Cambeva perobana, Martins & Reis & Stabile & Graça, 2024

Cambeva perobana , new species urn:lsid:zoobank.org:act:C0BF9D84-4F67-4B57-8BCA-267A8F608AC9

( Figs. 1–8; Tab. 1)

Trichomycterus sp. 1 — Delariva, Silva, 2013:552 (Checklist of fishes from Perobas Biological Reserve).

Trichomycterus sp. 2 — Delariva, Silva, 2013:552 (Checklist of fishes from Perobas Biological Reserve).

Trichomycterus sp. 3 — Delariva, Silva, 2013:552 (Checklist of fishes from Perobas Biological Reserve; partim:

NUP 11703).

Trichomycterus sp. — Delariva, Silva, 2014:61 (Checklist of fishes from Perobas Biological Reserve; fig. in p. 61).

Holotype. NUP 23907, 75.9 mm SL, Brazil, Paraná State, municipality of Tuneiras do Oeste, rio Mouro, tributary of rio Goioerê , rio Piquiri basin, upper rio Paraná basin, 23°52’54”S 52°49’46”W, 15 Jul 2011, A. G. Bifi & G. C. Deprá. GoogleMaps

Paratypes. All from Brazil, Paraná State, upper rio Paraná basin. Rio Ivaí basin: NUP 11020, 2, 56.1–70.0 mm SL, municipality of Araruna , rio Ligeiro , tributary of rio Ivaí , 23°50’52”S 52°33’47”W, 25 Oct 2010, C. H. Zawadzki, L. Raisi & G. C. Zawadzki. NUP 11726, 2, 57.8–71.9 mm SL, municipality of Cianorte , rio dos Índios, 23°51’11”S 52°42’03”W, 6 Dec 2010, R. Delariva. NUP 24163, 1, 56.3 mm SL, municipality of Cianorte , rio dos Índios, tributary of rio Ivaí , 23°51’11”S 52°42’03”W, 6 Dec 2010, R. Delariva. NUP 24274, 1 c&s, 50.0 mm SL, municipality of Cianorte , rio dos Índios, 23°51’11”S 52°42’03”W, 6 Dec 2010, R. Delariva. Rio Piquiri basin: MCP 54936 View Materials , 3 View Materials , 67.6–79.5 mm SL, municipality of Tuneiras do Oeste , rio Mouro , tributary of rio Goioerê , 23°53’10”S 52°49’19”W, 1 May 2017, R. Delariva & C. Larentis. NUP 11703, 5, 38.4–67.5 mm SL, municipality of Tuneiras do Oeste , rio Concórdia , tributary of rio Mouro , 23°52’51”S 52°49’56”W, 4 Dec 2010, R. Delariva. NUP 11707, 2, 23.3–58.8 mm SL, municipality of Tuneiras do Oeste , rio Concórdia , tributary of rio Piquiri , 23°52’51”S 52°49’56”W, 4 Dec 2010, R. Delariva. NUP 11712, 4, 27.3–68.8 mm SL, municipality of Tuneiros do Oeste , rio Mouro , tributary of rio Piquiri , 23°52’07”S 52°48’56”W, 6 Dec 2010, R. Delariva. NUP 14648, 5, 22.0– 63.8 mm SL, municipality of Tuneiras do Oeste , rio Saquarema , tributary of rio Piquiri , 23°52’02”S 52°46’30”W, 28 Aug 2011, R. Delariva. NUP 16051, 2, 31.0– 58.3 mm SL, municipality of Janiópolis, NN stream, tributary of rio Barreiro , 24°12’44”S 52°47’50”W, 25 Mar 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 16086, 1, 52.5 mm SL, municipality of Tuneiras do Oeste , rio Água Cinquenta e Cinco, tributary of rio Goioerê , 23°56’02”S 52°45’52”W, 29 Jan 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 