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Research Article
The ornate rubbernose pleco (Siluriformes, Loricariidae, Chaetostoma), a new species from the Ucayali River Basin, Peru
expand article infoVanessa Meza-Vargas, Jorge L. Ramirez, Nathan K. Lujan§|
‡ Universidad Nacional Mayor de San Marcos, Lima, Peru
§ University of Toronto, Toronto, Canada
| Royal Ontario Museum, Toronto, Canada
Open Access

Abstract

A new species in the rubbernose catfish genus Chaetostoma is described from the Aguaytia, Pisqui and Palcazu Rivers, which drain the Pampa de Sacramento Region in the Ucayali River drainage of central Peru. The new species is distinguished from congeners, except C. anomalum, C. branickii, C. dorsale, C. leucomelas, C. microps, C. nudirostre, C. palmeri and C. thomsoni by having distinct, white, variably-shaped spots or vermiculations ½–2× nostril diameter on dark grey to black background on the head (vs. spots absent or black on light-coloured background). The new species is distinguished from C. anomalum, C. branickii, C. dorsale, C. microps, C. nudirostre and C. thomsoni by having highly variable, distinct white spots, vermiculations or bands ½–10× nostril diameter on the body, from C. leucomelas by having dorsal and caudal fin indistinctly and variably-patterned with zero to four bands (vs. dorsal and caudal fin consistently having five or more uniform bands) and from C. palmeri by having two predorsal plates (vs. three), supraoccipital excrescence present (vs. absent) and pelvic-fin insertion slightly posterior to dorsal-fin insertion (vs. pelvic-fin insertion at middle of dorsal-fin base). Species delimitation analyses of the COI and Cytb genes further support the recognition of this new species.

Key Words

Amazon, Andes, Chaetostoma clade, freshwater, molecular, Neotropical, taxonomy

Introduction

With over 1060 currently valid species, 19% of which (200) were described in the last decade, the suckermouth armoured catfish family Loricariidae is the fifth largest vertebrate family and one of the fastest growing (Fricke et al. 2024). Of the four vertebrate families that are more species-rich (Characidae, Cichlidae, Cyprinidae and Gobiidae), none is growing as quickly or is as geographically restricted as Loricariidae, which naturally occurs only in fresh or estuarine waters from southern Costa Rica to northern Argentina. Within Loricariidae, the rubbernose catfish genus Chaetostoma Tschudi 1846 is the third most species-rich genus with 49 valid species (Meza-Vargas et al. 2022), being superseded only by Hypostomus (> 150 valid species) and Ancistrus (76 valid species) (de Queiroz et al. 2020; Neuhaus et al. 2023). Chaetostoma is also the most geographically restricted of these species-rich genera, with 46 species being restricted to flanks of the Andes and Caribbean Coastal mountains from Panama to southern Peru and three species (C. jegui, C. orientale and C. vasquezi) occurring in rivers draining the Guiana or Brazilian shields (Meza-Vargas et al. 2022).

All but one species of Chaetostoma (C. platyrhynchus) are externally distinguished from most other loricariid genera by having the anterior and anterolateral snout margins free of plates, with this region instead being covered by a broad band of naked (i.e. unplated) skin, lacking tentacles. Five other loricariid genera also have unplated snout margins (Ancistrus Kner, 1854, Corumbataia (Britski, 1997), Paulasquama Armbruster & Taphorn, 2011, Soromonichthys Lujan & Armbruster, 2011 and Transancistrus Lujan, Meza-Vargas & Barriga-Salazar, 2015), but only Ancistrus and Transancistrus are sympatric with Chaetostoma. Chaetostoma can be easily distinguished from Ancistrus by lacking snout tentacles (vs. tentacles present), having at least five longitudinal series of plates at the shallowest part of the caudal peduncle (vs. three) and having eight or more branched dorsal-fin rays (vs. seven). The only Chaetostoma with a fully-plated snout (C. platyrhynchus) is restricted to the upper Caqueta and Napo drainages of Colombia and Ecuador, respectively; it shares all other traits above with congeners.

Chaetostoma has never undergone a comprehensive species-level taxonomic revision, though Lujan et al. (2015b) reviewed species from the centre of the genus’s geographic range in and around Ecuador and provided a multi-locus molecular phylogenetic hypothesis spanning many species and the entire geographic range of the genus. This analysis was recently enlarged and updated by Meza-Vargas et al. (2022). Both studies used combinations of up to 18 external morphological traits including colouration, meristics and morphometrics to distinguish amongst 13 species from rivers draining east of the Andes (cis-Andean), making taxonomy of the cis-Andean distribution of Chaetostoma the most well-studied (Salcedo 2006a, 2006b; Salcedo and Ortega 2015; Ballen et al. 2016). Within this distribution, the Huallaga River in central Peru is the epicentre of species diversity, with seven species currently recognised: C. changae, C. daidalmatos, C. lexa, C. marmorescens, C. stroumpoulos, C. taczanowskii, and C. trimaculineum. In the much larger Ucayali River drainage, which borders the Huallaga to the east and south, only two species are currently known: C. lineopunctatum and C. loborhynchos. In this paper, we describe a new, third species for the Ucayali River drainage. Seidel (2011) first reported this species as undescribed in the ornamental aquarium fish literature, simultaneously highlighting the species’ considerable colour pattern variation by assigning it three different L-numbers (L455, L456, L457). L-numbers are codes used by the ornamental aquarium fish industry to differentiate amongst and market distinct, often scientifically undescribed colour morphs (Stawikowski 1988).

