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Research Article
Top mountain areas of subtropical southern Brazil sheltering four new small-ranged catfishes (Siluriformes, Trichomycteridae): relationships and taxonomy
expand article infoWilson J. E. M. Costa‡, Caio R. M. Feltrin‡, José Leonardo O. Mattos‡, Axel M. Katz‡
‡ Federal University of Rio de Janeiro, Rio de Janeiro, Brazil

Corresponding author: Wilson J. E. M. Costa ( wcosta@acd.ufrj.br )

Academic editor: Oliver Hawlitschek
© 2024 Wilson J. E. M. Costa, Caio R. M. Feltrin, José Leonardo O. Mattos, Axel M. Katz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Costa WJEM, Feltrin CRM, Mattos JLO, Katz AM (2024) Top mountain areas of subtropical southern Brazil sheltering four new small-ranged catfishes (Siluriformes, Trichomycteridae): relationships and taxonomy. Evolutionary Systematics 8(2): 199-218. https://doi.org/10.3897/evolsyst.8.126393
ZooBank: urn:lsid:zoobank.org:pub:973DECB5-340A-41CE-9AEE-B968664489B2
Open Access

Abstract

Mountainous regions typically host a great diversity of small-ranged species, often contributing for delineating world biodiversity hotspots. Species of trichomycterine catfishes have been recorded for several high-altitude areas of tropical South America, but field inventories in top mountains of southern Brazil are still rare. Here we report four new small-ranged species collected in streams of the Rio Iguaçu at Serra do Espigão (RISE) in altitudes between about 970 and 1020 m asl, one in the eastern portion of RISE and three in the western portion. A molecular phylogenetic analysis indicated that these species belong to the Cambeva beta-clade, which comprises all species endemic to the Rio Iguaçu drainage, but together not forming a monophyletic group. The analysis also indicated that species endemic to high altitudes are variably related to species from lower altitudes. The only eastern RISE species appears in a basal position of a well-supported clade (Cambeva beta1-clade), with the western RISE species appearing in a subclade of the Cambeva beta1-clade with species occurring in a vast area of southern Brazil. New species are diagnosed by combinations of morphological character states, including meristic, colouration, latero-sensory system, and osteological data.

Key Words

Cambeva, Molecular phylogeny, Mountain biodiversity, Rio Iguaçu drainage, Serra do Espigão

Introduction

Mountainous areas of tropical and subtropical regions of the world host a great biodiversity, commonly concentrating numerous small-ranged species (Rahbek et al. 2007, 2019) and contributing for delineation of the most important biodiversity hotspots (Myers et al. 2000). Among fish groups, Trichomycterinae (hereafter trichomycterines) is the most diverse in South American river mountains (e.g. Costa et al. 2021), with species being sporadically recorded from high altitude localities of Andes since the 18th century (e.g. Valenciennes 1832) until the 20th one (e.g. Arratia and Menu Marque 1984). Recent studies, however, have consistently recorded the occurrence of numerous new trichomycterines from various Andean regions with geographic distribution restricted to small areas in altitudes between around 1,200 and 4,000 m asl (e.g. Fernández and Schaefer 2003; DoNascimiento et al. 2014; Fernández and Liota 2016; Fernández et al. 2023). Species occurring in high Andean areas are often confined to particular environments such as caves (e.g. Castellanos-Morales 2008, 2018; Mesa et al. 2018), phreatic waters (e.g. Fernández and de Pinna 2005) and thermal water wells (Fernandéz and Vari 2012). On the other hand, in the mountain ranges of subtropical southern Brazil, where higher altitudes barely surpass 1800 m asl, recent studies also have revealed some small-ranged species of Cambeva Katz, Barbosa, Mattos & Costa, 2018 only found at altitudes above 900 m asl (e.g. Ferrer and Malabarba 2011; Costa et al. 2021, 2023a), but field inventories at higher areas of this region are still rare and many sites remain unsampled.

With a surface area of about 72,600 km2, the Rio Iguaçu drainage, a main tributary of the Rio Paraná basin, is situated in a region characterized by a sequence of mountain ranges and plateaus (i.e. Serra da Esperança, Serra do Espigão, Serra do Mar; e.g. Ab’Saber 2007). This landscape contains extensive areas of rapids and waterfalls inhabited by a remarkable diversity of species of Cambeva, with a great concentration reported for some areas. For example, Wosiacki and collaborators described six species and reported the occurrence of other two already described species from the Rio Jordão, a tributary of the middle Rio Iguaçu (Wosiacki and Garavello 2004; Wosiacki and de Pinna 2008a, b). A total of 13 nominal species have been recorded for this drainage (Haseman 1911; Miranda Ribeiro 1968; de Pinna 1992; Wosiacki and Garavello 2004; Wosiacki and de Pinna 2008a, b; dos Reis et al. 2021, 2023; Costa et al. 2022) and phylogenetic studies have indicated that all species belong to a single large intrageneric clade, called Cambeva beta-clade (Costa et al. 2023b). However, all records of Cambeva for the Rio Iguaçu drainage are concentrated in the northern part of the drainage, with no data available about its occurrence in the southern part of the drainage, where rivers and streams drain the Serra do Espigão, which is part of a long chain of mountain ranges that stretches across southern Brazil under the name Serra Geral. The section known as Serra do Espigão forms a long plateau running east-west, with higher average altitudes between 900 and 1200 m, having its eastern portion in contact with the coastal mountain range chain called Serra do Mar.

A great diversity of species of Cambeva was found during a detailed recent field inventory (March/April 2023) in rivers and streams of an area about 9,000 km2 belonging to the Rio Iguaçu drainage at Serra do Espigão (hereafter RISE) by one of us (CRMF). Most of these species were found in about 25 localities of a broad area at altitudes below about 840 m asl and were conspecific or morphologically similar to species widely distributed in other parts of the drainage (i. e. Cambeva naipi (Wosiacki & Garavello, 2004), Cambeva papillifera (Wosiacki & Garavello, 2004), Cambeva stawiarski (Miranda Ribeiro, 1968) or occurring in adjacent coastal basins to the east (i.e. C. barbosae Costa, Feltrin & Katz, 2021). Contrastingly, four undescribed species were found in isolated points of RISE at altitudes between about 970 and 1020 m asl, one of them in the eastern portion of RISE, and three in neighbouring drainages at the western portion. All these species were found only in small areas, despite sampling efforts at various points in different altitudes of their sub-drainages. Furthermore, they were rare in their habitats, making necessary a second collecting trip (June/July 2023) to supplement the material necessary for adequate descriptions. The objectives of the present study are to perform a molecular phylogenetic analysis to positioning the new species among the main intrageneric lineages and to describe the four new species.

Materials and methods

Specimens

Specimens were captured using dip nets during daylight. Collecting permits were provided by ICMBio (Instituto Chico Mendes de Conservação da Biodiversidade; permit number: 38553-13). Methods for collections were approved by CEUA-CCS-UFRJ (Ethics Committee for Animal Use of Federal University of Rio de Janeiro; permit numbers: 065/18 and 084/23). Fixation and preservation of specimens as described in Costa et al. (2023b). Osteological preparations followed Taylor and Van Dyke (1985). Specimens were deposited in Instituto de Biologia, Universidade Federal do Rio de Janeiro (UFRJ) and Centro de Ciências Agrárias e Ambientais, Universidade Federal do Maranhão (CICCAA). Abbreviations used in list of specimens are: C&S, cleared and stained specimens for osteological examination; SL, standard length. In localities where specimens were collected, geographical names are according to Portuguese names used in the region. Comparative material is listed in our previous studies on the genus Cambeva (Costa et al. 2023a, b and included references).

Morphological data

Measurements were according to landmarks described in Costa (1992), modified in Costa et al. (2020a) and were made in well-preserved specimens above about 40 mm SL. Meristic data and fin ray formulae followed Costa et al. (2020a) based on Costa (1992) and Bockmann and Sazima (2004). Osteological morphology was primarily approached using a stereomicroscope Zeiss Stemi SV 6 with camera lucida. Terminology for osteological structures is according to Costa (2021), with more recent updates described in Kubicek (2022). Nomenclature for pores of the latero-sensory system followed Arratia and Huaquin (1995), except for post-orbital pores as proposed by Bockmann and Sazima (2004).

