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A new species of Harttia from the rio São Francisco basin (Siluriformes: Loricariidae)

Abstract

The genus Harttia belongs to the subfamily Loricariinae and has to date 27 described species, distributed in the drainages of Guiana Shield, Amazon and southeastern Brazil. The new species is distinguished from its congeners by the combination of: canal plate present; abdominal plates absent; trapezoid preanal plates; males with elongated and conspicuous odontodes on the first pectoral-fin ray and on the lateral region of the head, close to the canal plate; and dorsal-fin spinelet present. An identification key for the species of the rio São Francisco basin and a comparison between the new taxon and all other species of the genus are presented.

Keywords:
Brazilian Crystalline Shield; Brazilian Plateau; Identification key; Streams; Waterfalls.

Resumo

O gênero Harttia pertence à subfamília Loricariinae e possui 27 espécies descritas, distribuídas nas drenagens do Escudo das Guianas, Amazônica e do sudeste brasileiro. A nova espécie distingue-se das congêneres pela combinação de: placa do canal presente; placas abdominais ausentes; placas pré-anais trapezóides; machos com odontódeos alongados e conspícuos no primeiro raio da nadadeira peitoral e na região lateral da cabeça, próximo a placa do canal; e spinelet da nadadeira dorsal presente. Apresenta-se uma chave de identificação para as espécies da bacia do rio São Francisco e a comparação com todas as demais espécies do gênero.

Palavras-chave:
Cachoeiras; Chave de identificação; Escudo Cristalino Brasileiro; Planalto Brasileiro; Riachos

INTRODUCTION

The genus Harttia Steindachner, 1877 belongs to the subfamily Loricariinae, comprising 27 species (Fricke et al., 2022Fricke R, Eschmeyer WN, Van der Laan R. Eschmeyer’s catalog of fishes: genera, species, references [Internet]. San Francisco: California Academy of Science; 2022. Available from: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp
http://researcharchive.calacademy.org/re...
; Tab. 1), that inhabit the upper portions of medium and large rivers, with rocky or sandy bottoms, in the drainages of the rivers Doce, Jequitinhonha, Paraíba do Sul, Paraná, Ribeira de Iguape, São Francisco, Orinoco, rivers of Guiana Shield, and Amazon (Covain et al., 2012Covain R, Fisch-Müller S, Montoya-Burgos JI, Mol JH, Le Bail PY, Dray S. The Harttiini (Siluriformes, Loricariidae) from the Guianas: a multi-table approach to assess their diversity, evolution, and distribution. Cybium. 2012; 36(1):115–61.; Oyakawa et al., 2018Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
). Rapp Py-Daniel, (1997)Rapp Py-Daniel LH. Phylogeny of the neotropical armored catfishes of the subfamily Loricariinae (Siluriformes: Loricariidae). [PhD Thesis]. Tucson: The University of Arizona, EUA; 1997. diagnosed the genus as monophyletic by having: anterior internal process of the basipterygium oriented mesially and not connected to the contralateral element medially, bifurcation point of the infraorbital and supraorbital canals between the sphenotic and the compound pterotic, first ceratobranchial with a wide flap, parapophysis of the fourth vertebra not connected to the transcapular ligament, terminal exit of the parietal branch in the supraoccipital. Other authors have diagnosed the genus Harttia by having an elongated and flattened body, lateral keel of the body absent; large bony plates surrounding the anal papilla, and caudal fin emarginated (Oyakawa, 1993Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. ; Rapp Py-Daniel, Oliveira, 2001Rapp Py-Daniel LH, Oliveira EC. Seven New Species of Harttia from the Amazonian-Guyana Region (Siluriformes: Loricariidae). Ichthyol Explor Freshw. 2001; 12(1):79–96. ; Rapp Py-Daniel, Ohara, 2013Rapp Py-Daniel LH, Ohara WM. Loricariidae. In: Queiroz L, Torrente-Vilara G, Ohara W, Pires T, Zuanon J, Doria C, editors. Peixes do rio Madeira. São Paulo: Dialeto Latin America Documentary; 2013. p.225–301.).