In scientific literature, the species was first recognised as new by Lujan et al. (2015a), who referred to it as ‘Chaetostoma n.sp. Ucayali’ and included it in a multi-locus molecular phylogenetic analysis. This analysis found the new species to be part of a polytomy containing C. breve, C. dermorhynchus, C. dorsale, C. formosae, C. orientale (then Chaetostoma n.sp. Xingu), C. trimaculineum and an undescribed species from the Orinoco River Basin. In a re-analysis and expansion of the Lujan et al. (2015c) dataset, Meza-Vargas et al. (2022) found the new species to be sister to Chaetostoma breve Regan, 1904, a species whose nomenclature we revisit in light of these phylogenetic results and a large series of more recently collected topotype material for Chaetostoma branickii Steindachner, 1881.

Material and methods

Taxon sampling

This study is based mostly on 12 formalin-fixed, 70% ethanol-preserved, museum-catalogued specimens of the undescribed species collected from the Aguaytia River and the Raya and Omaiz Rivers in the Palcazu River drainage (Fig. 1) in August and September 2022 and on 313 specimens of Chaetostoma branickii collected in the Marañon River drainage in August 2018 (collection permit: Nº 00680-2022-PRODUCE/DGPCHDI). Specimens were euthanised prior to preservation by overdose of eugenol (clove oil) until ventilation stopped for at least 30 minutes in adherence with an institutional animal use protocol approved by Auburn University (protocol 2018–3354; Lucena et al. (2013)). Institutional codes follow Sabaj (2020).

Figure 1. 

Distribution of Chaetostoma sacramento type specimen sample sites along the Pampa del Sacramento in Andean left-bank tributaries of the Ucayali River Basin. Holotype (red star), green dots (paratypes), purple dots (aquarist trade).

Gene amplification and sequencing

Genetic samples of fin or muscle tissue were dissected prior to specimen fixation and individually linked to source specimens using unique, matching alphanumeric tags on voucher specimens and cryosafe tubes, then preserved in 95% ethanol. Whole genomic DNA was extracted from tissue subsamples using a salt precipitation protocol (Alijanabi and Martinez 1997). DNA sequence data were obtained for two mitochondrial markers: Cytochrome C oxidase I (COI) and Cytochrome b (CytB). A 658-base region of COI was amplified using the primers ANOSCOIF and ANOSCOIR (Ramirez and Galetti 2015) and a 1150-base region of the mitochondrial CytB gene was amplified using the primers and protocol of Lujan et al. (2015a). Amplified gene regions were sequenced on an ABI 3730 capillary sequencer at the Royal Ontario Museum (Toronto, Canada) or sent for sequencing at Macrogen Inc. (Seoul, South Korea).

Molecular analyses

Sequences for each gene were automatically aligned using Clustal W (Larkin et al. 2007) then manually edited and checked to ensure the absence of internal stop codons. De novo CytB sequences were compared with existing data for the Chaetostoma Clade from Lujan et al. (2015b) and additional COI sequences were obtained from BOLD (Table 1). Both gene datasets were used to run three species delimitation methods: the General Mixed Yule Coalescent model (GMYC) with a single threshold (Pons et al. 2006), the Bayesian implementation of the PTP model (bPTP) (Zhang et al. 2013) and the Assemble Species by Automating Partitioning (ASAP) method (Puillandre et al. 2021). As input for the first two methods, an ultrametric tree was built using BEAST v.2.7.4 (Bouckaert et al. 2014) with a GTR+I molecular evolution model for COI and an HKY+I+G model for CytB, a fast log-normal relaxed clock and a birth-death tree model. The analyses were run for 100 million Markov Chain Monte Carlo (MCMC) generations with the first 10% discarded as burn-in. Convergence of MCMC chains within a stable range and effective sample sizes (ESS) greater than 200 were confirmed in Tracer v.1.6 (Rambaut et al. 2014). The pipeline SPdel (Ramirez et al. 2023) was used to compute and compare all delimitation methods.

Table 1.

Tissue samples used in the molecular species delimitation analyses, including GenBank accession number (coI = Cytochrome oxidase subunit I, cytb = cytochrome b), voucher specimen catalogue number, country and drainages of origin. Bold accession numbers = new sequences for this study.