DNA extraction, amplification and sequencing

DNA extraction, amplification and sequencing followed the same methods as described in our recent phylogenetic studies on Cambeva (e.g. Costa et al. 2023b), using the same markers: the mitochondrially encoded genes cytochrome b (CYTB) and cytochrome c oxidase I (COX1) and the nuclear encoded gene recombination activating 2 (RAG2). We used the following primers: Cytb Siluri F and Cytb Siluri R (Villa-Verde et al. 2012), CatThr29 and Glu 31 (Unmack et al. 2009), and Glu 5 and Cb23 (Barros et al. 2015) for CYTB; FISHF1 and FISHR1 (Ward et al. 2005) for COX1; MHRAG2-F1 and MHRAG2-R1 (Hardman and Page 2003), RAG2 TRICHO F and RAG2 TRICHO R (Costa et al. 2020b), and RAG2 MCF and RAG2 MCR (Cramer et al. 2011) for RAG2. PCR reaction parameters are those described in Costa et al. (2023b). Reading and interpretation of sequencing chromatograms and sequence edition were performed using MEGA 11 (Tamura et al. 2021). GenBank accession numbers appear in Table 1.

Table 1.
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Terminal taxa, and GenBank accessions numbers by gene used in molecular analysis. Asterisks (*) indicate the newly added sequences.

COI CYTB RAG2
Trichogenes longipinnis OQ810037 MK123704 MF431117
Listrura tetraradiata JQ231083 JQ231088.1 MN385826.1
Ituglanis boitata OQ810038 MK123706 MK123758
Trichomycterus itatiayae MW671552 MW679291 OL779233
Scleronema minutum MK123685 MK123707 MK123759.1
Cambeva variegata PP319019 PP328534 PP333217
Cambeva zonata KY857986 KY858053 –
Cambeva brachykechenos MN995669 MN995758 –
Cambeva diatropoporos KY857996 KY858065 KY858213
Cambeva stawiarski MN995720 MN995779 –
Cambeva perkos KY857981 KY858050 –
Cambeva poikilos KY857995 KY858064 –
Cambeva guaraquessaba MN995662 MN995749 –
Cambeva naipi MN995699 MN995771 –
Cambeva taroba MN995708 MN995757 –
Cambeva tupinamba MN995656 MN995751 –
Cambeva tropeira MN995674 MN995752 –
Cambeva grisea MN995671 MN995760 –
Cambeva imaruhy MN995700 MN995766 OQ814191
Cambeva orbitofrontalis MN995703 MN995764 OQ814192
Cambeva cubataonis OQ095914 OQ110814 OQ110815
Cambeva iheringi GU701893 KY858074 KY858223
Cambeva barbosae MK123689.1 OQ110808 OQ110815.1
Cambeva diabola JN989258 OQ110812 –
Cambeva balios OQ810040 OQ814186 OQ814193
Cambeva chrysornata MN995726 OQ110810 OQ110819
Cambeva pascuali MF034463 OQ110811 OQ110820
Cambeva guaratuba MN995721 MN995792 –
Cambeva panthera OQ810041 OQ814187 OQ814194
Cambeva flavopicta OQ810042 OQ814188 OQ814195
Cambeva podostemophila OQ810043 OQ814189 OQ814196
Cambeva tourensis MN995697 OQ814190 OQ814197 13283
Cambeva davisi KR140345 MK123714 MK123762
Cambeva luteoreticulata PP448186 * PP449083 * PP449087 *
Cambeva atrobrunnea PP448187 * PP449084 * PP449088 *
Cambeva rotundipinna PP448188 * PP449085 * PP449089 *
Cambeva galactica PP448189 * PP449086 * PP449090 *
Cambeva biseriata PP448190 * OQ110806 OQ110817
Cambeva ventropapillata PP448191 * OQ110807 OQ110818
Cambeva guareiensis PP448192 * OQ110813 OQ110821
Cambeva castroi – MK123712 OQ110816

Phylogenetic analyses

Terminal taxa comprised the four new species and other 19 species of the Cambeva beta-clade, besides 13 congeners of other lineages and five outgroup species belonging to other trichomycterid lineages. The analysis comprised both DNA sequences here generated and those taken from our previous studies (Katz et al. 2018; Costa et al. 2023a, b) and from GenBank, first published in Ochoa et al. (2017a, 2017b) and Donin et al. (2022). The concatenated molecular data matrix comprised 2469 bp (COX1 685 bp, CYTB 993 bp, RAG2 791 bp). Each gene data set was individually aligned using the Clustal W algorithm (Chenna et al. 2003) implemented n MEGA 11. No stop codons and gaps were found. PartitionFinder2.1.1 (Lanfear et al. 2016) algorithm was used to calculate best-fit evolutive model schemes (Table 2), under the Corrected Akaike Information Criterion. Phylogenetic analyses were conducted using Bayesian Inference (BI) and Maximum Likelihood (ML) approaches. BI was performed in Beast 1.10.4 (Suchard at al. 2018), using two independent Markov Chain Monte Carlo (MCMC) runs with 3 × 107 generations, with sampling frequency of 1000 generation, using Tracer 1.7.2 (Rambaut et al. 2018) to verify convergence of the MCMC chains and the proper burn-in value. LogCombiner v.1.10.4 (Suchard et al. 2018) and Tree Annotator version 1.10.4 (Suchard et al. 2018) were used to combine and calculate the consensus tree, apply the 25% burn-in, and annotate the Bayesian posterior probabilities. ML was performed using IQTREE 2.2.0 (Minh et al. 2020), with node support evaluated by ultrafast bootstrap (UFBoot) (Hoang et al. 2018) and the Shimodaira-Hasegawa-like approximate likelihood ratio test (SH-aLRT), each using 1000 replicates.

Table 2.
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Optimal partition schemes with their respective quantities of base pairs and the best-fitting evolutive models.

Partition Base pairs Evolutive Model
COI 1st 229 GTR+G
COI 2nd 228 TRN+G
COI 3rd 228 HKY+I
CYTB 1st 331 TRNEF+I+G
CYTB 2nd 331 HKY+I+G
CYTB 3rd 331 TRN+G
RAG2 2nd, 3rd 527 GTR+I
RAG2 1st 263 K80+I

New species diagnoses

Species here described were primarily diagnosed using unique morphological character states (i.e. not occurring in all other species of Cambeva), followed by unique combinations of morphological character states (see Davis and Nixon 1992). Morphological diagnoses were followed by molecular diagnoses based on the partial sequences of the mitochondrial genes COX1 and CYTB used the phylogenetic analysis (see above), following Costa et al. (2014) and Aguiar et al. (2022), using PAUP* 4 (Swofford 2002) to track base pair transformations. The position of each base pair relative to the entire gene was estimated by aligning the fragments with the nearly complete mitochondrial DNA genome of Trichomycterus areolatus Valenciennes, 1846 (accession number AP012026). In addition, although genetic distances alone are not informative to delimit species (e.g. DeSalle et al. 2005), we provided pairwise COX1 genetic distances between each new species and other species of the Cambeva beta-clade, calculated using the Kimura 2-parameter (Kimura 1980) model algorithm, implemented in MEGA 11, to illustrate interspecific ranges, indicating taxa with shortest distances).

Results

Phylogenetic relationships and comparative morphology

Both analyses generated similar topologies (Fig. 1), in which the Cambeva beta-clade is corroborated, including Cambeva naipi Wosiacki & Garavello, 2004 as the sister group to all other species. All the four RISE species are corroborated as members of the Cambeva beta-clade, but appearing in different sections of the tree, with the western RISE species (C. galactica Costa, Feltrin & Katz, sp. nov.) supported as a basal species of an inclusive well-supported clade (hereafter Cambeva beta-1 clade) that also includes the three other RISE species. The three eastern RISE species are positioned in a subclade that also includes other species endemic to the Rio Iguaçu drainage (Cambeva castroi (Pinna, 1992), Cambeva davisi (Haseman, 1911), Cambeva stawiarski (Miranda Ribeiro, 1968)), as well as species that occur in adjacent coastal areas (Cambeva barbosae Costa, Feltrin & Katz, 2020) or in other drainages of the Paraná River basin (Cambeva diabola (Bockmann, Casatti & de Pinna, 2004), Cambeva guareiensis Katz & Costa, 2020). The analyses indicated that three western RISE species do not form a monophyletic group, but their position was weakly supported. In spite of a superficial similarity in their external morphology, the osteological analysis revealed a highly divergent bone morphology (see descriptions below), suggesting that they are not close relatives.

Figure 1.

Bayesian phylogenetic tree obtained by BEAST for 36 species of Cambeva species and five outgroups, using three genes (COI, CYTB, and RAG2; total of 2469 bp). Species in red are the new species herein described. Numbers separated by bars (/) above branches indicate posterior probabilities from the Bayesian Inference, and ultrafast bootstrap (UFBoot) and the Shimodaira-Hasegawa-like approximate likelihood ratio test (SH-aLRT) from the Maximum Likelihood analysis inferred in IQTREE2. Asterisks (*) indicate maximum support values, and dashes (-) indicate support values below 50.

