppens H (2010) Evolutionary morphology of trichomycterid catfishes: About hanging on and diggingin. In: Nelson JS, Schultze HP, Wilson MVH (Eds) Origin and Phylogenetic Interre- lationships of Teleosts. Verlag Dr. Friedrich Pfeil, Mtinchen, 337-362. Abilhoa V, Braga RR, Bornatowski H, Vitule JRS (2011) Fishes of the Atlantic Rain Forest streams: ecological patterns and conservation. In: Grillo O, Venora G (Eds) Changing Diversity in Changing Envi- ronmental. 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