Taxa Tissues Number cytb coI Catalogue Number Country Drainage
Chaetostoma anale T24822 OL303592 ROM 107267 Colombia Orteguasa
Chaetostoma anale T24946 OL303593 OK514631 ROM 107837 Colombia Orteguasa
Chaetostoma anale T24906 OL303604 OK514628 ROM 107811 Colombia Caqueta
Chaetostoma anale T24907 OL303605 OK514629 ROM 107811 Colombia Caqueta
Chaetostoma anomalum T631 OL303608 INHS 55455 Venezuela Maracaibo
Chaetostoma bifurcum T13602 KP960196 ROM 93687 Ecuador Esmeraldas
Chaetostoma bifurcum T13603 OL303609 ROM 93687 Ecuador Esmeraldas
Chaetostoma bifurcum T13665 OL303610 ROM 93721 Ecuador Guayas
Chaetostoma bifurcum T13896 OL303611 ROM 93787 Ecuador Santa Rosa
Chaetostoma bifurcum T13897 OL303612 ROM 93787 Ecuador Santa Rosa
Chaetostoma breve P6292 KP960190 AUM 46515 Peru Marañon
Chaetostoma breve PE08208 OL303613 MHNG 2712.047 Peru Huallaga
Chaetostoma breve PE08213 OL303614 MHNG 2712.048 Peru Huallaga
Chaetostoma breve PE08648 OL303615 MHNG 2712.074 Peru Huallaga
Chaetostoma breve PE08673 OL303616 MHNG 2712.078 Peru Huallaga
Chaetostoma breve P6294 OL303617 AUM 46516 Peru Marañon
Chaetostoma breve T14360 OL303618 ROM 93950 Ecuador Napo S
Chaetostoma breve T19715 OL303619 ROM 100323 Ecuador Napo N
Chaetostoma breve T19716 OL303620 ROM 100320 Ecuador Napo N
Chaetostoma breve T14224 OL303621 ROM 93923 Ecuador Pastaza
Chaetostoma breve T14005 OL303622 ROM 93848 Ecuador Santiago
Chaetostoma brevilabiatum 6630 OL303623 ANSP 189597 Colombia Honda
Chaetostoma brevilabiatum 6631 OL303624 ANSP 189597 Colombia Honda
Chaetostoma cf. anale T19707 OL303594 ROM 100322 Ecuador Napo
Chaetostoma cf. anale T19708 OL303595 OK514634 ROM 100322 Ecuador Napo
Chaetostoma cf. anale T19606 OL303628 OK514632 ROM 100250 Ecuador Napo
Chaetostoma cf. anale T19647 OL303629 ROM 100263 Ecuador Napo
Chaetostoma cf. fischeri T9034 KP960197 STRI 11581 Panama Tuira
Chaetostoma cf. lineopunctatum CH153 OL303630 MUSM 44253 Peru Huallaga
Chaetostoma cf. lineopunctatum PE08318 OL303631 MHNG 2712.057 Peru Huallaga
Chaetostoma cf. lineopunctatum PE08545 OL303632 MHNG 2712.069 Peru Huallaga
Chaetostoma cf. loborhynchos CH200 OL303633 MUSM 44899 Peru Huallaga
Chaetostoma cf. loborhynchos CH202 OL303634 MUSM 44899 Peru Huallaga
Chaetostoma cf. loborhynchos CH2023 OL303635 MUSM 44899 Peru Huallaga
Chaetostoma cf. loborhynchos CH204 OL303636 MUSM 44899 Peru Huallaga
Chaetostoma changae PE08442 OL303637 MHNG 2712.064 Peru Huallaga
Chaetostoma changae PE08543 OL303638 MHNG 2712.067 Peru Huallaga
Chaetostoma chimu TICT-FCA-07 FBCH009-21 Colombia Meta
Chaetostoma chimu TICT-FCA-223 FBCH118-21 Colombia Meta
Chaetostoma chimu TICT-FCA-148 FBCH143-21 Colombia Meta
Chaetostoma daidalmatos CH140 OL303639 MUSM 44845 Peru Huallaga
Chaetostoma daidalmatos CH141 OL303640 MUSM 44845 Peru Huallaga
Chaetostoma daidalmatos PE08191 OL303641 MHNG 2712.045 Peru Huallaga
Chaetostoma daidalmatos PE08207 OL303642 MHNG 2712.051 Peru Huallaga
Chaetostoma daidalmatos PE08347 OL303644 MHNG 2712.055 Peru Huallaga
Chaetostoma dermorhynchus T14258 KP960191 ROM 93656 Ecuador Pastaza
Chaetostoma dermorhynchus T14287 OL303647 ROM 93656 Ecuador Pastaza
Chaetostoma dermorhynchus T14293 OL303648 ROM 93946 Ecuador Bobonaza
Chaetostoma dermorhynchus T14296 OL303649 ROM 93946 Ecuador Bobonaza
Chaetostoma dermorhynchus T14308 OL303650 ROM 93946 Ecuador Bobonaza
Chaetostoma dorsale TICT-FCA-08 FBCH008-21 Colombia Meta
Chaetostoma dorsale TICT-FCA-168 FBCH078-21 Colombia Meta
Chaetostoma dorsale TICT-FCA-185 FBCH086-21 Colombia Meta
Chaetostoma dorsale TICT-FCA-244 FBCH091-21 Colombia Meta
Chaetostoma fischeri T9026 KP960195 STRI 7604 Panama Chagres
Chaetostoma fischeri T9027 OL303651 STRI 905 Panama Bayano
Chaetostoma fischeri T9036 OL303652 STRI 12274 Panama Bayano
Chaetostoma fischeri T9037 OL303653 STRI Panama Chagres
Chaetostoma fischeri STRI-01740 BSFFA489-07 Panama Tuira
Chaetostoma fischeri STRI-01732 BSFFA490-07 Panama Acla
Chaetostoma fischeri STRI-01735 BSFFA491-07 Panama Bayano
Chaetostoma fischeri STRI-01754 BSFFA492-07 Panama Tuira
Chaetostoma fischeri STRI-01760 BSFFA493-07 Panama Bayano
Chaetostoma fischeri STRI-07104 BSFFA735-07 Panama Chagres
Chaetostoma formosae T17431 OL303654 ROM 95260 Colombia Meta
Chaetostoma formosae TICT-FCA-64 FBCH052-21 Colombia Meta
Chaetostoma guairense VZ122 OL303655 INHS 34786 Venezuela Limon
Chaetostoma jegui Tec63785 OL303596 OK514635 LBP 15478 Brazil Branco
Chaetostoma jegui Tec63786 OL303597 OK514636 LBP 15478 Brazil Branco
Chaetostoma joropo T17428 OL303656 ROM 95259 Colombia Meta
Chaetostoma joropo T17429 OL303657 ROM 95259 Colombia Meta
Chaetostoma joropo TICT-FCA-208 FBCH134-21 Colombia Meta
Chaetostoma joropo TICT-FCA-224 FBCH136-21 Colombia Meta
Chaetostoma joropo TICT-FCA-245 FBCH137-21 Colombia Meta
Chaetostoma leucomelas CIUA550-20 Colombia Cauca
Chaetostoma leucomelas CIUA557-20 Colombia Cauca
Chaetostoma leucomelas CIUA-7439 UDEA074-18 Colombia Cauca
Chaetostoma leucomelas CIUA-7441 UDEA075-18 Colombia Cauca
Chaetostoma leucomelas CIUA-7482 UDEA076-18 Colombia Magdalena