More recently, Londoño-Burbano, Reis, (2021)Londoño-Burbano A, Reis RE. A combined molecular and morphological phylogeny of the Loricariinae (Siluriformes: Loricariidae), with emphasis on the Harttiini and Farlowellini. PLoS ONE. 2021; 16(3):e0247747. https://doi.org/10.1371/journal.pone.0247747
https://doi.org/10.1371/journal.pone.024...
in a combined molecular and morphological phylogeny of the Loricariinae tested all the synapomorphies proposed by Rapp Py-Daniel, (1997)Rapp Py-Daniel LH. Phylogeny of the neotropical armored catfishes of the subfamily Loricariinae (Siluriformes: Loricariidae). [PhD Thesis]. Tucson: The University of Arizona, EUA; 1997., and only one, the point of bifurcation of the infraorbital and supraorbital canals on the sphenotic (their character 37) resulted as a non-exclusive synapomorphy of Harttia, which was confirmed as monophyletic, corroborating Rapp Py-Daniel, (1997)Rapp Py-Daniel LH. Phylogeny of the neotropical armored catfishes of the subfamily Loricariinae (Siluriformes: Loricariidae). [PhD Thesis]. Tucson: The University of Arizona, EUA; 1997., Provenzano, (2011)Provenzano F. Estudio sobre las relaciones filogenéticas de las especies incluidas en la subfamilia Loricariinae (Siluriformes, Loricariidae). [PhD Thesis]. Universidad Central de Venezuela, Caracas. 2011. Available from: http://saber.ucv.ve/handle/123456789/1552
http://saber.ucv.ve/handle/123456789/155...
, and Covain et al., (2016)Covain R, Fisch-Müller S, Oliveira C, Mol JH, Montoya-Burgos JI, Dray S. Molecular phylogeny of the highly diversified catfish subfamily Loricariinae (Siluriformes, Loricariidae) reveals incongruences with morphological classification. Mol Phylogenet Evol. 2016; 94:492–517. http://dx.doi.org/10.1016/j.ympev.2015.10.018
http://dx.doi.org/10.1016/j.ympev.2015.1...
. Londoño-Burbano, Reis, (2021)Londoño-Burbano A, Reis RE. A combined molecular and morphological phylogeny of the Loricariinae (Siluriformes: Loricariidae), with emphasis on the Harttiini and Farlowellini. PLoS ONE. 2021; 16(3):e0247747. https://doi.org/10.1371/journal.pone.0247747
https://doi.org/10.1371/journal.pone.024...
also differentiate Harttia from other Loricariini by a combination of plates on caudal peduncle smooth, without keels; odontodes usually not well developed on sides of head or predorsal region; canal-bearing cheek plate with short, broad ventral process; caudal peduncle strongly depressed (vs. plates on caudal peduncle keeled, not smooth; odontodes well developed on sides of head or predorsal region; canal-bearing cheek plate with long and thin ventral process; caudal peduncle oval in cross-section in Harttiella Boeseman, 1971) and dark blotch at caudal-fin base; tip of snout naked, devoid of plates; abdominal plates absent or present as small platelets, partially or completely covering the abdomen (vs. dark transverse, half-moon shaped band at caudal-fin base, occupying the base of all rays of upper and lower lobes; tip of snout covered with plates; abdominal plates, as medium-sized polygonal plates completely covering the abdomen in Cteniloricaria Isbrücker & Nijssen, 1979).

Oyakawa et al., (2018)Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
divided the genus into three groups based on the coverage of plates in the abdominal region: 1) Harttia loricariformis group, whose members have a naked abdomen (without plates); 2) H. rhombocephala group, with abdomen partially covered by plates; and 3) H. fowleri group, with fully plated abdomen (for more information on the species composition of each group see Tab. 1).

To date, the rio São Francisco basin has the greatest diversity of the genus in southeastern Brazil, housing four species: Harttia leiopleura Oyakawa, 1993, H. longipinna Langeani, Oyakawa & Montoya-Burgos, 2001, H. novalimensisOyakawa, 1993Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. , and H. torrenticolaOyakawa, 1993Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. . Our analyses indicate the occurrence of a new species from the Serra da Canastra National Park region, already mentioned in the literature as Harttia sp. (Casatti, Castro, 1998Casatti L, Castro RMC. A fish community of the São Francisco River headwaters riffles, southeastern Brasil. Ichthyol Explor Freshw. 1998; 9(3):229–42.), Harttia cf. novalimensis (Romero, Casatti, 2012Romero RM, Casatti L. Identification of key microhabitats for fish assemblages in tropical Brazilian savanna streams. Int Rev Hydrobiol. 2012; 97(6):526–41. https://doi.org/10.1002/iroh.201111513
https://doi.org/10.1002/iroh.201111513...
; Roa-Fuentes et al., 2015Roa-Fuentes CA, Casatti L, Romero RM. Phylogenetic signal and major ecological shifts in the ecomorphological structure of stream fish in two river basins in Brazil. Neotrop Ichthyol. 2015; 13(1):165–78. https://doi.org/10.1590/1982-0224-20140045
https://doi.org/10.1590/1982-0224-201400...
), and H. novalimensis (Angulo, 2019Angulo A. Gross brain morphology of the Loricariinae (Siluriformes: Loricariidae): comparative anatomy, ecological implications and phylogenetic analysis. [PhD Thesis]. São José do Rio Preto: Universidade Estadual Paulista; 2019. Available from: http://hdl.handle.net/11449/182045
http://hdl.handle.net/11449/182045...
). The description of the new species, its comparison with all other species of the genus and an identification key for the species in the basin are presented here.

TABLE 1 |
Currently recognized species of Harttia presented in three groups based on abdominal covering according to Oyakawa et al., (2018)Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
and their type locality.