Taxonomic accounts

The four species here described share several morphological character states of the external morphology that are common among congeners of the beta-clade. In order to avoid unnecessarily repeating the same characteristics in each individual species description, below is a general description of character states shared by all four species.

General description of species from RISE

Body moderately slender, subcylindrical in anterior region, compressed in posterior region. Greatest body depth in area midway between pectoral-fin and pelvic-fin bases. Dorsal and ventral profile slightly convex between snout and dorsal-fin origin, about straight along caudal peduncle. Anus and urogenital papilla at vertical through middle part of dorsal-fin base or immediately posterior to it. Head sub-trapezoidal in dorsal view. Anterior profile of snout slightly convex in dorsal view. Eye small, dorsally positioned on head, in its anterior half. Distance between anterior and posterior nostrils shorter than distance between posterior nostril and orbit. Minute skin papillae on head surface. Mouth subterminal.

Supraorbital sensory canal continuous, posteriorly connected to posterior section of infraorbital canal, with three pores: s1, adjacent to medial margin of anterior nostril; s3, adjacent and just posterior to medial margin of posterior nostril; s6, in transverse line through posterior half of orbit. Pore s6 nearer orbit than its paired homologous pore. Posterior infraorbital sensory canal with two pores: pore i10, adjacent to ventral margin of orbit, and pore i11, posterior to orbit. Postorbital canal with two pores: po1, at vertical through posterior portion of interopercular patch of odontodes, and po2, at vertical through posterior portion of opercular patch of odontodes. Lateral line with two pores situated above and slightly posterior to pectoral-fin base.

Eastern RISE species

Cambeva galactica Costa, Feltrin & Katz, sp. nov.

https://zoobank.org/D7C5FF3B-FE1F-4F06-A8E4-B3F0D1ABF829
Figs 2, 3, 4A–C, Table 3

Type material

Holotype. Brazil • 1 ex., 76.9 mm SL; Santa Catarina State: Rio Negrinho Municipality: village of Volta Grande: stream tributary of Rio Preto, itself a tributary of Rio Negro, Rio Iguaçu drainage, Rio Paraná basin; 26°33'04"S, 49°40'14"W; about 970 m asl; 29 Mar. 2023; C.R.M. Feltrin, leg.; UFRJ 14064.

Paratypes. Brazil • 4 ex., 18.7–40.6 mm SL; same data as for holotype; UFRJ 13516. • 3 ex. (C&S), 40.5–64.9 mm SL; same data as for holotype; UFRJ 14065. • 2 ex. (DNA), 16.5–38.8 mm SL; same data as for holotype; UFRJ 13517. • 5 ex., 28.8–108.0 mm SL; same locality and collector as holotype; 28 Jun. 2023; UFRJ 13847.

Diagnosis

Cambeva galactica is distinguished from all congeners by a unique colour pattern in adult specimens consisting of flank and dorsum with longitudinal rows of interconnected yellowish white vermiculate diffuse marks (vs. never a similar colour pattern), and the presence of a distinctive projection on the anterior portion of the medial margin of the sesamoid supraorbital, connected by thin ligamentous tissue to a dorsal projection on the articulatory shell of the autopalatine for the lateral ethmoid (Fig. 4A; vs. never a similar process connected to that articulation). Cambeva galactica is also distinguished from all other species of the Cambeva beta-clade, except Cambeva flavopicta Costa, Feltrin & Katz, 2020, Cambeva naipi (Wosiacki & Garavello, 2004), Cambeva taroba (Wosiacki & Garavello, 2004), and Cambeva tourensis Costa, Feltrin & Katz, 2023 by having six pectoral-fin rays (vs. five, seven, or eight). Cambeva galactica is distinguished from C. flavopicta and C. tourense by having well-developed pelvic fins (vs. absent); from C. naipi and C. taroba by having more interopercular odontodes 29–32 vs. 11 or 12 in C. naipi and 17–21 in C. taroba) and more jaw teeth (41–43 on the premaxilla and 39–46 on the dentary, vs. 25–34 and 23–32, respectively); from C. naipi by having 14 or 15 ribs (vs. 12 or 13), fewer opercular odontodes (eight or nine vs 12 or 13), and from C. taroba by having a minute pectoral-fin filament, its length less than 5% of the pectoral-fin length (vs. about 20%), fewer procurrent caudal-fin rays (18 or 19 dorsal and 12 or 13 ventral, vs. 26 or 27 and 21–23, respectively), more vertebrae (39 or 40 vs 36). Molecular diagnosis: 34 nucleotide substitutions, nine of them unique among taxa analysed*, and five unique for the Cambeva beta-clade **: COX1 103 (T→C)**, COX1 103 (G→A), COX1 117 (A→T)*, COX1 243 (A→G)*, COX1 309 (G→A), COX1 312 (C→A), COX1 321 (C→A), COX1 330 (C→A)*, COX1 483 (T→C), COX1 540 (A→G), COX1 547 (C→T), COX1 684 (A→C)*, CYTB 195 (T→C), CYTB 219 (T→C), CYTB 282 (C→T)**, CYTB 283 (T→C), CYTB 339 (C→T), COX1 342 (C→T), CYTB 394 (C→T), CYTB 399 (T→C)**, CYTB 495 (G→A), CYTB 528 (A→G), CYTB 585 (T→C)**, CYTB 711 (C→T)*, CYTB 715 (G→A)*, CYTB 735 (C→T)*, CYTB 771 (C→T)*, CYTB 822 (A→G), CYTB 849 (T→C), CYTB 891 (T→C)*, CYTB 900 (T→C)*, CYTB 909 (A→G), CYTB 994 (A→C), CYTB 1032 (C→T)**. COX1 p-distances among congeners of the Cambeva beta-clade ranging from 2.9 (Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov. and Cambeva ventropapillata Costa, Feltrin & Katz, 2022) and 4.7.

Description

Morphometric data appear in Table 3.

Table 3.
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Morphometric data of Cambeva galactica Costa, Feltrin & Katz, sp. nov.

Holotype Paratypes (n = 6)
Standard length (SL) 76.9 40.1–108.0
Percentage of standard length
Body depth 16.4 15.1–16.2
Caudal peduncle depth 11.8 10.5–12.7
Body width 11.6 9.8–12.6
Caudal peduncle width 4.5 3.4–5.5
Pre-dorsal length 65.3 62.6–65.8
Pre-pelvic length 60.0 58.5–62.4
Dorsal-fin base length 10.9 11.1–12.2
Anal-fin base length 9.3 8.4–10.0
Caudal-fin length 17.6 15.4–18.3
Pectoral-fin length 13.3 11.6–14.4
Pelvic-fin length 9.9 8.7–9.4
Head length 22.5 20.6–23.1
Percentage of head length
Head depth 44.3 42.2–52.0
Head width 80.4 78.5–90.5
Snout length 42.1 35.1–43.6
Interorbital width 24.4 25.5–26.9
Preorbital length 11.9 9.7–11.7
Eye diameter 9.7 6.9–11.9

Head morphology. Barbels moderate in length. Nasal barbel reaching area just anterior to opercle, maxillary barbel reaching between interopercular patch of odontodes and pectoral-fin base, and rictal barbel reaching posterior portion of interopercular patch of odontodes. Jaw teeth pointed, irregularly arranged. Premaxillary teeth 41–43, dentary teeth 39–46. Opercular and interopercular odontodes pointed, about straight. Opercular odontodes 8 or 9; interopercular odontodes 29–32. Anterior infraorbital sensory canal present.

Fin morphology. Dorsal and anal fins subtriangular, margins slightly convex. Total dorsal-fin rays 11 or 12 (ii–iii + II–III + 6–7), total anal-fin rays 9 or 10 (ii–iii + II + 5). Anal-fin origin at vertical just posterior to middle dorsal-fin base, at base of 3rd or 4th branched dorsal-fin ray. Pectoral fin sub-triangular in dorsal view, margins slightly convex, first pectoral-fin ray slightly longer than second ray, weakly extending beyond fin membrane forming minute filament. Total pectoral-fin rays 6 (I + 5). Pelvic fin rounded, its tip reaching vertical through middle portion of dorsal-fin base. Total pelvic-fin rays 5 (I + 4). Caudal fin subtruncate, corners rounded. Total principal caudal-fin rays 13 (I + 11 + I), total dorsal procurrent rays 18 or 19 (xvii–xviii + I), total ventral procurrent rays 12 or 13 (xi–xii + I).