Chaetostoma leucomelas CIUA-7491 UDEA077-18 Colombia Magdalena
Chaetostoma lineopunctatum P4772 GU569899 EU359409 ANSP 180446 Peru Urubamba
Chaetostoma lineopunctatum P4814 GU569913 EU359410 ANSP 180448 Peru Madre de Dios
Chaetostoma lineopunctatum PE08047 KP960199 MHNG 2712.041 Peru Ucayali
Chaetostoma lineopunctatum T10088 OL303658 AUM 51166 Peru Madre de Dios
Chaetostoma lineopunctatum T10089 OL303659 AUM 51166 Peru Madre de Dios
Chaetostoma loborhynchos TK70561 KJ947862 MUSM 20291 Peru Tulumayo
Chaetostoma loborhynchos TK70563 KJ947864 MUSM 20291 Peru Tulumayo
Chaetostoma loborhynchos TK70579 KJ947865 MUSM 20307 Peru Paucartambo
Chaetostoma loborhynchos TK70580 KJ947866 MUSM 20307 Peru Paucartambo
Chaetostoma loborhynchos TK70581 KJ947867 MUSM 20307 Peru Paucartambo
Chaetostoma marmorescens CH198 KP960194 MUSM 44898 Peru Huallaga
Chaetostoma marmorescens CH197 OL303660 MUSM 44898 Peru Huallaga
Chaetostoma marmorescens CH199 OL303661 MUSM 44898 Peru Huallaga
Chaetostoma microps T14125 KP960189 ROM 93895 Ecuador Santiago
Chaetostoma microps CH121 OL303662 MUSM 44313 Peru Huallaga
Chaetostoma microps CH149 OL303665 MUSM 44869 Peru Huallaga
Chaetostoma microps CH150 OL303666 MUSM 44869 Peru Huallaga
Chaetostoma microps PE08190 OL303667 MHNG 2712.046 Peru Huallaga
Chaetostoma microps PE08580 OL303668 MHNG 2712.07 Peru Huallaga
Chaetostoma microps P6034 OL303669 AUM 45518 Peru Marañon
Chaetostoma microps T14364 OL303670 ROM 93949 Ecuador Napo S
Chaetostoma microps MEPN1255 OL303671 MEPN 1255 Ecuador Napo S cave
Chaetostoma microps MEPN1259 OL303672 MEPN 19165 Ecuador Napo S cave
Chaetostoma microps ROM 93902 OL303673 ROM 93902 Ecuador Santiago
Chaetostoma microps T14096 OL303674 ROM 93877 Ecuador Yungantza
Chaetostoma microps T14097 OL303675 ROM 93877 Ecuador Yungantza
Chaetostoma milesi JAM1984 OL303677 MPUJ Colombia Suaza
Chaetostoma milesi CIUA-8907 UDEA079-18 Colombia Magdalena
Chaetostoma milesi T24605 OL303676 ROM 106963 Colombia Suaza
Chaetostoma milesi CIUA680-20 Colombia Magdalena
Chaetostoma nudirostre T2084 OL303682 ANSP 191471 Venezuela Valencia
Chaetostoma orientale B1464 OL303679 OK514638 ANSP 199686 Brazil Xingu
Chaetostoma orientale B1472 OL303680 OK514639 ANSP 199686 Brazil Xingu
Chaetostoma orientale B1487 OL303681 OK514640 ANSP 199686 Brazil Xingu
Chaetostoma orientale 2606 OK514647 INPA-ICT 58146 Brazil Xingu
Chaetostoma sacramento sp. nov. PE08121 OL303678 MHNG 2712.042 Peru Ucayali
Chaetostoma sacramento sp. nov. LGBBF358 OR875871 OR859576 MUSM 72046 Peru Ucayali
Chaetostoma sacramento sp. nov. LGBBF359 OR875872 OR859577 MUSM 72046 Peru Ucayali
Chaetostoma sacramento sp. nov. MUSMT1564 OR859578 MUSM 71392 Peru Ucayali
Chaetostoma sacramento sp. nov. MUSMT1566 OR859579 MUSM 71392 Peru Ucayali
Chaetostoma sp. Apure T08954 OL303683 AUM 54034 Venezuela Apure
Chaetostoma sp. Apure T08955 OL303684 AUM 54034 Venezuela Apure
Chaetostoma sp. Apure T491 OL303685 AUM 41073 Venezuela Apure
Chaetostoma sp. Apure T621 OL303686 INHS 56147 Venezuela Apure
Chaetostoma sp. CuruaUna 525 OL303599 OK514641 MCP 53005 Brazil Curua-Una
Chaetostoma sp. L445 T12930 KP960193 ROM 94925 Colombia Meta
Chaetostoma sp. L445 T17424 OL303687 ROM 95257 Colombia Meta
Chaetostoma sp. L445 T17425 OL303688 ROM 95257 Colombia Meta
Chaetostoma sp. Meta CIUA698-20 Colombia Cauca
Chaetostoma sp. Meta CIUA707-20 Colombia Cauca
Chaetostoma stroumpoulos CH120 OL303689 MUSM 44303 Peru Huallaga
Chaetostoma stroumpoulos CH142 OL303691 MUSM 44847 Peru Huallaga
Chaetostoma stroumpoulos PE08210 OL303693 MHNG 2712.049 Peru Huallaga
Chaetostoma thomsoni JAM1966 OL303695 ICNMHN 17768 Colombia Chucuri
Chaetostoma thomsoni CIUA541-20 Colombia Cauca
Chaetostoma thomsoni CIUA542-20 Colombia Cauca
Chaetostoma thomsoni CIUA543-20 Colombia Cauca
Chaetostoma thomsoni CIUA544-20 Colombia Cauca
Chaetostoma thomsoni CIUA545-20 Colombia Cauca
Chaetostoma thomsoni CIUA546-20 Colombia Cauca
Chaetostoma thomsoni CIUA547-20 Colombia Cauca
Chaetostoma thomsoni CIUA553-20 Colombia Cauca
Chaetostoma thomsoni CIUA554-20 Colombia Magdalena
Chaetostoma thomsoni CIUA559-20 Colombia Cauca
Chaetostoma thomsoni CIUA560-20 Colombia Cauca
Chaetostoma thomsoni CIUA563-20 Colombia Cauca
Chaetostoma thomsoni CIUA565-20 Colombia Cauca
Chaetostoma thomsoni CIUA566-20 Colombia Magdalena
Chaetostoma thomsoni CIUA570-20 Colombia Cauca
Chaetostoma thomsoni CIUA572-20 Colombia Cauca
Chaetostoma thomsoni CIUA574-20 Colombia Cauca
Chaetostoma thomsoni CIUA576-20 Colombia Cauca
Chaetostoma thomsoni CIUA579-20 Colombia Cauca
Chaetostoma thomsoni CIUA580-20 Colombia Cauca
Chaetostoma thomsoni UDEA080-18 Colombia Cauca
Chaetostoma thomsoni UDEA083-18 Colombia Cauca
Chaetostoma thomsoni UDEA085-18 Colombia Cauca
Chaetostoma trimaculineum P6047 OL303696 AUM 45524 Peru Marañon
Chaetostoma trimaculineum T14136 OL303697 ROM 93894 Ecuador Santiago
Chaetostoma vasquezi T09945 KP960192 AUM 53812 Venezuela Caura
Chaetostoma vasquezi V27 OL303698 OK514643 AUM 36555 Venezuela Caroni
Chaetostoma vasquezi V28 OL303699 OK514644 AUM 36555 Venezuela Caroni