MATERIAL AND METHODS

Measurements and counts, taken under a stereomicroscope, follow Oyakawa et al., (2018)Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
. The morphometric data were taken with a digital caliper, from point-to-point, approximation of 0.1 mm, and on the left side of the specimens; central tendency measures, such as minimum and maximum values, mean (for measurements), mode (for counts) and standard deviation were calculated in Microsoft Excel 2010 software. The standard length (SL) is expressed in millimeters, the other measurements are presented as a percentage of the standard length (measurements referring to the body, except head parts) or head length (HL; measurements referring to the head); the numbers in parentheses refer to the mean or the mode. The nomenclature and the counting of the body plates follow Schaefer, (1997)Schaefer SA. The Neotropical cascudinhos: Systematics and biogeography of the Otocinclus catfishes (Siluriformes: Loricariidae). Proc Acad Nat Sci. 1997; 148:1–120. Available from: https://www.jstor.org/stable/4065046
https://www.jstor.org/stable/4065046...
. According to Oyakawa et al., (2018)Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
and Oliveira, Oyakawa, (2019)Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
, the abdomen comprises the area between the pectoral girdle, the lateral thoracic plates and the pre-anal plates, or the urogenital pore, in case the pre-anal plates are lacking. The list of the examined material includes the acronym of the collection and the catalog number, followed by the number of copies in alcohol and, in parentheses, the specimens cleared and stained (c&s) according to Taylor, Van Dyke, (1985)Taylor WR, Van Dyke GC. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium. 1985; 9(2):107–19.. Non-types are small or dissected specimens, not used for taking measures and counts.

Data referring to Harttia depressa Rapp Py-Daniel & Oliveira, 2001, H. fluminensis Covain & Fisch-Muller, 2012, H. merevari Provenzano, Machado-Allison, Chernoff, Willink & Petry, 2005, H. rhombocephala Miranda Ribeiro, 1939, and H. tuna Covain & Fisch-Muller, 2012 were taken from the original descriptions. In the identification key, some data are from Oyakawa, (1993)Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. and Langeani et al., (2001)Langeani F, Oyakawa OT, Montoya-Burgos JI. New species of Harttia (Loricariidae, Loricariinae) from the rio São Francisco basin. Copeia. 2001(1):136–42. https://doi.org/10.1643/0045-8511(2001)001[0136:NSOHLL]2.0.CO;2
https://doi.org/10.1643/0045-8511(2001)0...
.

The material examined comes from the fish collections of the Departamento de Ciências Biológicas, Universidade Estadual Paulista “Júlio de Mesquita Filho”, São José do Rio Preto (DZSJRP); Instituto Nacional de Pesquisas da Amazônia, Manaus (INPA); Laboratório de Ictiologia da Universidade de São Paulo, Ribeirão Preto (LIRP); Museu de Ciências Naturais, Pontifícia Universidade Católica de Minas Gerais, Belo Horizonte (MCNIP); Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre (MCP); Museum d’Histoire Naturelle, Département d’Herpétologie et Ichthyologie, Ville de Genève, Genève (MHNG); Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro (MNRJ); and Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP).

RESULTS

Harttia canastra, new species

urn:lsid:zoobank.org:act:B141D47F-D458-4951-A3DE-6EBC61441A46

(Figs. 14; Tab. 2)

Harttia sp. —Casatti, Castro, 1998Casatti L, Castro RMC. A fish community of the São Francisco River headwaters riffles, southeastern Brasil. Ichthyol Explor Freshw. 1998; 9(3):229–42.:232, 233, 236 [tabs. 1–2; fig. 3m, paratype, male; 5 ecology, spatial distribution, behavior, figure]. —Casatti, Castro, 2006Casatti L, Castro RMC. Testing the ecomorphological hypothesis in a headwater riffles fish assemblage of the rio São Francisco, southeastern Brazil. Neotrop Ichthyol. 2016; 4(2):203–14. https://doi.org/10.1590/S1679-62252006000200006
https://doi.org/10.1590/S1679-6225200600...
:204, 206, 210–211 (tabs. 1–2; figs. 2–3, 8, ecomorphology).

Harttia cf. novalimensis. —Romero, Casatti, 2012Romero RM, Casatti L. Identification of key microhabitats for fish assemblages in tropical Brazilian savanna streams. Int Rev Hydrobiol. 2012; 97(6):526–41. https://doi.org/10.1002/iroh.201111513
https://doi.org/10.1002/iroh.201111513...
:541 [ecology]. —Roa-Fuentes et al., 2015:170, 178 [ecomorphology, feeding behavior, list].

Harttia novalimensis (not Oyakawa). —Angulo, 2019Angulo A. Gross brain morphology of the Loricariinae (Siluriformes: Loricariidae): comparative anatomy, ecological implications and phylogenetic analysis. [PhD Thesis]. São José do Rio Preto: Universidade Estadual Paulista; 2019. Available from: http://hdl.handle.net/11449/182045
http://hdl.handle.net/11449/182045...
:208 [brain comparative anatomy].

Harttia sp. São Roque. —Cherobim, 2022Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
:282 (morphological phylogenetic relationships), 307 (appendix 1, material examined).

Harttia sp. São Francisco. —Cherobim, 2022Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
:344 (in table), 356 (molecular phylogenetic relationships).

Holotype. LIRP 651, male, 99.2 mm SL, Brazil, Minas Gerais State, rio São Francisco, São Roque de Minas municipality, Fazenda Casca D’Anta, 20o30’00”S 46o50’00”W, Dec 1993 – May 1995, L. Casatti, R. M. C. Castro & H. F. Santos.