17219, 2, 34.5–68.6 mm SL, municipality of Farol, NN stream, tributary of rio Farol , 24°22’45”S 52°40’53”W, 12 Sep 2014, C. H. Zawadzki. NUP 17223, 1, 35.0 mm SL, municipality of Farol, rio Farol , tributary of rio Goioerê , 24°22’45”S 52°40’ 54”W, 12 Sep 2014, C. H. Zawadzki. NUP 17228, 1, 45.7 mm SL, municipality of Farol, NN stream, tributary of córrego Água da Granada, 24°22’53”S 52°35’58”W, 12 Sep 2014, C. H. Zawadzki. NUP 17232, 3, 27.6–43.1 mm SL, municipality of Farol, Córrego Água da Granada, tributary of rio Goioerê , 24°16’55”S 52°41’31”W, 13 Sep 2014, C. H. Zawadzki. NUP 23908, 2, 29.0– 68.3 mm SL, municipality of Tuneiras do Oeste , NN stream, tributary of rio Mouro , 23°52’54”S 52°49’46”W, 23 Mar 2022, A. G. Bifi & G. C. Deprá. NUP 24159, 2, 60.9–67.5 mm SL, same data as holotype. NUP 24160, 1, 41.2 mm SL, municipality of Farol, NN stream, tributary of córrego Água da Granada, 24°22’53”S 52°35’58”W, 12 Sep 2014, C. H. Zawadzki. NUP 24161, 1, 49.9 mm SL, municipality of Farol, NN stream, tributary of rio Farol , 24°22’45”S 52°40’53”W, 12 Sep 2014, C. H. Zawadzki. NUP 24162, 1, 74.7 mm SL, municipality of Tuneiras do Oeste , rio Água Cinquenta e Cinco, tributary of rio Goioerê , 23°56’02”S 52°45’52”W, 29 Jan 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 24164, 2 c&s, 45.7–50.0 mm SL, municipality of Janiópolis, NN stream, tributary of rio Barreiro , 24°12’44”S 52°47’50”W, 25 Mar 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 24165, 1, 68.8 mm SL, rio Concórdia , tributary of rio Mouro , 23°52’54”S 52°49’46”W, 15 Jul 2011, A. G. Bifi & G. C. Deprá. NUP 24166, 2, 46.0– 63.2 mm SL, rio Mouro , tributary of rio Goioerê , 23°52’54”S 52°49’46”W, 9 Sep 2022, I. C. Martins, Renan B. Reis, B. H. M. Stabile & M. Z. Roloff. NUP 24167, 6, 25.9–54.6 mm SL, rio Concórdia , tributary of rio Mouro , 23°53’0.31”S 52°49’52.00”W, 9 Sep 2022, I. C. Martins, R. B. Reis, B. H. M. Stabile & M. Z. Roloff. NUP 24271, 1 c&s, 48.1 mm SL, municipality of Tuneiras do Oeste , rio Água Cinquenta e Cinco, tributary of rio Goioerê , 23°56’02”S 52°45’52”W, 29 Jan 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 24272, 1 c&s, 50.5 mm SL, municipality of Tuneiras do Oeste , rio Água Cinquenta e Cinco, tributary of rio Goioerê , 23°56’02”S 52°45’52”W, 29 Jan 2014, W. J. da Graça, W. M. Domingues, F. A. Teixeira & R. J. da Graça. NUP 25070, 7, 25.7–57.8 mm SL, municipality of Tuneiras do Oeste , rio Concórdia , tributary of rio Mouro , 23°53’03”S 52°49’52”W, 9 Sep 2022, I. C. Martins, R. B. Reis, B. H. M. Stabile & M. Z. Roloff. NUP 25181, 3 c&s, 52.4–75.1 mm SL, municipality of Tuneiros do Oeste , rio Mouro , tributary of rio Piquiri , 23°52’07”S 52°48’56”W, 6 Dec 2010, R. Delariva GoogleMaps .