Morphological analysis

Morphometric landmarks follow Armbruster (2003) with lateral trunk plate row terminology following Schaefer (1997) and cheek odontode type following Lujan et al. (2015b). Standard length (SL) is expressed in mm and other measurements are expressed as percentages of either standard length or, for subunits of the head, head length. Measurements and counts were taken on the left side of specimens when possible.

Results

Taxonomic accounts

Chaetostoma sacramento sp. nov.

Fig. 2, Tables 2, 3

Chaetostoma sp. nov. Ucayali: Lujan et al. (2015b) [molecular phylogeny].

Type material

Holotype • Adult MUSM 72045; 65.1 mm SL; Peru, Ucayali Department, Padre Abad Province, Boqueron District, Shambillo, Amazon Basin, Ucayali River, unnamed left-bank tributary of Aguaytia River; 09°0'29.88"S, 75°37'1.92"W; alt. 365 m; 07 Aug 2022; D. Faustino, J. Chuctaya, C. Nolasco, O. Quispe.

Paratype : All Peru, Amazon – Ucayali River Basin • MHNG 2712.042, 1; 32.2 mm SL; tissues PE08-122, 123, 124; Padre Abad Province, Aguaytia River at mouth of the Boca Yurac River; 11 September 2008; S. Fisch- Muller, R. Covain, P. de Rham, H. Ortega, J. Figuerosa Minaya, J. Sanchez Ramirez • MUSM 71392, 4; 50.5–79.4 mm SL; Pasco Department, Oxapampa Province, Palcazu District, Raya River; 10°22'13.61"S, 75°7'52.90"W; 7 September 2022; R. Olivera, R. Quispe, J. Arana, M. Paniagua • MUSM 72046, 34; 44.2–12.5 mm SL; Ucayali Department, Padre Abad Province, Padre Abad District, Aguaytia River; 9°4'8.58"S, 75°30'51.48"W; 8 August 2022; D. Faustino, J. Chuctaya, C. Nolasco, R. Quispe • ANSP 182805, 3; 57.8–70.5 mm SL; same data as MUSM 71392 • ROM 114668, 4; 54.3–76.7 mm SL; same data as MUSM 71392.

Genseq-2 COI

GenBank accession number. Obtained from paratypes MUSM 72046 (OR859576 and OR859577) and MUSM 71392 (OR859578 and OR859579).

Genseq-2 CytB

GenBank accession number. Obtained from paratypes MUSM 72046 (OR875871 and OR875872).

Diagnosis

Chaetostoma sacramento can be diagnosed from all congeners, except C. anomalum, C. branickii, C. dorsale, C. leucomelas, C. microps, C. nudirostre, C. palmeri and C. thomsoni by having distinct, white, variably-shaped spots or vermiculations ½–2× nostril diameter on grey to brown background on the head (vs. spots absent or black on light-coloured background). Chaetostoma sacramento is distinguished from C. anomalum, C. branickii, C. dorsale, C. nudirostre and C. thomsonii by having highly variable, small to large distinct white spots, vermiculations or bands on the body (vs. spots, vermiculations or bands absent or black on light-coloured background), from C. anomalum by having adipose fin fully formed (vs. rudimentary), from C. dorsale by having uniformly brown adipose fin (vs. adipose fin with black spot), from C. leucomelas by having golden spots across the dorsal fin rays (vs. light bands), from C. microps by having eight branched dorsal-fin rays (vs. nine), from C. nudirostre by having curved cheek odontodes (vs. strait), from C. palmeri by having two predorsal plates (vs. three), excrescence present (vs. absent) and pelvic-fin insertion slightly posterior of dorsal-fin insertion (vs. pelvic-fin insertion at middle of dorsal-fin base).

Description

Morphometric data in Table 2 and meristic data in Table 3. Snout moderately depressed, dorsal profile of head convex from snout tip to dorsal-fin origin, then straight and gradually descending to adipose fin. Caudal peduncle dorsal profile slightly concave. Ventral profile straight from snout to caudal fin. Body depressed, deepest at dorsal-fin origin, shallowest at caudal peduncle; greatest width at pectoral girdle. Caudal peduncle slightly compressed, roundly triangular in cross-section.