Osteology (Fig. 4A–C). Mesethmoid narrow anteriorly, with lateral expansion in area just anterior to lateral ethmoid, anterior mesethmoid margin slightly concave, with minute anterior projection on its middle portion. Mesethmoid cornu extremity pointed. Lateral ethmoid with small lateral projection immediately posterior to articular facet for autopalatine. Anterodorsal portion of lateral ethmoid widened, projecting laterally. Lacrimal thin, elliptical. Sesamoid supraorbital gently curved, its longitudinal length about two times and half longer than lacrimal longitudinal length, its largest width about equal to lacrimal width. Medial margin of anterior portion of sesamoid supraorbital with distinctive projection, connected by thin ligamentous tissue to dorsal projection on articulatory shell of autopalatine for lateral ethmoid. Premaxilla long, laterally narrowing. Maxilla slender, with rudimentary posterior process, slightly curved, its length about four fifths of premaxilla. Autopalatine sub-trapezoidal in dorsal view, medial margin sinuous, lateral margin weakly concave. Autopalatine postero-lateral process triangular, short, its length about half autopalatine length.

Metapterygoid trapezoid, deeper than long, large, its surface about twice quadrate lateral surface. Quadrate with deep anterior constriction at dorsal process base. Hyomandibula long, anterior outgrowth horizontal length longer than largest horizontal metapterygoid length. Posterior margin of hyomandibula with small projection just above articular facet for opercle. Dorsal margin of hyomandibula outgrowth concave. Opercle elongate, longer than interopercle. Opercular odontode patch very slender, its depth about one third hyomandibula articular facet length. Dorsal process of opercle short, subtriangular, its extremity rounded. Opercular articular facet for hyomandibula with dorsal, rounded laminar projection. Articular facet for preopercle rounded, well-developed. Interopercle relatively long, interopercular odontode patch length longer than hyomandibula outgrowth length. Preopercle slender, with minute ventral projection.

Parurohyal thin, lateral process narrow, slightly curved posteriorly, with rounded extremity. Parurohyal head with prominent anterolateral paired process. Parurohyal middle foramen small, rounded. Parurohyal posterior process moderate in length, about three fourths of distance between anterior margin of parurohyal and anterior insertion of posterior process. Branchiostegal rays 8. Vertebrae 39 or 40. Ribs 14 or 15. Dorsal-fin origin at vertical through centrum of 21st or 22nd vertebra; anal-fin origin at vertical through centrum of 21st or 22nd vertebra. Two dorsal and single ventral hypural plate.

Colouration in alcohol. In adult specimens (Fig. 2), flank, dorsum, and head side pale brown; three longitudinal rows of interconnected yellowish white, diffuse, vermiculate marks: row on dorsum, with marks forming reticulate pattern along pre-dorsal midline; one row on dorsal portion of flank, comprising minute marks; and row on ventral portion of flank. Venter and ventral surface of head yellowish white. Barbels pale brown. Fins greyish hyaline. In juvenile specimens between about 25 and 50 mm SL (Fig. 3), flank and dorsum pale yellow, with broad dark brown stripe along flank longitudinal midline and dark brown reticulate pattern on dorsum, and dorsal and ventral portions of flank. In juvenile specimens smaller than 20 mm SL, flank and dorsum pale yellow, with narrow black stripe along flank longitudinal midline and longitudinal series of black blotches between dorsum and flank, and longitudinal series of small black dots on ventral portion of flank.

Figure 2.

Cambeva galactica Costa, Feltrin & Katz, sp. nov., holotype, UFRJ 14064, 76.9 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Figure 3.

Cambeva galactica Costa, Feltrin & Katz, sp. nov., paratype, UFRJ 13847, 33.4 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Figure 4.

Osteological structures of: A–C. Cambeva galactica Costa, Feltrin & Katz, sp. nov.; D–F. Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov.; G–I. Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov., and J–L. Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov. A, D, G, J. Mesethmoidal region and adjacent structures, left and middle portions, dorsal view; B, E, H, K. Left jaw suspensorium and opercular series, lateral view. C, F, I, L. Parurohyal, ventral view. Abbreviation of structure indicated by arrow: aes, anteromedial expansion of sesamoid supraorbital. Larger stippling represents cartilaginous areas.

Distribution

Cambeva galactica is only known from its type locality in the upper Rio Preto drainage, which is a tributary of the Rio Negro, Rio Iguaçu drainage, Rio Paraná basin, at about 970 m asl (Fig. 5).

Figure 5.

Geographical distribution of four new species of Cambeva from the Rio Iguaçu drainage, Serra do Espigão, southern Brazil: 1. Cambeva galactica Costa, Feltrin & Katz, sp. nov.; 2. Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov.; 3. Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov.; 4. Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov.

Etymology

The name galactica is derived from the Ancient Greek word galaktikós meaning milky, an allusion to the rows of yellowish white diffuse vermiculate marks present in the flank of the new species, reminiscent of the Milky Way.

Western RISE species

Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov.

https://zoobank.org/A2FF0368-0311-4739-9CB2-415343090897
Figs 4D–F, 6, 7, Table 4

Type material

Holotype. Brazil • 70.1 mm SL; Santa Catarina State: Timbó Grande Municipality: stream tributary of Rio Timbó, Rio Iguaçu drainage, Rio Paraná basin; 26°34'41"S, 50°40'28"W; about 970 m asl; 29 Jun. 2023; C.R.M. Feltrin, leg; UFRJ 14066.

Paratypes. Brazil • 7 ex., 26.6–70.1 mm SL; same data as holotype; UFRJ 13821. • 2 ex. (C&S), 27.4–44.2 mm SL; same data as holotype; UFRJ 14067. • 2 ex., 29.9–46.8 mm SL. same locality and collector as holotype; 31 Mar. 2023; UFRJ 13553. • 3 ex. (DNA), 27.4–44.2 mm SL; same locality and collector as holotype; 31 Mar. 2023; UFRJ 13554.

Diagnosis

Cambeva atrobrunnea is distinguished from all other congeners by having the two posterior-most dorsal and ventral procurrent caudal-fin rays segmented (vs. only the posterior-most ray segmented). Cambeva atrobrunnea is also distinguished from the two other species of western RISE, Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov. and Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov., by having subtruncate caudal fin (vs. rounded), fewer interopercular odontodes (20–22 vs. 29–34), and specimens below about 40 mm SL having flank light grey with small black dots that are arranged in irregular rows, coalesced on the anterior portion of the longitudinal midline to form a stripe (Fig. 7A; vs. pale yellow with large, irregularly shaped dark brown to black blotches, sometimes forming longitudinal stripes in the area between dorsum and flank in C. luteoreticulata Costa, Feltrin & Katz, sp. nov., and light brownish yellow with small black dots irregularly arranged in C. rotundipinna Costa, Feltrin & Katz, sp. nov. (see descriptions below); from C. luteoreticulata Costa, Feltrin & Katz, sp. nov. by specimens above 40 mm SL having the nasal barbel reaching area just anterior to opercle (vs. reaching area anterior to orbit), fewer ventral procurrent caudal-fin rays (13 or 14 vs. 15 or 16), and jaw teeth irregularly arranged (vs. arranged in three rows); and from C. rotundipinna Costa, Feltrin & Katz, sp. nov. by having more procurrent caudal-fin rays (21 dorsal and 13 or 14 ventral, vs. 15–17 and 10 or 11, respectively), and more opercular odontodes (12 vs. eight or nine). Cambeva atrobrunnea is also distinguished from all the remaining congeners from the Rio Iguaçu drainage by the following combination of character states: seven pectoral-fin rays (vs. eight in Cambeva castroi (de Pinna, 1992), Cambeva melanoptera Costa, Abilhoa, Dalcin & Katz, 2022, Cambeva crassicaudata (Wosiacki & de Pinna, 2008), Cambeva stawiarski (Miranda Ribeiro, 1968), and six in C. galactica, C. naipi, and C. taroba); absence of the anterior infraorbital (vs. presence in C. galactica, C. naipi, Cambeva papillifera (Wosiacki & Garavello, 2004), Cambeva plumbea (Wosiacki & Garavello, 2004), and C. taroba); posterior margin of the caudal fin slightly convex (vs. about straight in C. crassicaudata, C. davisi, C. galactica, C. melanoptera, C. papillifera, Cambeva piraquara dos Reis, Wosiacki, Ferrer, Donin & da Graça, 2022, C. plumbea, C. igobi (Wosiacki & de Pinna, 2008); straight to slightly concave in C. stawiarski; bilobed in C. crassicaudata; and emarginate in Cambeva cauim dos Reis, Ferrer & Graça, 2021); absence of hypertrophied papillae on the ventral surface of the head (vs. presence in C. papillifera); absence of pectoral-fin filament (vs. presence a well-developed filament in C. taroba and a rudimentary filament in C. davisi, C. galactica, and C. piraquara); 12 opercular odontodes (vs. broad, with 17 or 18 in Cambeva mboycy (Wosiacki & Garavello, 2004); 15 or 16 in C. davisi; seven or eight in C. taroba); 20–22 interopercular odontodes (vs. 12 or 13 in C. naipi); 21 dorsal procurrent caudal-fin rays (vs. 15–17 in C. castroi and C. melanoptera; 18 or 19 in C. naipi; 25–29 in C. crassicaudata, C. igobi, C. mboycy, C. stawiarski, and C. taroba; 30 or 31 in C. cauim); 13 or 14 ventral procurrent caudal-fin rays (vs. 21 – 23 in C. taroba); dorsal-fin origin at a vertical through the centrum of the 21st or 22nd vertebra (vs. 19th or 20th in C. crassicaudata, C. mboycy, and C. stawiarski); jaw teeth pointed, irregularly arranged (vs. incisiform and arranged in rows in C. davisi); and 39 or 40 vertebrae (36 in C. taroba). Molecular diagnosis: combination of 17 nucleotide substitutions, five of them unique among taxa analysed*, and four unique for the Cambeva beta-clade **: COX1 216 (A→C)*, COX1 249 (T→C), COX1 252 (G→A), COX1 471 (G→A), COX1 534 (G→A), CYTB 108 (C→T), CYTB 186 (C→T)**, CYTB 237 (C→T)*, CYTB 360 (T→C)**, CYTB 378 (C→T)*, CYTB 442 (C→T)*, CYTB 765 (C→T), CYTB 909 (A→T)**, CYTB 942 (A→G), CYTB 960 (A→T)*, CYTB 994 (A→C). COX1 p-distances among congeners of the Cambeva beta-clade ranging from 1.2 (Cambeva chrysornata Costa, Feltrin & Katz, 2022 and Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov.) and 3.9.