Paratypes. Minas Gerais State, rio São Francisco. Arapuá municipality. DZSJRP 20129, 5, 19.3–55.4 mm SL (1, 57.9 mm SL) brigde over the ribeirão Bebedouro, on land access to Chaves upstream from Arapuá, 19o02’50”S 46o09’26”W, 920 m asl, 27 Aug 2014, F. Langeani, A. M. Cherobim, B. N. Andrade & A. C. Rosa. São Roque de Minas municipality: LIRP 16932, 25, 19.3–105.1 mm SL (16, 48.8–105.1 mm SL), same data as holotype. MZUSP 50751, (1, 104.3 mm SL), same data as holotype. MZUSP 58677, 10, 37.0–103.3 mm SL (8, 50.2–103.3 mm SL), same data as holotype. DZSJRP 11432, 8, 45.2–86.5 mm SL (3, 61.3–86.5 mm SL), Córrego Grande, surroundings of Parque Nacional Serra da Canastra, owned by Mr. Elmo, 20o20’25”S 46o27’56”W, 19 Jun 2008, R. M. Romero & C. P. Ferreira. DZSJRP 11450, 3, 31.8–74.3 mm SL (2, 65.9–74.3 mm SL), same locality as DZSJRP 11432, 9 Dec 2008, R. M. Romero & G. H. Baviera. DZSJRP 11462, (3, 48.9–79.1 mm SL), Córrego Grande, surroundings of Parque Nacional Serra da Canastra, owned by Mr. Elmo, 20o20’25”S 46o27’56”W, 23 Mar 2009, R. W. Romero & C. A. Roa-Fuentes. DZSJRP 11562, 3, 39.2–108.9 mm SL (2, 102.0–108.9 mm SL), Córrego Cachoeirinha, surroundings of Parque Nacional Serra da Canastra, in front of Brazil Diamond, 20o19’27”S 46o32’15”W, 25 Oct 2009, R. M. Romero & G. H. Baviera. DZSJRP 11568, 1, 61.7 mm SL, same locality as DZSJRP 11562, 7 Nov 2009, R. M. Romero & G. H. Baviera. DZSJRP 11585, 28, 24.4–66.3 mm SL (7, 55.2–66.3 mm SL; 1, c&s), Córrego Lavra, surroundings of Parque Nacional Serra da Canastra, property of Mr. Vitor, upstream bridge, 20o18’36”S 46o25’59”W, 9 Aug 2008, R. M. Romero & A. R. Manzotti. DZSJRP 11597, 7, 50.9–53.9 mm SL (2, 50.9–53.9 mm SL), same locality as DZSJRP 11585, 27 Mar 2009, R. M. Romero & C. A. Roa-Fuentes. DZSJRP 11604, 5, 44.1–68.5 mm SL (2, 51.9–64.2 mm SL), same locality as DZSJRP 11585, 5 Nov 2009, R. M. Romero & G. H. Baviera. DZSJRP 11650, 5, 44.4–84.6 mm SL (4, 48.3–84.6 mm SL), Córrego Luciano, surroundings of Parque Nacional Serra da Canastra, park entrance under bridge, 20o18’46”S 46o31’46”W, 24 Mar 2009, R. M. Romero & C. A. Roa-Fuentes. DZSJRP 20172, (2, 72.1–80.9 mm SL), São José do Barreiro, riacho Grande, road from Vargem Bonita to São José do Barreiro, towards cachoeira da Casca d’Anta, 20o20’10”S 46o27’47”W, 18 Nov 2014, F. Langeani, A. M. Cherobim & B. N. Andrade. DZSJRP 20180, (7, 56.0–82.7 mm SL), São José do Barreiro, on the road from Vargem Bonita to São José do Barreiro, 20o20’00”S 46o28’06”W, 18 Nov 2014, F. Langeani, A. M. Cherobim & B. N. Andrade. DZSJRP 20188, 2, 45.4–81.9 mm SL (1, 81.9 mm SL), stream before the entrance of the Parque Nacional Serra da Canastra, São José do Barreiro, 20o19’16”S 46o31’58”W, 18 Nov 2014, F. Langeani, A. M. Cherobim & B. N. Andrade. INPA 59845, 5, 44.4–60.0 mm SL (1, 60.0 mm SL), same locality as DZSJRP 11585, 5 Nov 2009, R. M. Romero & G. H. Baviera. MCP 54879, 5, 29.8–75.9 mm SL (2, 50.3–75.9 mm SL), same data as DZSJRP 20188. MHNG 2787.091, 5, 38.0–73.0 mm SL, same data as DZSJRP 20180. MNRJ 53314, 5, 30.8–63.2 mm SL (3, 49.6–63.2 mm SL), same data as DZSJRP 20188.

FIGURE 1 |
Harttia canastra, LIRP 651, male, holotype, 99.2 mm SL, Brazil, Minas Gerais State, rio São Francisco, São Roque de Minas municipality, Fazenda Casca D’Anta. A. Dorsal view; B. Lateral view; C. Ventral view.