Table 2.

Morphometrics of Chaetostoma sacramento. Morphometric values as percentages of standard length (SL) or head length (HL). H = holotype, SD = standard deviation, (n = 11).

H Min Max Mean SD
Standard length (mm) 65.13 50.49 76.74 63.7
Percent of standard length
Head length 33.9 34.6 37.6 36.0 1.0
Predorsal length 43.4 41.9 47.3 45.1 1.5
Head-dorsal length 9.8 8.7 11.1 9.9 0.7
Cleithral width 33.6 29.3 33.4 32.0 1.2
Cleithral widest distance 37.1 34.8 36.5 35.7 0.5
Head-pectoral length 31.2 30.1 34.3 31.8 1.3
Thorax length 21.7 19.2 23.9 21.3 1.2
Pectoral-spine length 36.3 26.2 29.1 28.1 0.9
Abdominal length 25.4 23.0 25.5 24.5 0.7
Pelvic-spine length 25.7 24.8 28.5 26.4 1.0
Postanal length 30.2 29.1 34.8 32.0 1.4
Anal-fin spine length 15.0 8.7 10.4 9.8 0.5
Dorsal-pectoral distance 27.9 28.5 31.0 29.8 0.7
Dorsal spine length 26.7 25.5 29.4 27.8 1.0
Dorsal-pelvic distance 24.0 21.5 24.1 23.1 0.7
Dorsal-fin base length 28.7 25.0 26.4 25.5 0.4
Dorsal-fin base length 26.2 22.8 26.3 23.8 0.9
Dorsal-adipose distance 16.1 13.2 15.6 14.4 0.8
Adipose-spine length 8.2 7.7 9.4 8.2 0.5
Adipose-up. caudal distance 12.2 12.2 17.4 15.1 1.6
Caudal peduncle depth 15.2 12.4 13.9 13.1 0.6
Adipose-low. caudal distance 21.3 21.2 24.8 23.3 1.2
Adipose-anal distance 20.9 18.6 22.4 20.7 1.0
Dorsal-anal distance 17.1 15.6 17.0 16.3 0.5
Pelvic-dorsal distance 28.9 26.9 29.7 27.9 0.9
Percent of head length
Head-eye length 29.7 10.0 11.1 10.6 0.3
Orbit diameter 14.0 5.2 6.6 5.9 0.4
Snout length 71.4 22.7 24.9 23.6 0.7
Internares width 14.5 4.6 5.7 5.2 0.4
Interorbital width 45.0 13.8 16.1 15.1 0.7
Head depth 71.9 24.7 26.3 25.6 0.5
Head width 95.6 33.3 35.6 34.5 0.9
Mouth length 60.3 21.1 23.5 22.2 0.8
Mouth width 81.3 29.5 33.2 30.9 1.0
Barbel length 12.3 2.5 4.4 3.6 0.6
Dentary tooth cup length 32.0 11.5 13.4 12.3 0.7
Premaxillary tooth cup length 27.1 10.4 11.6 11.0 0.3
Occipital length 54.1 17.9 20.0 19.1 0.8
Caudal peduncle length 24.3 24.5 28.2 26.8 1.3
Opercle length 9.7 3.3 5.0 4.0 0.5
Interbranchial distance 25.9 24.2 26.3 24.9 0.7
Table 3.

Meristics of Chaetostoma sacramento. H = holotype, SD = standard deviation, (n = 11).

Count H Min Max Mode SD
Median plates 24 22 24 23 0.5
Supramedian plates 23 22 24 24 0.7
Inframedian plates 24 24 25 24 0.5
Caudal plates rows 5 5 5 5 0.0
Dorsal-fin branched rays 8 8 8 8 0.0
Pectoral-fin branched rays 6 6 6 6 0.0
Pelvic-fin branched rays 5 5 5 5 0.0
Anal-fin branched rays 4 4 4 4 0.0
Caudal–fin branched rays 14 13 14 14 0.3
Dorsal procurrent caudal–fin rays 4 4 4 4 0.0
Ventral procurrent caudal–fin rays 4 2 4 3 0.6
Dorsal fin base plates 7 7 7 7 0.0
Preadipose plates 6 5 5 5 0.0
Adipose-caudal plates 5 6 6 6 0.0
Infraorbital plates 6 6 6 6 0.0
Left dentary teeth 186 82 214 104 40.3
Left premaxillary teeth 115 63 137 86 19.4
Cheek odontodes 4 3 5 5 0.6

Head wide, anteriorly rounded in dorsal view; snout anterior margin unplated, lacking odontodes and tentacles. Orbit small (5.2–6.6% HL), dorsolaterally positioned, posterior margin aligned with vertical through anterior margin of cleithrum; skull roof flat. Oral disc wide, elliptical, occupying most of head width, lower lip ending just anterior to origin of opercular opening. Oval papillae covering upper lip, roundish papillae covering lower lip; all papillae smaller towards outer lip margins; lower lip margin crenulate. Buccal cavity with digitate papillae present posterior to premaxillary symphysis, one large patch of rugose papillae dorsomedial to each dentary tooth row. Maxillary barbel short (2.5–4.4% HL). Premaxillary tooth row straight, joining contralateral tooth row at 160º angle. Dentary straight, joining contralateral tooth row at 170º angle. Teeth small, villiform, asymmetrically bicuspid. Opercle external border having row of six straight odontodes, odontodes slightly larger than elsewhere on body. Three to five type 3 hypertrophied hook-like evertible cheek odontodes (mode 5), odontodes hooked anteriorly, not reaching cleithrum. Supraoccipital excrescence restricted to vestigial longitudinal unplated patch, keel absent.