Non-types. CIG 869, 2, 51.0–67.0 mm SL, municipality of Campo Mourão , rio Mourão, 24°11’18”S 52°22’43”W, 1 Jul 2010, V. A. Frana. NUP 24273, 1 c&s, 54.9 mm SL, municipality of Cianorte, rio dos Índios, 23°51’11”S 52°42’03”W, 6 Dec 2010, R. Delariva GoogleMaps .

Diagnosis. Cambeva perobana can be distinguished from most congeners by having six branched pectoral-fin rays (vs. four in C. alphabelardense Costa, Feltrin & Katz, 2022 ; C. betabelardense Costa, Feltrin & Katz, 2022 , C. pascuali (Ochoa, Silva, Costa e Silva, Oliveira & Datovo, 2017) ; five in C. brachykechenos ( Ferrer & Malabarba, 2013) , C. flavopicta Costa, Feltrin & Katz, 2020 , C. grisea Costa, Feltrin & Katz, 2021 , C. mboycy (Wosiacki & Garavello, 2004) , C. naipi (Wosiacki & Garavello, 2004) , C. podostemophila Costa, Feltrin & Katz, 2023 , C. poikilos ( Ferrer & Malabarba, 2013) , C. taroba (Wosiacki & Garavello, 2004) and C. tourensis Costa, Feltrin & Katz, 2023 ; or seven in C. barbosae Costa, Feltrin & Katz, 2021 , C. castroi (de Pinna, 1992) , C. concolor ( Costa, 1992) , C. crassicaudata (Wosiacki & de Pinna, 2008) , C. diabola ( Bockmann, Casatti & de Pinna, 2004) , C. difficilis Costa, Feltrin & Katz, 2024 , C. guaraquessaba (Wosiacki, 2005) , C. igobi (Wosiacki & de Pinna, 2008) , C. iheringi (Eigenmann, 1917) , C. melanoptera Costa, Abilhoa, Dalcin & Katz, 2022 , C. tupinamba (Wosiacki & Oyakawa, 2005) , C. variegata ( Costa, 1992) , and C. ytororo (Terán, Ferrer, Benitez, Alonso, Aguilera & Mirande, 2017)) . Cambeva perobana can be distinguished from C. chrysornata Costa, Feltrin, Mattos, Dalcin, Abilhoa & Katz, 2023 , C. davisi (Haseman, 1911) , C. orbitofrontalis Costa, Feltrin & Katz, 2021 , and C. stawiarski (Miranda Ribeiro, 1968) by the lower number of interopercular odontodes (28–29 vs. 30–38), or from C. balios ( Ferrer & Malabarba, 2013) , C. biseriata Costa, Feltrin, Mattos, Dalcin, Abilhoa & Katz, 2023 , C. diatropoporos ( Ferrer & Malabarba, 2013) , C. diffusa Costa, Feltrin & Katz, 2021 , C. duplimaculata Costa, Feltrin & Katz, 2021 , C. horacioi , C. imaruhy Costa, Feltrin & Katz, 2021 , C. longipalata Costa, Feltrin & Katz, 2021 , C. notabilis Costa, Feltrin & Katz, 2021 , C. panthera Costa, Feltrin & Katz, 2021 , C. pericoh Costa, Feltrin & Katz, 2021 , C. perkos (Datovo, Carvalho & Ferrer, 2012) , C. plumbea (Wosiacki & Garavello, 2004) , C. tropeiro ( Ferrer & Malabarba, 2011) , C. urubici Costa, Feltrin & Katz, 2021 , and C. ventropapillata Costa, Feltrin, Mattos, Dalcin, Abilhoa & Katz, 2023 by the greater number of interopercular odontodess (28–29 vs. 11–26); from C. balios , C. botuvera Costa, Feltrin & Katz, 2021 , C. chrysornata , C. diffusa , C. duplimaculata , C. gamabelardense Costa, Feltrin & Katz, 2022 , C. guaratuba Costa, Feltrin, Mattos, Dalcin, Abilhoa & Katz, 2023 , C. horacioi , C. imaruhy , C. longipalata , C. notabilis , C. orbitofrontalis , C. paolence (Eigenmann, 1917) , C. pericoh , C. perkos , C. piraquara Reis, Wosiacki, Ferrer, Donin & Graça, 2023 , C. stawiarski , and C. urubici differs by the number of vertebrae (34–37 vs. more than 37); from C. biseriata , C. botuvera , C. chrysornata , C. davisi , C. imaruhy , C. notabilis , C. papillifera (Wosiacki & Garavello, 2004) , C. plumbea , C. stawiarski , and C. urubici differs by the lower number of opercular odontodes (13 vs. 14–28), or from C. cubataonis (Bizerril, 1994) , C. longipalata , and C. panthera differs by the lower number of opercular odontodes (13 vs. 7–11); from C. cubataonis , C. diffusa , C. guaratuba , C. guareiensis , C. imaruhy , C. longipalata , C. notabilis , C. orbitofrontalis , C. panthera , C. pericoh , and C. zonata (Eigenmann, 1918) differs by the number of branchiostegal rays (9 vs. 7–8); from C. papillifera , and C. ventropapillata differs by the absent of conspicuous papillae on ventral region of head (vs. present); and from C. tropeiro is distinguished by the presence of pelvic girdle and fin (vs. absent).