FIGURE 2 |
Harttia canastra, DZSJRP 20172, female, paratype, 80.9 mm SL, São José do Barreiro, riacho Grande, road Vargem Bonita to São José do Barreiro, towards Cachoeira Casca D’Anta. A. Dorsal view; B. Lateral view; C. Ventral view.

Non-types. Minas Gerais State, rio São Francisco, São Roque de Minas municipality: same locality as DZSJRP 11432: DZSJRP 11441, 4, 27.6–66.5 mm SL, 7 Aug 2008, R. M. Romero & A. R. Manzotti. DZSJRP 11445, 7, 15.3–58.0 mm SL, 8 Oct 2008, R. M. Romero & V. H. M. Prado. DZSJRP 11456, 4, 43.1–98.6 mm SL, 8 Feb 2009, R. M. Romero & G. H. Baviera. DZSJRP 11468, 6, 15.6–40.1 mm SL, Córrego Cerrado, surroundings of Parque Nacional Serra da Canastra, owned by Mr. Joaquim Evaristo, 20o20’0”S 46o28’31”W, 18 Jun 2009, R. M. Romero & C. P. Ferreira. Same locality as DZSJRP 11468: DZSJRP 11474, 12, 16.2–57.2 mm SL, 8 Ago 2008, R. M. Romero & A. R. Manzotti. DZSJRP 11483, 6, 11.7–46.8 mm SL, 9 Oct 2008, R. M. Romero & V. H. M. Prado. DZSJRP 11491, 10, 21.2–46.0 mm SL, 9 Dec 2008, R. M. Romero & G. H. Baviera. DZSJRP 11500, 2, 36.7–39.5 mm SL, 7 Feb 2009, R. M. Romero & G. H. Baviera. DZSJRP 11509, 1, 19.3 mm SL, 25 Mar 2009, R. M. Romero & C. A. Roa-Fuentes. DZSJRP 11516, 12, 52.4–52.5 mm SL, Córrego da Mandioca, surroundings of Parque Nacional Serra da Canastra, São José do Barreiro, Fazenda Passaredo, 20o18’53”S 46o26’22”W, 10 Jun 2008, R. M. Romero & C. P. Ferreira. Same locality as DZSJRP 11516: DZSJRP 11522, 16, 20.0–52.0 mm SL, 9 Aug 2008, R. M. Romero & A. R. Manzotti. DZSJRP 11535, 2, 29.4–31.6 mm SL, 25 Aug 2009, R. M. Romero & G. H. Baviera. DZSJRP 11528, 15, 29.9–53.5 mm SL, Córrego da Mandioca, bridge on the road, surroundings of Parque Nacional Serra da Canastra, São José do Barreiro, Fazenda Passaredo, 20o18’53”S 46o27’22”W, 8 Jun 2008, R. M. Romero & V. H. M. Prado. DZSJRP 11540, 3, 23.2–43.8 mm SL, same locality as DZSJRP 11528, 5 Nov 2009, R. M. Romero & G. H. Baviera. DZSJRP 11591, 7, 28.1–31.0 mm SL, same locality as DZSJRP 11585, 9 Out 2008, R. M. Romero & V. H. M. Prado. DZSJRP 15826, 1, 28.8 mm SL, same locality as DZSRP 11516, 8 Jun 2008, R. M. Romero and G. H. Baviera. Same locality as DZSJRP 11620: DZSJRP 11626, 6, 55.2–73.0 mm SL, 7 Aug 2008, R. M. Romero & A. R. Manzotti. DZSJRP 11632, 4, 31.1–97.4 mm SL, 24 Mar 2009, R. M. Romero & C. A. Roa-Fuentes.