Figure 2. 

Live holotype of Chaetostoma sacramento MUSM 72045, 65.1 cm, collected from the Yamino River, a tributary of the Aguaytía River. Photos by D. Faustino.

Flanks covered by five longitudinal plate series. Plates absent from abdomen and around dorsal-fin base. Body plates flat or gently curved, lacking keel or ridge. Dorsal-fin base bordered laterally by seven dorsal plates. Medial interdorsal plates five. Dorsal plate series with 20 plates, supramedian with 24, median series with 23, mid-ventral series with 24, ventral series with 19.

Dorsal fin II,8; locking mechanism functional, spinelet V-shape, dorsal-fin origin slightly anterior to pelvic-fin origin, last dorsal-fin ray not reaching adipose fin when adpressed. Pectoral-fin rays I,6; pectoral-fin spine reaching first third of pelvic fin when adpressed. Pelvic-fin rays i,5; unbranched rays surpassing anal-fin origin; dorsal skin folds present on proximalmost two-thirds of unbranched pelvic-fin rays. Pectoral-fin spine with thicker odontodes irregularly distributed along entire dorsal, anterior and ventral surface of spine; one aligned row of larger odontodes along posterodorsal margin. Odontodes present on all, but posteriormost branched pectoral- and pelvic-fin rays. Anal-fin rays ii,4, first unbranched ray almost as long as second. Second branched anal-fin ray longest, with remaining rays successively shorter. Caudal fin obliquely forked, lower lobe longer than upper; i,7+7,i. Dorsal procurrent caudal-fin rays four, ventral two to four.

Colour in alcohol

Head and body base colour brown dorsally, yellowish-white ventrally. Head with dense small white dots. In some specimens, scattered white spots on the body may be fused. Supraoccipital excrescence black or grey. Dorsal fin with white scattered spots on brownish base composed of melanophores distributed on rays and membrane. Remaining fins with brownish base composed of scattered melanophores. Tip of unbranched dorsal- and caudal-fin rays whitish in some individuals.

Sexual dimorphism

Males with fleshy dorsal fold on the pelvic-fin leading ray. Males have more distinct, intense and contrasting white patterns on darker brown to black base colour. Females with duller colour patterns on lighter brown to black base colour.

Distribution

Chaetostoma sacramento is known exclusively from the Pampa de Sacramento valley east of the eastern cordillera of the Andes in Peru, inhabiting the Negro River, a tributary of the Pisqui River in Loreto Department; the Yamino River, a tributary of the Aguaytia River; the Chui River, a tributary of upper San Alejandro River in Ucayali Department; and the Raya River, a left-bank tributary of Iscozacin River in Pasco Department (Fig. 1).

Etymology

The species epithet sacramento refers to the plain (pampa) in central Peru between the Huallaga and Ucayali Rivers, approximately delimited by the Pisqui River in the north and Palcazu River in the south. Chaetostoma sacramento is currently known exclusively from this region, known as the Pampa de Sacramento, which occupies a valley between Huánuco and Ucayali provinces that is part of the Peruvian subandean belt and surrounds Boqueron del Padre Abad in the Cordillera Azul. The Pampa de Sacramento was first encountered by Europeans on 21 June 1726, by an expedition led by Don Juan Nunez Lobo and was christened Pampa del Sacramento to commemorate the Catholic ceremony of the Corpus Christi. Subsequent Franciscan missionaries highlighted the rich ethnic diversity of this region (IBC 2016). A noun in apposition.

Molecular species delimitation

With 626 total bases and 72 sequences from 12 nominal species, the COI dataset yielded 158 parsimony informative sites (25%). The ASAP species delimitation analysis yielded 20 Molecular Operational Taxonomic units (MOTUs) (score = 2.5), whereas GMYC and bPTP yielded 23 and 22 MOTUs, respectively. These results can be summarised with 22 consensus MOTUs (Fig. 3).

Figure 3. 

Results of species delimitation analyses, based on COI (left) and CytB (right) gene regions.

With 1048 total bases and 108 sequences from 24 nominal species, the CytB dataset yielded 355 parsimony informative sites (34%). ASAP (score = 8.0), GMYC and bPTP species delimitation analyses yielded 46, 45, 49 MOTUs, respectively, for a total of 46 consensus MOTUs (Fig. 3).

In all species delimitation analyses of both genes, Chaetostoma sacramento was identified as an MOTU distinct from all other examined Chaetostoma species. The minimum genetic distance to the nearest species was 6.15% for CytB, with C. branickii being the closest species. For COI, the closest species was C. leucomelas, with a genetic distance of 4.48%.

Chaetostoma branickii Steindachner 1881

Figs 4, 5

Chaetostoma breve Regan 1904 [synonym]

Diagnosis and description

As for Chaetostoma breve in Lujan et al. (2015a).

Comments

Regan (1904) based the description of Chaetostoma breve on specimens from the Zamora River in south-eastern Ecuador. Lujan et al. (2015a) re-described Chaetostoma breve based on specimens from throughout most of the species’ range, from the Napo River in the north (although the species is also now known from more northern Caqueta River headwaters in Colombia; ROM 107831, 107845) to the Marañon River in the south. Lujan et al. (2015a) also generated a multi-locus molecular phylogenetic hypothesis spanning over 24 valid and undescribed species, with Chaetostoma breve represented by samples from the Napo, Pastaza, Marañon, Santiago and Huallaga River Basins. Bayesian and Maximum Likelihood analyses found uniformly strong support for monophyly of the entire Chaetostoma breve clade (Bayesian posterior probability: 1.00, Maximum Likelihood bootstrap: 100).