Additionally, C. perobana can be distinguished from all congeners from the upper rio Paraná and rio Iguaçu basin by the presence of pelvic girdle (vs. absence of pelvic girdle in C. pascuali ); by the long rictal barbel, 40.0–73.0% of HL (vs. short, 12.0–30.0% in C. papillifera , 33.0–40.0% in C. castroi ); by the pelvic fins not reaching the urogenital opening (vs. reaching to anal-fin origin in C. crassicaudata and C. paolence , reaching urogenital opening in C. cauim Reis, Ferrer & Graça, 2021 , C. davisi , C. guareiensis Katz & Costa, 2020 , and C. taroba ); by the caudal fin truncate to rounded (vs. caudal fin forked in C. crassicaudata ; distal margin strongly concave in C. cauim , or slightly concave in C. stawiarski ); by the first pectoral-fin ray not prolonged as a filament (vs. prolonged as a rudimentary filament in C. pascuali , and C. piraquara ; short filament in C. paolence , and some specimens of C. piraquara ; and long filament in C. taroba ); by the color pattern of the caudal fin, presenting blotches, spots, or a homogeneous dark-brown or gray pattern in the proximal region (vs. presenting a pale-yellow to unpigmented stripe in the proximal region in C. castroi , C. diabola , C. difficilis , and C. melanoptera ); by the presence of a narrow mid-lateral stripe in some specimens (vs. presence of four narrow stripes on the body: mid-sagital, mid-dorsal, mid-lateral, and ventro-lateral stripes in C. naipi ); by the shorter head, 17.2–20.6% of SL (vs. longer, 23.8–26.8% of SL in C. igobi ); by the longer pelvic and pectoral fins, 10.3–16.0% of SL and 7.5–9.8%, respectively (vs. 7.7–9.1%, and 5.6–7.3%, respectively, in C. mboycy ); by the color pattern of some specimens, consisting in a dark-gray to dark-brown, becoming lighter towards ventral portion of the body and head, and absence of chromatophores, or some specimens with irregular dark-brown blotches on the inner skin layer, larger than two or three times the diameter of the orbit, sometimes coalescing and forming larger blotches, and an interrupted stripe in the mid-lateral region of the flank, over a plain yellowish to brown background (vs. color pattern consisting in dorsal and lateral surface of body with scattered circular well-defined dark-brown blotches, variable in size in inner skin layer and small black spots on the outer skin layer in C. horacioi ; or lateral and dorsal surface of body with numerous small spots, irregularly distributed, and well-defined dark-brown blotches, larger than orbit diameter, along dorsal and mid-lateral surface of body in C. iheringi ); and by the number of branchiostegal rays (9 vs. 7–8 in C. iheringi ).