Diagnosis. The naked abdomen between pectoral- and pelvic-fin girdles readily discriminates Harttia canastra from Harttia absaberi Oyakawa, Fichberg & Langeani, 2013, H. dissidens Rapp Py-Daniel & Oliveira, 2001, H. duriventris Rapp Py-Daniel & Oliveira, 2001, H. fluminensis, H. fowleri (Pellegrin, 1908), H. longipinna, H. panara Oyakawa, Fichberg & Rapp Py-Daniel, 2018, H. punctata Rapp Py-Daniel & Oliveira, 2001, H. rhombocephala, H. rondoni Oyakawa, Fichberg & Rapp Py-Daniel, 2018, H. surinamensis Boeseman, 1971, H. trombetensis Rapp Py-Daniel & Oliveira, 2001, H. tuna, and H. villasboas Oyakawa, Fichberg & Rapp Py-Daniel, 2018 (vs. abdomen partially or completely covered by plates; Figs. 3B,C). Harttia canastra can be distinguished from H. carvalhoi Miranda Ribeiro, 1939, H. garavelloi Oyakawa, 1993, H. intermontana Oliveira & Oyakawa, 2019, H. kronei Miranda Ribeiro, 1908, H. leiopleura, and H. novalimensis by having preanal plates (vs. preanal plates absent; Figs. 3A,D,E). Furthermore, Harttia canastra can be distinguished from H. torrenticola by having two to four large trapezoidal preanal plates and dorsal-fin spinelet present (vs. two to four circular preanal plates and dorsal-fin spinelet absent; Fig. 3F). It differs from Harttia gracilis Oyakawa, 1993 in that its lower and upper caudal rays are of same size (vs. upper ray slightly longer than the lower one). Harttia canastra can be distinguished from H. guianensis Rapp Py-Daniel & Oliveira, 2001, H. loricariformis Steindachner, 1877, and H. uatumensis Rapp Py-Daniel & Oliveira, 2001 by the presence of the dorsal-fin spinelet and by the hypertrophy of odontodes in the lateral region of the head, close to the canal plate, and in the unbranched pectoral fin ray in adult males (vs. absence of the dorsal-fin spinelet and males without hypertrophied odontodes). Distinguished from Harttia depressa by body depth 40.0–66.7% and head depth 35.7–52.6% of HL (vs. extremely depressed body and head, respectively 27.0–33.3% and 31.2–35.7% of HL). It differs from H. merevari by having anterior region of head more rounded, darker coloration and adult males with hypertrophied odontodes (vs. anterior region of head more triangular, light or dark yellow with many spots and males without hypertrophied odontodes). Finally, in relation to the other species of the rio São Francisco basin, H. canastra is distinguished from H. longipinna by the absence of plates in the abdominal region between pectoral- and pelvic-fin girdles and by the equivalent anal-fin length both in males and females (respectively 11.7–18.2% and 11.7–18.7% of SL; vs. partially covered abdomen and anal-fin length longer in males than in females (respectively 20.0–25.0% and 13.9–16.1% of SL; Fig. 3B); from H. leiopleura and H. novalimensis for presenting preanal plates (vs. preanal plates absent; Figs. 3D,E); and from H. torrenticola by having two large preanal trapezoidal plates (vs. two to four circular minute preanal plates; Fig. 3F) and the dorsal-fin spinelet (vs. dorsal-fin spinelet absent).

FIGURE 3 |
Abdominal (yellow), thoracic (blue), preanal (pink), and canal (purple) plates: (A) Harttia intermontana, DZSJRP 20978, male, 53.6 mm SL, with a naked abdomen (Group 1), thoracic and canal plates present; (B) Harttia longipinna, DZSJRP 3666, paratype, male, 82.3 mm SL, with abdomen partially covered by plates (Group 2), preanal, thoracic and canal plates present; (C) Harttia duriventris, MZUSP 106199, male, 114.9 mm SL, with fully plated abdomen (Group 3), preanal, thoracic and canal plates present; (D) Harttia leiopleura, MZUSP 109373, 48.3 mm SL, with a naked abdomen (Group 1), preanal, thoracic, and canal plates absent; (E) Harttia novalimensis, MZUSP 99481, male, 49.0 mm SL, with a naked abdomen (Group 1), thoracic plates present, and preanal and canal plates absent; (F) Harttia torrenticola, MZUSP 94482, male, 85.5 mm SL, with a naked abdomen (Group 1), preanal, thoracic, and canal plates present; (G) Harttia canastra, LIRP 651, male, holotype, 99.2 mm SL, with a naked abdomen (Group 1), preanal, thoracic, and canal plates present.

Description. Examined material: 247 specimens (68 measured and counted). Small size, 48.3–108.9 mm SL (Tab. 2). Body depressed and elongated, wider at cleithrum, progressively narrower from origin of pectoral fin to caudal peduncle. Dorsal profile convex from tip of snout to anterior region of dorsal fin, gently descending to caudal peduncle. Ventral profile of body straight from tip of snout to origin of anal fin, then gently ascending to caudal peduncle. Anterior region of head elliptical in dorsal view. Orbit rounded, preorbital crest rudimentary (crest not as prominent as in Amazonian Harttia species), inferior margin of orbit approximately straight (margin not as straight as Harttia loricariformis). Snout tip naked. Maxillary barbel rudimentary and joined to lips. Lower lip larger than upper lip, both papillose. Lower lip usually reaching posterior margin of coracoid. Infraorbital sensory canal in six plates. Canal plate present. Premaxillary with 30–67 (40) and dentary with 29–70 (38) rounded, bicuspid, and subequal teeth. Abdominal plates between pectoral- and pelvic-fin girdles absent; two to four large and trapezoidal preanal plates, sometimes bordered by small plates; specimens smaller than 60 mm SL usually with no preanal plates. Five longitudinal series of plates on body: dorsal series, 19–27 (24) plates; mid-dorsal series, 12–20 (17) plates; median series, 25–30 (27) perforated plates; mid-ventral series, 14–21 (17) plates; ventral series, 19–23 (22) plates. Three to eight (five) rectangular and elongated lateral thoracic plates; plates between dorsal and caudal fin, 14–18 (16); plates from anal to caudal fin, 12–16 (14); two long, poreless supracaudal plates on the lateral line. Dorsal fin, II,7; dorsal-fin spinelet present; anal fin, i,5; pectoral fin, I,6; pelvic fin, i,5; caudal fin, i,12,i, forked, upper and lower unbranched rays subequal. Caudal peduncle, viewed from above, slightly more tapered after confluence of lateral plates series.