Description

Morphometric data appear in Table 4.

Table 4.
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Morphometric data of Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov.

Holotype Paratypes (n = 5)
Standard length (SL) 70.1 46.8–78.5
Percentage of standard length
Body depth 14.2 12.2–15.0
Caudal peduncle depth 13.7 11.2–13.1
Body width 12.1 10.3–12.8
Caudal peduncle width 4.3 3.0–5.0
Pre-dorsal length 67.7 63.5–67.2
Pre-pelvic length 62.7 57.9–60.3
Dorsal-fin base length 13.4 10.9–12.5
Anal-fin base length 10.0 9.0–12.7
Caudal-fin length 17.1 16.0–19.0
Pectoral-fin length 12.0 11.6–13.1
Pelvic-fin length 8.7 9.1–12.4
Head length 22.3 19.5–24.7
Percentage of head length
Head depth 43.9 39.7–51.9
Head width 82.7 65.6–88.0
Snout length 39.9 36.1–46.6
Interorbital width 20.3 18.3–23.6
Preorbital length 12.2 10.6–15.4
Eye diameter 8.4 8.2–12.4

Head morphology. Barbels moderate in length. Nasal barbel reaching between orbit and opercular patch of odontodes, maxillary barbel reaching posterior portion of interopercular patch of odontodes, and rictal barbel reaching middle of interopercular patch of odontodes. Jaw teeth variable in shape, smaller teeth slightly pointed, larger teeth sub-incisiform with slightly rounded tip, irregularly arranged. Premaxillary teeth 39–41, dentary teeth 40. Opercular and interopercular odontodes pointed, about straight. Opercular odontodes 12; interopercular odontodes 20–22. Anterior infraorbital sensory canal absent.

Fin morphology. Dorsal and anal fins subtriangular, distal margin slightly convex. Total dorsal-fin rays 10 or 11 (ii + II–III + 6–7), total anal-fin rays 9 or 10 (ii + II + 5–6). Anal-fin origin at vertical just posterior to middle of dorsal-fin base, at base of 4th branched dorsal-fin ray. Pectoral fin sub-triangular in dorsal view, margins slightly convex, first pectoral-fin ray shorter than second ray, not forming terminal filament. Total pectoral-fin rays 7 (I + 6). Pelvic fin rounded, its tip reaching vertical through anterior portion of dorsal-fin base. Total pelvic-fin rays 5 (I + 4). Caudal fin subtruncate, posterior margin weakly convex. Total principal caudal-fin rays 13 (I + 11 + I), total dorsal procurrent rays 21 (xix + II), total ventral procurrent rays 13 or 14 (xi–xii + II).

Osteology (Fig. 4D–F). Mesethmoid broader anteriorly, without lateral expansions in its main axis, anterior mesethmoid margin slightly convex. Mesethmoid cornu extremity rounded. Lateral ethmoid with small lateral projection immediately posterior to articular facet for autopalatine and small, twisted expansion on anterior margin. Lacrimal narrow, short and thin. Sesamoid supraorbital rod-shaped, narrower than lacrimal, its longitudinal length about two times and half longer than lacrimal longitudinal length, without lateral expansions. Premaxilla long, laterally narrowing, slightly curved. Maxilla slender, with rudimentary posterior process, slightly curved, its length about three fourths of premaxilla. Autopalatine sub-trapezoidal in dorsal view, medial margin sinuous, lateral margin weakly concave. Autopalatine postero-lateral process triangular, short, its length about half autopalatine length excluding anterior cartilage. Autopalatine articulation for lateral ethmoid with laminar shovel-shaped expansion.

Metapterygoid sub-rectangular, longer than deep, relatively large, its surface greater than quadrate lateral surface. Areas anterior and posterior to cartilaginous articulation between metapterygoid and quadrate with small laminar overlapped expansions forming additional points of articulation. Quadrate with deep anterior constriction at dorsal process base. Hyomandibula long, anterior outgrowth horizontal length slightly longer than largest horizontal metapterygoid length; dorsal margin of hyomandibula outgrowth concave. Opercle elongate, longer than interopercle. Opercular odontode patch slender, its depth about half hyomandibula articular facet length. Dorsal process of opercle short, subtriangular, its extremity rounded. Opercular articular facet for hyomandibula with dorsal, trapezoidal laminar projection, articular facet for preopercle small, rounded. Interopercle moderate in length, interopercular odontode patch length about equal hyomandibula outgrowth length. Preopercle slender, with minute ventral projection.

Parurohyal robust, lateral process subtriangular, slightly curved posteriorly, with pointed tip. Parurohyal head with prominent anterolateral paired process. Parurohyal middle foramen relatively large, oval. Parurohyal posterior process moderate in length, about half of distance between anterior margin of parurohyal and anterior insertion of posterior process. Branchiostegal rays 9. Vertebrae 39 or 40. Ribs 14 or 15. Dorsal-fin origin at vertical through centrum of 21st or 22nd vertebra; anal-fin origin at vertical through centrum of 25th or 26th vertebra. Two or one dorsal and single ventral hypural plate.

Colouration in alcohol. In adult specimens (Fig. 6), flank, dorsum, and head side light brownish yellow, with great concentration of small dark brown to black dots. Venter and ventral surface of head brownish white, with minute dark brown dots in area just anterior to pelvic fin. Nasal and maxillary barbels brown, rictal barbel brownish white. Fins hyaline, with minute dark brown dots on basal region of dorsal, anal and pectoral fins, and on whole caudal fin. In juvenile specimens below about 50 mm SL (Fig. 7), flank, dorsum and head side light grey, with small black dots arranged in irregular longitudinal rows, coalesced on anterior portion of longitudinal midline of flank to form black stripe.

Figure 6.

Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov., holotype, UFRJ 14066, 70.1 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Figure 7.

Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov., paratype, UFRJ 13821, 33.4 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Distribution

Cambeva atrobrunnea is known from a single locality in a stream tributary of the Rio Timbó, Rio Iguaçu drainage, Rio Paraná basin, at about 970 m asl (Fig. 5).

Etymology

From the Latin ater (dull black, dark) and brunneus (brown), referring to the predominant colour of the flank in adult specimens of the new species.

Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov.

https://zoobank.org/E144F2B3-9DEE-4A23-930B-B5B336070D65
Figs 4G–I, 8, 9, Table 5

Type material

Holotype. Brazil • 81.6 mm SL; Santa Catarina State: Matos Costa Municipality: village of Colônia Cerne: stream tributary of Rio Liso, itself a tributary of Rio Pintado, Rio Iguaçu drainage, Rio Paraná basin; 26°24'25"S, 51°00'45"W; about 1,015 m asl; 1 Apr. 2023; C.R.M. Feltrin and L. Sebben, leg.; UFRJ 14068.

Paratypes. Brazil • 7 ex. 25.2–70.8 mm SL; same data as holotype; UFRJ 13562. • 3 ex. (DNA), 22.7–42.6 mm SL; same data as holotype; UFRJ 13563. • 15 ex., 27.6–77.2 mm SL; same locality and collectors as holotype; 1 Jul. 2023; UFRJ 13828. • 4 ex. 33.7–75.9 mm SL; same locality and collectors as holotype; 1 Jul. 2023; CICCAA 08268. • 4 ex. (C&S), 38.2–65.9 mm SL; same locality and collectors as holotype; 1 Jul. 2023; UFRJ 14069.