Description. Summarized morphometric data of two morphotypes in Tab 1. Body elongate, trunk roughly cylindrical close to head and gradually becoming laterally compressed towards caudal fin. Dorsal profile of trunk slightly convex along anterior half of body to insertion of dorsal fin. Ventral profile of trunk slightly convex. Dorsal and ventral profiles of caudal peduncle slightly convex.

Head depressed, trapezoidal in dorsal view, wider posteriorly and anterior portion slightly rounded. Dorsal and ventral profiles of head straight to slightly convex in lateral view. Eyes located dorsolaterally on anterior region of head, at same longitudinal line of nasal barbel. Eyes with a round to elliptical shape anteroposteriorly, covered by thin and translucent skin. Orbital rim not free. Eyes visible from lateral view.

Anterior nostril slightly smaller than diameter of eye, surrounded by flap of integument posterolaterally continuous with base of nasal barbel. Posterior nostril slightly smaller than diameter of eye, surrounded anterolaterally by thin flap of integument. Gill openings not constricted, forming free fold reaching pectoral-fin insertion. Mouth subterminal and slightly curved with corners posteriorly oriented in ventral view. Upper lip thicker laterally. Lower lip with conspicuous fleshy lobes at corners of mouth, continuous with base of rictal barbels. Lips with small rounded papillae of approximately same size.

Barbels with broad bases, tapering gradually towards tips. Nasal barbel emerging from posterolateral region of anterior nostril with tip surpassing infraorbital pores i10 when adpressed to head. Maxillary barbel emerging from corner of mouth with tip reaching to anterior region of interopercular odontodophore when adpressed to head. Rictal barbel emerging from corner of mouth, shorter than maxillary barbel.

Pectoral fin with distal margin rounded, I,6*(40), first ray unbranched, not prolonged as filament. Pelvic fin with distal margin rounded, not covering anterior margin of urogenital papilla; with I,4* rays (40). Pelvic-fin insertion anterior to dorsal-fin origin. Inner margins of pelvic fins close basally. Urogenital papilla closer to distal margin of pelvic fin than to origin of anal fin. Dorsal fin with distal margin rounded, ii(5), II,7* rays (40). Origin of dorsal fin located at vertical through last third of pelvic fin. Anal fin elongated with distal margin rounded and slightly smaller than dorsal fin, ii(5), II,5*(29) or II,6(11) rays. Origin of anal fin located at vertical through half or last third of dorsal-fin base. Caudal fin with distal margin rounded or truncate in specimens smaller than 35.0 mm SL; upper caudal plate with I,5* rays (40), lower caudal plate with I,6* rays (40).

Osteology. Mesethmoid with anterior margin straight to slightly concave and cornua short, with tapering distal ends. Anterior cranial fontanel restricted to small,

rounded opening situated between frontals and epiphyseal bar. Posterior cranial fontanel long and wide extending from posterior portion of frontals to parieto-supraoccipital.

Epiphyseal bar longer than wide. Antorbital anteriorly expanded and posteriorly elongated, extending over anterior third of autopalatine. Barbular bone elongate, with medial process in anterior third. Sphenotic, prootic, and pterosphenoid fused, anterior portion anterolaterally directed in dorsal view ( Fig. 2). Vomer arrow-shaped with long posterior process extending to parasphenoid. Parasphenoid with long and pointed posterior process extending to basioccipital. Weberian capsule with lateral openings and anterior margin fused to the basioccipital.