TABLE 2 |
Measurements and counts of Harttia canastra. Standard length (mm), percentages of body measurements in relation to standard length (SL) or percentage of head measurements in relation to head length (HL). Dorsal fin origin (D), anal fin origin (A), caudal fin origin (C), standard deviation (SD). Total number of analyzed type-specimens=68.

Coloration in alcohol. Light brown body with brown spots along dorsal region, forming four inconspicuous transverse bands, starting at origin of dorsal fin; ventral region light yellow. Pectoral, dorsal, pelvic, anal and caudal-fin rays with dark brown spots, interradial membrane hyaline.

Sexual dimorphism. Males with hypertrophied odontodes in the first pectoral-fin ray and in the lateral region of the head, close to the opercular opening, and an elongated urogenital papilla (vs. females without developed odontodes and with a short urogenital papilla, resembling a pore) (Figs. 12).

Geographical distribution. Several locations in the rio São Francisco basin, mainly headwater streams around the Parque Nacional Serra da Canastra, Municipality of São Roque de Minas, State of Minas Gerais (Fig. 4).

Ecological notes.Harttia canastra type locality consisted of an approximately 60 m riffle stretch with substrate consisting of sand, gravel, and rocks, current ranging 0.3–2.3 m.s-1, width from 4–13 m, depth from 0.5–1.2 m, discharge from 1.8–3.2 m3.s-1, subaquatic horizontal transparency from 1.6– 9.6 m, air temperature from 10–31.5oC, water temperature from 12.4–23oC, and dissolved oxygen from 8.9–21.7 mg.l-1 (Casatti, Castro, 1998Casatti L, Castro RMC. A fish community of the São Francisco River headwaters riffles, southeastern Brasil. Ichthyol Explor Freshw. 1998; 9(3):229–42., 2006Casatti L, Castro RMC. Testing the ecomorphological hypothesis in a headwater riffles fish assemblage of the rio São Francisco, southeastern Brazil. Neotrop Ichthyol. 2016; 4(2):203–14. https://doi.org/10.1590/S1679-62252006000200006
https://doi.org/10.1590/S1679-6225200600...
). Romero, Casatti, (2012)Romero RM, Casatti L. Identification of key microhabitats for fish assemblages in tropical Brazilian savanna streams. Int Rev Hydrobiol. 2012; 97(6):526–41. https://doi.org/10.1002/iroh.201111513
https://doi.org/10.1002/iroh.201111513...
and Roa-Fuentes et al., (2015)Roa-Fuentes CA, Casatti L, Romero RM. Phylogenetic signal and major ecological shifts in the ecomorphological structure of stream fish in two river basins in Brazil. Neotrop Ichthyol. 2015; 13(1):165–78. https://doi.org/10.1590/1982-0224-20140045
https://doi.org/10.1590/1982-0224-201400...
presented additional information about other localities where most of the paratypes were collected, and also about other species inhabiting the same sites. Other species occurring with Harttia canastra include: Apareiodon ibitiensis Amaral Campos, 1944, A. piracicabae (Eigenmann, 1907), Cetopsorhamdia iheringi Schubart & Gomes, 1959, Characidium fasciatum Reinhardt, 1867, C. zebra Eigenmann, 1909, Creagrutus varii Ribeiro, Benine & Figueiredo, 2004, Harttia cf. novalimensis, Hisonotus alberti Roxo, Silva, Waltz & Melo, 2016, Hoplias malabaricus (Bloch, 1794), Hypostomus garmani (Regan, 1904), Neoplecostomus canastra Roxo, Silva, Zawadzki & Oliveira, 2017, Oreochromis niloticus (Linnaeus, 1758), Parodon hilarii Reinhardt, 1867, Phalloceros sp., Piabarchus stramineus (Eigenmann, 1908), Piabina argentea Reinhardt, 1867, Psalidodon rivularis (Lütken, 1875), Rhamdia aff. quelen (Quoy & Gaimard, 1824), Trichomycterus brasiliensis Lütken, 1874, T. reinhardti (Eigenmann, 1917), T. variegatus Costa, 1992 (all of them deposited at the DZSJRP fish collection).

Etymology. The name “canastra” refers to the Serra da Canastra, a mountain range located in the center-south of the state of Minas Gerais, which houses the headwaters of the rio São Francisco, where most of the specimens were collected. A noun in apposition.

Conservation status. The species was sampled in various localities in the rio São Francisco basin, some of them very near the Parque Nacional da Serra da Canastra, a national conservation unit. This large distribution, together with its abundance and frequency of capture, suggests that Harttia canastra can be categorized as Least Concern (LC) according to the International Union for Conservation of Nature (IUCN) categories and criteria (IUCN Standards and Petitions Subcommittee, 2022International Union for Conservation of Nature (IUCN). Standards and petitions subcommittee. Guidelines for using the IUCN Red List categories and criteria. Version 15 [Internet]. 2022. Available from: https://www.iucnredlist.org/resources/redlistguidelines
https://www.iucnredlist.org/resources/re...
).