Diagnosis

Cambeva luteoreticulata differs from all other congeners by its unique rounded, stapula-shaped caudal fin in specimens above about 40 mm SL (Fig. 8A). Cambeva luteoreticulata is also distinguished from all other congeners of the Cambeva beta-clade, except Cambeva chrysornata Costa, Feltrin, Mattos, Dalcin, Abilhoa & Katz, 2023 and C. papillifera, by having short barbels, with the nasal barbel not reaching the orbit in specimens above 60 mm SL and maxillary and rictal barbels not reaching the interopercular patch of odontodes. Cambeva luteoreticulata also differs from C. chrysornata and C. papillifera by the absence of the anterior segment of the infraorbital series (vs. presence), from C. chrysornata by having more procurrent caudal-fin rays (21 or 22 dorsal and 15 or 16 ventral, vs. 16 or 17 and 11 or 12, respectively) and fewer opercular odontodes (10–12 vs. 18), and from C. papillifera by the absence of hypertrophied papillae on the head surface (vs. presence) and narrow nasal barbel (vs. broad laminar, ribbon-shaped). Cambeva luteoreticulata is also distinguished from all other congeners by a unique pattern of ontogenetic colouration change consisting of flank pale yellow with irregularly shaped and arranged, dark brown to black blotches in specimens below about 40 mm SL (Fig. 9A), becoming dark brown, with small, irregularly shaped pale yellow marks forming reticulate pattern in larger specimens (Fig. 8A). Cambeva luteoreticulata also differs from all the remaining congeners from the Rio Iguaçu drainage by the following combination of character states: seven pectoral-fin rays (vs. eight in C. castroi, C. melanoptera, C. crassicaudata, C. stawiarski; and six in C. galactica, C. naipi, and C. taroba); absence of the anterior infraorbital (vs. presence in C. galactica, C. naipi, C. plumbea, and C. taroba); posterior margin of the caudal fin convex (vs. about straight in C. crassicaudata, C. davisi, C. galactica, C. melanoptera, C. papillifera, C. piraquara, C. plumbea, C. igobi; straight to slightly concave in C. stawiarski; bilobed in C. crassicaudata; and emarginate in Cambeva cauim); absence of pectoral-fin filament (vs. presence a well-developed filament in C. taroba and a rudimentary filament in C. davisi and C. piraquara); nine or 10 opercular odontodes (vs. broad, with 17 or 18 in C. mboycy; 15 or 16 in C. davisi; seven or eight in C. taroba); 29–32 interopercular odontodes (vs. 12 or 13 in C. naipi; 17–21 in C. mboycy and C. taroba); 25 dorsal procurrent caudal-fin rays (vs. 15–17 in C. castroi and C. melanoptera; 18 or 19 in C. naipi;; 30 or 31 in C. cauim); 15 or 16 ventral procurrent caudal-fin rays (vs. 21–23 in C. taroba); dorsal-fin origin at a vertical through the centrum of the 21st or 22nd vertebra (vs. 19th or 20th in C. crassicaudata, C. mboycy, and C. stawiarski); larger jaw teeth incisiform, teeth arranged in rows (vs. pointed to slight rounded, irregularly arranged in all species, except C. davisi); and 38–40 vertebrae (36 in C. taroba). Molecular diagnosis: 17 nucleotide substitutions, five of them unique among taxa analysed* and one unique for the Cambeva beta-clade **: COX1 252 (A→G), COX1 564 (A→C)*, COX1 648 (C→T)*, COX1 678 (A→G), CYTB 123 (C→T), CYTB 206 (G→C)*, CYTB 294 (C→T)*, CYTB 474 (G→A), CYTB 495 (G→A), CYTB 636 (C→T)*, CYTB 675 (T→C), CYTB 696 (A→G), CYTB 696 (A→G), CYTB 879 (A→G)**, CYTB 988 (G→A), CYTB 996 (A→G), CYTB 1038 (C→T). COX1 p-distances among congeners of the Cambeva beta-clade ranging from 1.5 (Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov. and Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov.) and 4.3.

Figure 8.

Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov., holotype, UFRJ 14068, 81.6 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Figure 9.

Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov., paratype, UFRJ 13562, 33.1 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Description

Morphometric data appear in Table 5.

Table 5.
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Morphometric data of Cambeva luteoreticulata Costa, Feltrin & Katz, sp. nov.

Holotype Paratypes (n = 10)
Standard length (SL) 81.6 44.4–77.2
Percentage of standard length
Body depth 13.6 13.3–16.0
Caudal peduncle depth 12.7 11.8–13.8
Body width 10.1 10.3–12.8
Caudal peduncle width 4.1 3.3–5.4
Pre-dorsal length 67.9 61.0–67.8
Pre-pelvic length 61.3 59.3–65.2
Dorsal-fin base length 11.0 10.6–12.3
Anal-fin base length 8.6 7.8–9.5
Caudal-fin length 13.1 11.5–16.1
Pectoral-fin length 9.6 8.1–12.7
Pelvic-fin length 7.2 5.8–8.9
Head length 21.6 21.4–23.8
Percentage of head length
Head depth 45.0 42.6–51.7
Head width 80.3 78.1–84.1
Snout length 40.3 37.9–42.4
Interorbital width 21.2 18.7–24.6
Preorbital length 13.7 9.8–14.1
Eye diameter 8.2 8.6–12.2

Head morphology. Barbels short. Nasal barbel reaching area anterior to orbit in specimens above 60 mm SL, between orbit and area just posterior to it in smaller specimens, and maxillary and rictal barbels reaching area just anterior to interopercular patch of odontodes. Jaw teeth variable in shape, smaller teeth slightly pointed, larger teeth incisiform with slightly rounded tip, arranged in three series. Premaxillary outer row with 14 or 15 teeth, middle row with 16 or 17 teeth, and inner row with 18 teeth; total premaxillary teeth 49. Dentary outer row with 10 or 11 teeth, middle row with 14 or 15 teeth, and inner row with 17–20 teeth; total dentary teeth 42–45. Opercular and interopercular odontodes pointed, about straight. Opercular odontodes 10–12; interopercular odontodes 29–33.

Fin morphology. Dorsal and anal fins subtriangular, distal margin slightly convex. Total dorsal-fin rays 11 (ii + II + 7), total anal-fin rays 9 (ii + II + 5). Anal-fin origin at vertical through posterior portion of dorsal-fin base, at base of 5th branched dorsal-fin ray. Pectoral fin rounded in dorsal view, first pectoral-fin ray shorter than second ray, not forming terminal filament. Total pectoral-fin rays 7 (I + 6). Pelvic fin rounded, its tip reaching vertical through dorsal-fin origin. Total pelvic-fin rays 5 (I + 4). Caudal fin short, rounded, forming spatula-shaped tail in specimens above about 40 mm SL. Total principal caudal-fin rays 13 (I + 11 + I), total dorsal procurrent rays 21 or 22 (xx-xxi + I), total ventral procurrent rays 15 or 16 (xiv–xv + I).

Osteology (Fig. 4G–I). Mesethmoid distinctively broader anteriorly, with lateral expansion in area just anterior to lateral ethmoid, anterior mesethmoid margin about straight to slightly convex. Mesethmoid cornu extremity rounded. Lateral ethmoid with small lateral projection immediately posterior to articular facet for autopalatine. Lacrimal thin, elliptical. Sesamoid supraorbital about two times and half longer than lacrimal, without lateral expansions, its width about equal to lacrimal width. Premaxilla long, laterally narrowing, slightly curved. Maxilla slender, without posterior process, slightly curved, its length about four fifths of premaxilla. Autopalatine sub-trapezoidal in dorsal view, medial margin sinuous, lateral margin weakly concave. Autopalatine postero-lateral process triangular, short, its length about half autopalatine length.

Metapterygoid sub-rectangular, deeper than long, relatively large, its surface greater than quadrate lateral surface. Quadrate with deep anterior constriction at dorsal process base. Hyomandibula long, anterior outgrowth horizontal length slightly longer than largest horizontal metapterygoid length; dorsal margin of hyomandibula outgrowth straight anteriorly, with pronounced U-shaped concavity posteriorly. Opercle elongate, longer than interopercle. Opercular odontode patch slender, its depth about half hyomandibula articular facet length. Dorsal process of opercle short, subtriangular, its extremity rounded. Opercular articular facet for hyomandibula with dorsal, broad, rounded laminar projection, articular facet for preopercle rudimentary. Interopercle moderate in length, interopercular odontode patch length about equal hyomandibula outgrowth length. Preopercle slender, with minute ventral projection.