Premaxilla rectangular with 38 or 40 (2) spatulate to conical teeth similar in size and roughly distributed in four irregular rows. Dentary teeth extending from coronoid process base to near dentary symphysis ( Figs. 3A–B). Maxilla boomerang-shaped,

shorter than premaxilla. Autopalatine with lateral margin concave; anterior margin slightly convex; medial margin slightly concave and long posterior process extending over posterior portion of metapterygoid. Metapterygoid large and laminar, connected to quadrate through cartilage; posterior portion with notch (arrow in Figs. 3C–D). Quadrate L-shaped with concavity in anterior portion. Hyomandibula well developed, dorsal margin with concavity ( Figs. 3C–D). Opercle longer than interopercle. Opercular odotodophores ovoid to rounded with 13 (2) conical odontodes, gradually curving medially and increasing in size posteriorly, arranged in five irregular transverse rows. Interopercular patch of odontodes elongate with 28 or 29 (2) conical odontodes, arranged in two transverse rows.

Ventral hypohyal trapezoid-shaped. Anterior ceratohyal elongate and wider at anterior and posterior ends. Posterior ceratohyal short, triangular, with rounded tips. Nine branchiostegal rays (2): six in contact with anterior ceratohyal, one with interceratohyal cartilage, and two with posterior ceratohyal. Four posteriormost branchiostegal rays wider distally ( Figs. 4A–B). Urohyal with expanded anterior head, two elongate lateral processes with wide bases and decreasing in width distally with rounded tips, and sharp and elongated posterior process. Posterior process of urohyal shorter than lateral processes.

Basibranchials 2 and 3 elongated, connected by cartilage; basibranchial 2 slightly wider than basibranchial 3. Basibranchial 4 hexagonal and entirely cartilaginous. Hypobranchial 1 elongated, with cartilaginous tips, approximately of same size than basibranchial 2. Hypobranchials 2 and 3 with narrow anterolateral ossified processes with large area of cartilage posteriorly; hypobranchials 2 and 3 equal in size. Five elongate ceratobranchials with cartilaginous tips. Ceratobranchial 3 with prominent concavity on posterior margin. Ceratobranchial 5 with 16 or 20 (2) conical, elongated, and pointed teeth, arranged in three irregular rows. Four epibranchials; anteriormost three elongated and narrow. Epibranchials 1 L shaped, with anterior pointed process; epibranchial 2 with mesial-anterior and distal-posterior process; epibranchial 3 with wider process on distal-posterior margin, slightly curved mesially. Epibranchial 4 rectangular. Pharyngobranchial 3 straight and elongated, shorter than hypobranchial 1. Pharyngobranchial 4 ossified and connected to curved plate with 23 or 28 (2) conical, elongated, and pointed teeth, arranged in up to three irregular rows; teeth increasing in size posteriorly ( Figs. 4C–D).

Dorsal fin with eight pterygiophores (4) or seven (2), first inserted anterior to neural spine of 18 th or 19 th post-weberian vertebrae. Anal fin with six pterygiophores (6), first inserted anterior to hemal spine of 22 nd post-weberian vertebrae. Procurrent caudal-fin rays 17(1), 18(1), or 19(4) dorsally and 12(1), 13(2), or 14(22) ventrally. Upper caudal plate with one unbranched ray and five branched rays; hypural 3 free and hypurals 4 and 5 fused to each other. Single lower caudal plate with one unbranched ray and six branched rays. Parhypural and hypurals 1 and 2 co-ossified and fused to compound caudal centrum. Post weberian vertebrae 34(1), 35(1), 36(3), 37(3), ribs 13(4), or 15(1).

Laterosensory system. Laterosensory canals with simple (non-dendritic) branches ending in single pores. Nasal and frontal branches of supraorbital canal continuous, with three paired pores s1, s3, and s6. Supraorbital pore s1 located at posterior portion of anterior nostrils, pore s3 at same longitudinal line of pore s1, posteriorly to posterior nostrils, and pore s6 aligned with posterior margin of eyes. Antorbital segment of infraorbital canal absent. Sphenotic canal present with two pores, i10 located behind eyes, and i11 located laterally to posterior margin of eye. Otic and postotic canals present with two pores associated: po1 located anterolaterally to opercular odontodophore and po2 located laterally to half-length of opercular odontodophore. Lateral line canal short with two* (36) pores located above pectoral-fin insertion and posterior to gill opening.