Chaetostoma branickii was described by Steindachner (1881), based on specimens from the Chota River near Cajamarca in northern Peru. That portion of the Marañon River Basin was poorly sampled at the time of Lujan et al. (2015c), thus few specimens and no tissues from that specific region were available, making it reasonable to assume that a distinct, endemic species might exist there. Moreover, the syntypes of Chaetostoma branickii were in Vienna and not directly accessible to the authors. Lujan et al. (2015c) concluded that Chaetostoma branickii might be more closely allied with Chaetostoma taczanowskii, which was described by Steindachner (1881), based on specimens from the Huallaga River drainage to the south. To further investigate the validity of Chaetostoma branickii, Lujan and colleagues sampled the upper middle Marañon River in 2018, collecting over 540 fresh specimens and 60 tissues of Chaetostoma. This new material allowed a more detailed comparison of fresh, nearly topotypic specimens with the specimens on which Lujan et al. (2015a) based their re-description of Chaetostoma breve and type images of Chaetostoma branickii (Figs 4, 5). Based on this comparison, it has become clear that these are the same species and that Chaetostoma branickii should thus be recognised as the senior synonym of Chaetostoma breve.

Figure 4. 

Original illustrations of Chaetostoma branickii from Steindachner 1881.

Figure 5. 

Photo comparison of a nearly 150-year-old preserved syntype of Chaetostoma branickii (NMW 47270-71, photos by Mark Sabaj & Kyle Luckenbill, copyright NMW) with a live specimen collected in 2018 from the Marañon River, Cajamarca Province, Peru (ROM 109213, photos by NKL). Scale bars: 1 cm.

Chaetostoma taczanowskii Steindachner 1882 is another taxonomically ambiguous species from northern Peru, described from the Huallaga River Basin, a southern tributary of the Marañon. Type images of this species suggest that it is also closely related to Chaetostoma branickii, if not another junior synonym, but a robust evaluation of the status of this species must await availability of fresh topotypic specimens and tissues.

Discussion

Fourteen Chaetostoma species are currently recognised from Peruvian drainages, most of which (12) inhabit the Marañon River Basin, including the Huallaga, Pastaza and Napo Rivers, with only three now described from the Ucayali River Basin: C. lineopunctatum from the Pachitea River, C. loborhynchos from the Tulumayo Rivers and C. sacramento. Until now, the colour pattern described for most species of Chaetostoma in Peru comprises black spots on the body and head (C. daidalmatos, C. lineopunctatum, C. stroumpoulos, C. trimaculineum) or irregular black spots on the body alone (C. loborhynchos, C. spondylus). There are also six species described as having uniform or marbled colouration (C. branickii, C. changae, C. marmorescens, C. lexa and C. taczanowskii). White patterns on a darker base colour are observed in only two species in Peru, C. dermorhynchus and C. microps, from which C. sacramento differs by having distinct and highly variable small to large white spots on the body, head and fins (vs. white spots on only the dorsal fin in C. dermorhynchus) and by having eight dorsal-fin branched rays (vs. nine in C. microps). Our recognition of C. sacramento is further supported by our molecular results, which consistently identify it as a distinct MOTU using the mitochondrial genes COI and CytB. Additionally, our findings demonstrate that C. dermorhynchus and C. microps each represent a separate MOTU and are phylogenetically distant species to C. sacramento, as shown in Lujan et al (2015b).

One of the fundamental criticisms of DNA-based approaches to species delimitation is that such analyses are incapable of distinguishing between genetic structure due to population-level processes and that due to species boundaries (Sukumaran and Knowles 2017). We advocate for a conservative approach in which MOTUs are treated as candidate species to be recognised taxonomically only if they are also morphologically diagnosable. Thus, the combined molecular and morphological evidence supports our recognition of Chaetostoma sacramento as a distinct species. Alternatively, the molecular species delimitation analyses consistently subdivided the Chaetostoma branickii clade into six MOTUs that we do not recognise as distinct species because of the overall morphologically similarity of these subunits and the strong monophyly of the parent clade. Future research may yet yield morphological characteristics capable of distinguishing these populations, thereby supporting their recognition as distinct species. Such a potentiality would be unlikely to resurrect Chaetostoma breve, however, given the very close genetic relationship between individuals from the Santiago (breve) and Marañon (branickii) river type locality populations, consistent with our decision to recognise Chaetostoma branickii as the senior synonym for this species.

Although Chaetostoma sacramento has been known in the aquarium fish trade for almost 30 years, it was originally assigned three L-numbers in 2011 (L455, L456 and L457; Seidel (2011)) and is known in the aquarium trade most commonly as L455. Other common aquarium trade names for the species include ‘Tiger Chaetostoma’, ‘Tingo Red Chaetostoma’ or ‘Ornate Chaetostoma’. Specimens on which the present description is based were collected in the San Carlos River, part of the Palcazu River Basin in Yanachaga Chemillen National Park (PNYC) in 2007 and in the Aguaytia River in 2008. According to aquarium fish exporters, this species also occurs in the Negro River, a small tributary of the upper Pisqui River near the Boqueron del Padre Abad and in the Chiu River, a tributary of San Alejandro River (Seidel 2011).

Acknowledgements

We thank Dario Faustino, Junior Chuctaya, Claudia Nolasco, Omar Quispe, Jerry Arana and Miguel Paniagua for field assistance, Raphael Covain at MHNG for sharing specimens and data and Julian Dignall at PlanetCatfish.com for sharing photos, aquarium literature and his thoughts on L455. This work was financed by CONCYTEC through the PROCIENCIA programme within the framework of the contest “Projects for the incorporation of postdoctoral researchers in Peruvian institutions” according to contract [074-2021] and [PE501084299-2023-PROCIENCIA-BM] within the framework of the call “Interinstitutional Alliances for Doctorate Programs”.

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