Parurohyal robust, lateral process subtriangular, slightly curved posteriorly, with pointed tip. Parurohyal head with prominent anterolateral paired process. Parurohyal middle foramen relatively large, oval. Parurohyal posterior process moderate in length, about three fifths of distance between anterior margin of parurohyal and anterior insertion of posterior process. Branchiostegal rays 9 or 10. Vertebrae 38–40. Ribs 14–16. Dorsal-fin origin at vertical through centrum of 21st or 22nd vertebra; anal-fin origin at vertical through centrum of 25th or 26th vertebra. Two dorsal and single ventral hypural plate.

Colouration in alcohol. In adult specimens (Fig. 8A), flank, dorsum and head side dark brown, with small, irregularly shaped, irregularly arranged, pale yellow marks forming reticulate pattern. Nasal and maxillary barbels brown, rictal barbel grey. Venter and ventral surface of head yellowish white. Fins pale grey with black spots on basal region, dark brown dots in middle region. In juvenile specimens below about 40 mm SL (Fig. 9A), flank, dorsum and head side pale yellow with large dark brown to black blotches, more concentrated and sometimes forming longitudinal stripes in area between dorsum and flank.

Distribution

Cambeva luteoreticulata is known from a single locality in a stream tributary of the Rio Liso, Rio Iguaçu drainage, Rio Paraná basin, at about 1,015 m asl (Fig. 5).

Etymology

From the Latin luteus (saffron yellow) and reticulata (reticulated), in reference to the flank colour pattern of adult specimens.

Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov.

https://zoobank.org/1CD4DCB4-9F79-489D-B33E-420B2B935A57
Figs 4J–L, 10, 11, Table 6

Type material

Holotype. Brazil • 78.0 mm SL; Brazil: Santa Catarina State: Matos Costa Municipality: Paca road: Rio da Paca, tributary of Rio Jangada, Rio Iguaçu drainage, Rio Paraná basin; 26°25'02"S, 51°15'42"W; about 1000 m asl; 1 Apr. 2023; C.R.M. Feltrin and L. Sebben, leg.; UFRJ 14070.

Paratypes. All from Santa Catarina State: Matos Costa Municipality: Paca road: Rio da Paca, tributary of Rio Jangada, Rio Iguaçu drainage, Rio Paraná basin. Brazil • 2 ex., 44.4–76.3 mm SL; same data as holotype; UFRJ 13820. • 3 ex. (C&S), 30.3–55.2 mm SL; collected with holotype; UFRJ 14071. • 1 ex., 30.8 mm SL; collected with holotype; UFRJ 13559. • 1 ex. (DNA), 27.8 mm SL; 26°25'06"S, 51°16'30"W; about 1000 m asl; 30 Jun. 2023; C.R.M. Feltrin, leg.; UFRJ 13560.

Diagnosis

Cambeva rotundipinna differs from all other congeners of the Cambeva beta-clade, except C. luteoreticulata, by having a relatively short and rounded caudal fin in specimens above about 60 mm SL (Fig. 10A; vs. subtruncate, truncate, emarginate or forked). Cambeva rotundipinna differs from C. luteoreticulata by having fewer procurrent caudal-fin rays (15–17 dorsal and 10 or 11 ventral, vs. 21 or 22 and 15 or 16, respectively), jaw teeth irregularly arranged (vs. arranged in three rows), more opercular odontodes (14–17 vs. nine or ten), longer nasal barbel in specimens above 60 mm SL, reaching area between orbit and opercular patch of odontodes (vs. reaching area anterior to orbit), and a colour pattern of juveniles, in which the flank is light brownish yellow with small black dots irregularly arranged (vs. pale yellow with large, irregularly shaped dark brown to black blotches, more concentrated on its dorsal portion). Cambeva rotundipinna also differs from C. atrobrunnea, another species from western RISE, by having more odontodes (14–17 opercular and 30–34 interopercular, vs. 12 and 20–22, respectively) and a different juvenile colour pattern, comprising black dots irregularly arranged on the flank (Fig. 11A; vs. black dots arranged in irregular longitudinal rows, coalesced to form stripe on the anterior flank midline, Fig. 7A). Cambeva rotundipinna also differs from both C. atrobrunnea and C. luteoreticulata by having a broader autopalatine, with its largest width about equal to its length (Fig. 4J, vs. narrower, Fig. 4D, G) and a shorter posterior process of the parurohyal, its length about one third of the distance between the anterior margin of the parurohyal and the anterior insertion of the posterior process (Fig. 4L, vs. longer, Fig. 4F, I). Cambeva rotundipinna is also distinguished from all other species of Cambeva endemic to the Rio Iguaçu drainage by the following combination of character states: seven pectoral-fin rays (vs. eight in C. castroi, C. melanoptera, C. crassicaudata, C. stawiarski; and six in C. galactica, C. naipi, and C. taroba); absence of the anterior infraorbital (vs. presence in C. galactica, C. naipi, C. papillifera, C. plumbea, and C. taroba); posterior margin of the caudal fin convex (vs. about straight in C. crassicaudata, C. davisi, C. galactica, C. melanoptera, C. papillifera, C. piraquara, C. plumbea, C. igobi; straight to slightly concave in C. stawiarski; bilobed in C. crassicaudata; and emarginate in C. cauim); absence of hypertrophied papillae on the ventral surface of the head (vs. presence in C. papillifera); absence of pectoral-fin filament (vs. presence a well-developed filament in C. taroba and a rudimentary filament in C. davisi and C. piraquara); nine or 10 opercular odontodes (vs. broad, with 17 or 18 in C. mboycy; 15 or 16 in C. davisi; seven or eight in C. taroba); 30–34 interopercular odontodes (vs. 12 or 13 in C. naipi; 17–21 in C. mboycy and C. taroba); 15–17 dorsal procurrent caudal-fin rays (vs. 25–29 in C. crassicaudata, C. igobi, C. mboycy, C. stawiarski, and C. taroba; 30 or 31 in C. cauim); 10 or 11 ventral procurrent caudal-fin rays (vs. 15 or 16 in C. naipi, 21–23 in C. taroba); dorsal-fin origin at a vertical through the centrum of the 21st or 22nd vertebra (vs. 19th or 20th in C. crassicaudata, C. mboycy, and C. stawiarski); jaw teeth pointed, irregularly arranged (vs. incisiform and arranged in rows in C. davisi); and 39 vertebrae (36 in C. taroba). Molecular diagnosis: 10 nucleotide substitutions, two of them unique among taxa analysed *: COX1 105 (G→A), COX1 252 (G→A), COX1 360 (G→A), COX1 462 (T→G)*, COX1 534 (G→A), CYTB 69 (A→G)*, CYTB 195 (T→C), CYTB 219 (T→C), CYTB 483 (A→G), CYTB 637 (G→A). COX1 p-distances among congeners of the Cambeva beta-clade ranging from 1.2 (Cambeva atrobrunnea Costa, Feltrin & Katz, sp. nov., Cambeva cubataonis (Bizerril, 1994), and Cambeva ventropapillata Costa, Feltrin & Katz, 2022) and 4.7.

Figure 10.

Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov., holotype, UFRJ 14070, 78.0 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Figure 11.

Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov., paratype, UFRJ 13820, 44.0 mm SL. A. Lateral view; B. Dorsal view; C. Ventral view.

Description

Morphometric data appear in Table 6.

Table 6.
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Morphometric data of Cambeva rotundipinna Costa, Feltrin & Katz, sp. nov.

Holotype Paratypes (n = 3)
Standard length (SL) 78.0 44.4–76.3
Percentage of standard length
Body depth 15.1 14.5–15.6
Caudal peduncle depth 12.1 11.9–13.0
Body width 12.0 12.2–13.3
Caudal peduncle width 4.1 3.7–4.9
Pre-dorsal length 64.6 65.3–69.7
Pre-pelvic length 57.2 59.9–60.9
Dorsal-fin base length 11.5 11.4–12.2
Anal-fin base length 8.2 8.2–9.1
Caudal-fin length 14.0 14.8–16.6
Pectoral-fin length 9.4 9.6–12.6
Pelvic-fin length 8.2 8.4–9.4
Head length 19.9 21.6–25.9
Percentage of head length
Head depth 50.7 40.7–52.7
Head width 85.3 68.6–80.1
Snout length 39.6 35.6–41.6
Interorbital width 22.2 18.7–23.7
Preorbital length 13.8 9.5–13.3
Eye diameter 8.3 7.9–10.2

Head morphology. Barbels moderate in length. Nasal barbel reaching area just posterior to orbit, maxillary and rictal barbels reaching middle of interopercular patch of odontodes. Jaw teeth with pointed to rounded extremities, irregularly arranged. Premaxillary teeth 37 or 38, dentary teeth 32–35 Opercular and interopercular odontodes pointed, about straight. Opercular odontodes 14–17, interopercular odontodes 30–34.

Fin morphology. Fins rounded. Total dorsal-fin rays 11 (ii + II + 7), total anal-fin rays 9 (ii + II + 5). Anal-fin origin at vertical through posterior portion of dorsal-fin base, at base of 6th branched dorsal-fin ray in specimens above about 50 mm SL, at vertical through posterior middle of dorsal-fin base, at base of 4th branched dorsal-fin ray in smaller specimens. Pectoral fin rounded in dorsal view, first pectoral-fin ray about equal in length to second ray, not forming terminal filament. Total pectoral-fin rays 7 (I + 6). Pelvic fin rounded, its tip reaching vertical through middle of dorsal-fin base. Total pelvic-fin rays 5 (I + 4). Caudal fin subtruncate, corners rounded. Total principal caudal-fin rays 13 (I + 11 + I), total dorsal procurrent rays 15–17 (xiv–xvi + I), total ventral procurrent rays 10 or 11 (ix–x + I).

Osteology (Fig. 4J–L). Mesethmoid broad anteriorly, with small lateral expansion in area just anterior to lateral ethmoid. Anterior mesethmoid margin convex, mesethmoid cornu broad, subtriangular, slightly curved posteriorly, abruptly narrowing at its extremity. Lateral ethmoid with small lateral projection immediately posterior to articular facet for autopalatine. Lacrimal thin, elliptical. Sesamoid supraorbital length about two times longer than lacrimal, without lateral expansions, its width about equal to lacrimal width. Premaxilla long, laterally narrowing, slightly curved. Maxilla slender, slightly curved, its length about four fifths of premaxilla, posterior process rudimentary. Autopalatine sub-trapezoidal in dorsal view, broad, its largest width about equal to its length, medial margin deeply sinuous with pronounced expansion on posterior margin, lateral margin weakly concave. Autopalatine postero-lateral process triangular, short, its length about half autopalatine length.

Metapterygoid sub-trapezoidal, longer than deep, relatively large, its surface greater than quadrate lateral surface. Area anterior to articulation between metapterygoid and quadrate with small laminar overlapped expansions of both bones. Quadrate with deep anterior constriction at dorsal process base. Hyomandibula long, anterior outgrowth horizontal length slightly longer than largest horizontal metapterygoid length. Dorsal margin of hyomandibula outgrowth concave. Opercle elongate, longer than interopercle. Opercular odontode patch moderately slender, its depth about two thirds of hyomandibula articular facet length. Dorsal process of opercle short, subtriangular, its extremity pointed. Opercular articular facet for hyomandibula with dorsal, small, rounded laminar projection, articular facet for preopercle rudimentary. Interopercle moderate in length, interopercular odontode patch length about equal hyomandibula outgrowth length. Preopercle slender, with minute ventral projection.

Parurohyal robust, lateral process sub-rectangular, slightly curved posteriorly, with truncate extremity. Parurohyal head with prominent anterolateral paired process. Parurohyal middle foramen relatively large, oval. Parurohyal posterior process short, about one third of distance between anterior margin of parurohyal and anterior insertion of posterior process. Branchiostegal rays 8 or 9. Vertebrae 39. Ribs 14 or 15. Dorsal-fin origin at vertical through centrum of 21st or 22nd vertebra; anal-fin origin at vertical through centrum of 25th or 26th vertebra. Two dorsal and single ventral hypural plate.

Colouration in alcohol. In adult specimens, above about 50 mm SL (Fig. 10), flank, dorsum and head side light brownish yellow, with great concentration of small dark brown to black dots. Venter and ventral surface of head pale yellow, with minute dark brown dots in area just anterior to pelvic fin and on branchiostegal region. Nasal and maxillary barbels brown, rictal barbel brownish white. Fins hyaline; dark chromatophores scattered over all fins, except pelvic fin. In juvenile specimens about 40 mm SL or less (Fig. 11), flank, dorsum and head side light brownish yellow with small black dots irregularly arranged.

Distribution

Cambeva rotundipinna is known from two close localities in the Rio da Paca, a tributary of the Rio Jangada, Rio Iguaçu drainage, Rio Paraná basin, at about 1000 m asl (Fig. 5).

Etymology

From the Latin rotundus (rounded) and pinna (fin or wing), an allusion to the rounded fins of this new species.

Discussion

Phylogenetic analyses indicated that the four species here described found at high altitudes do not form a monophyletic group, thus not supporting a common origin, but, in contrast, supported relationships of RISE species at different nodes of the phylogenetic tree (Fig. 1). On the other hand, all the four species are supported as members of the Cambeva beta1-clade, with C. galactica, endemic to eastern RISE as the sister group to all other species of this clade (Fig. 1). In C. galactica, there are six pectoral-fin rays, characteristic that never occurs in other species Cambeva beta1-clade (i.e. seven or eight pectoral-fin rays) but is present in species with a more basal position within the more inclusive Cambeva beta-clade, such as C. flavopicta Costa, Feltrin & Katz, 2020, C. naipi, and C. taroba, possibly consisting of a plesiomorphic feature for the Cambeva beta1-clade.

The position of C. atrobrunnea, C. luteoreticulata, and C. rotundipinna in a phylogenetic tree section with low resolution does not allow us to have an accurate view of their closest relationships. They appear as basal taxa relative to an apical clade including C. barbosae, C. diabola, C. castroi and C. davisi (Fig. 1, hereafter Cambeva davisi group), which are separated from each other by short genetic distances. Whereas C. atrobrunnea, C. luteoreticulata, and C. rotundipinna were only found at higher altitudes, between about 970 and 1,015 m asl, species of the C. davisi group were found at low and middle altitudes, between about 15 and 850 m asl (e.g. Bockmann et al. 2004; Costa et al. 2021b). The phylogenetic position of C. atrobrunnea, C. luteoreticulata, and C. rotundipinna relative to the C. davisi group suggests an older origin of those species living in higher altitudes. However, more robust phylogenies with the inclusion of yet undescribed species from neighbouring regions and species already described but not available for molecular analyses are necessary to infer the evolutionary history of the Cambeva beta-clade in southern Brazilian mountains.

Cambeva rotundipinna appears weakly supported as the sister group to C. stawiarski, but it was not possible to identify morphological characters corroborating relationships between these two species when examining comparative material of C. stawiarski (UFRJ 11846, 4 ex., 25°30'33"S, 53°40'59"W; UFRJ 11847, 3 ex., UFRJ 13620, 1 ex (C&S), 25°30'33"S, 53°40'59"W; UFRJ 11850, 2 ex. (C&S), about 25°40'S, 52°00'W; UFRJ 13541, 3 ex., 26°20'11"S, 49°32'18"W). However, the mesethmoidal region of C. rotundipinna bears great similarity to that illustrated for C. cauim, a species difficult to distinguish from C. stawiarski by external morphological characters and probably closely related to it (Reis et al. 2021). Cambeva cauim, also endemic to the Iguaçu River drainage (Reis et al. 2021), was not included in our phylogenetic analysis and specimens were not available for morphological examination, but its detailed original description allows us to make accurate comparisons. In both C. rotundipinna and C. cauim, the mesethmoid is wide in its anterior portion, with the cornua being subtriangular and slightly curved posteriorly, narrowing abruptly at its ends, and the autopalatine has a pronounced expansion on its postero-medial margin (Fig. 4J; Reis et al. 2021: fig. 2A), besides the urohyal having relatively short lateral and posterior processes (Fig. 4L; Reis et al. 2021: fig. 7A). Cambeva rotundipinna is easily distinguished from C. cauim by having a rounded caudal fin (vs. emarginate) and 15–17 dorsal procurrent caudal-fin rays (vs. 30 or 31), in addition to the unique diagnostic character states above described for C. rotundipinna. Cambeva atrobrunnea was weakly supported as sister to the Cambeva davisi group. However, in the present comparative morphological analysis, it was not possible to find any evidence of a close relationship between C. atrobrunnea and species of this group.

Acknowledgements

We are grateful to Luis E. H. Sebben and family for help and support during field studies and to Diego da Silva for sending comparative material. Special thanks are also due to M. Petrungaro, L. I. Chaves, B. R. dos Santos, M. R. dos Santos, and L. Neves for technical assistance in the fish collection. This study was funded by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; grant 304755/2020-6 to WJEMC) and Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ; grant E-26/201.213/2021 to WJEMC, E-26/203.524/2023 to JLOM; and E-26/202.005/2020 to AMK). This study was also supported by CAPES (Finance Code 001) through the Programa de Pós-Graduação em: Biodiversidade e Biologia Evolutiva/UFRJ and Genética/UFRJ.

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