Key to species of Harttia from the rio São Francisco basin

1a. Canal plate and preanal plates present 2

1b. Canal plate and preanal plates absent 4

2a. Abdomen region between pectoral- and pelvic-fin girdles naked; anal-fin size equivalent in males and females (respectively, 11.7–18.2% and 11.7–18.7% of SL) 3

2b. Abdomen region between pectoral- and pelvic-fin girdles partially covered by plates; anal-fin size in adult males longer than in females, respectively 20–25% and 13.9–16.1% of SL H. longipinna

3a. Two to four small and circular pre-anal plates; dorsal-fin spinelet absent H. torrenticola

3b. Two large and trapezoidal pre-anal plates, sometimes with small plates around them; dorsal-fin spinelet present H. canastra

4a. Thoracic plates absent; five infraorbital plates; seven branched rays on the pectoral fin H. leiopleura

4b. Three to five thoracic plates; six infraorbital plates; six branched rays on the pectoral fin H. novalimensis

FIGURE 4 |
Distribution map of Harttia species from the rio São Francisco basin.

DISCUSSION

In the last 10 years, seven new Harttia species were described, two for the Guiana Shield drainages (Covain et al., 2012Covain R, Fisch-Müller S, Montoya-Burgos JI, Mol JH, Le Bail PY, Dray S. The Harttiini (Siluriformes, Loricariidae) from the Guianas: a multi-table approach to assess their diversity, evolution, and distribution. Cybium. 2012; 36(1):115–61.) and five for the Brazilian Crystaline Shield drainages (Oyakawa et al., 2013Oyakawa OT, Fichberg I, Langeani F. Harttia absaberi, a new species of loricariid catifish (Siluriformes: Loricariidae: Loricariinae) from the upper rio Paraná basin, Brazil. Neotrop Ichthyol. 2013; 11(4):779–86. https://doi.org/10.1590/S1679-62252013000400005
https://doi.org/10.1590/S1679-6225201300...
, 2018Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
; Oliveira, Oyakawa, 2019Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
). The Brazilian Crystalline Shield southeastern basins, with the headwaters of the rios Doce, Grande, Jequitinhonha, Paraíba do Sul and São Francisco, and also smaller coastal basins, drain the southern portions of the Espinhaço Complex and western portions of the Serra da Mantiqueira. Most of its species occur in forested headwater streams, with sandy and rocky bottoms, and clear, cold and fast-flowing water (Casatti, Castro, 1998Casatti L, Castro RMC. A fish community of the São Francisco River headwaters riffles, southeastern Brasil. Ichthyol Explor Freshw. 1998; 9(3):229–42., 2006Casatti L, Castro RMC. Testing the ecomorphological hypothesis in a headwater riffles fish assemblage of the rio São Francisco, southeastern Brazil. Neotrop Ichthyol. 2016; 4(2):203–14. https://doi.org/10.1590/S1679-62252006000200006
https://doi.org/10.1590/S1679-6225200600...
; Romero, Casatti, 2012Romero RM, Casatti L. Identification of key microhabitats for fish assemblages in tropical Brazilian savanna streams. Int Rev Hydrobiol. 2012; 97(6):526–41. https://doi.org/10.1002/iroh.201111513
https://doi.org/10.1002/iroh.201111513...
; Oyakawa et al., 2013Oyakawa OT, Fichberg I, Langeani F. Harttia absaberi, a new species of loricariid catifish (Siluriformes: Loricariidae: Loricariinae) from the upper rio Paraná basin, Brazil. Neotrop Ichthyol. 2013; 11(4):779–86. https://doi.org/10.1590/S1679-62252013000400005
https://doi.org/10.1590/S1679-6225201300...
; Roa-Fuentes et al., 2015Roa-Fuentes CA, Casatti L, Romero RM. Phylogenetic signal and major ecological shifts in the ecomorphological structure of stream fish in two river basins in Brazil. Neotrop Ichthyol. 2015; 13(1):165–78. https://doi.org/10.1590/1982-0224-20140045
https://doi.org/10.1590/1982-0224-201400...
; Oliveira, Oyakawa, 2019Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
). Harttia longipinna, on the contrary, inhabits large rivers (Langeani et al., 2001Langeani F, Oyakawa OT, Montoya-Burgos JI. New species of Harttia (Loricariidae, Loricariinae) from the rio São Francisco basin. Copeia. 2001(1):136–42. https://doi.org/10.1643/0045-8511(2001)001[0136:NSOHLL]2.0.CO;2
https://doi.org/10.1643/0045-8511(2001)0...
; our data), and in this respect is more similar to species of the Amazon, Orinoco, and Guiana Shield basins (Rapp Py-Daniel, Oliveira, 2001Rapp Py-Daniel LH, Oliveira EC. Seven New Species of Harttia from the Amazonian-Guyana Region (Siluriformes: Loricariidae). Ichthyol Explor Freshw. 2001; 12(1):79–96. ; Provenzano et al., 2005Provenzano F, Machado-Allison A, Chernoff B, Willink P, Petry P. Harttia merevari, a new species of catfish (Siluriformes: Loricariidae) from Venezuela. Neotrop Ichthyol. 2005; 3(4):519–24. Available from: https://www.scielo.br/j/ni/a/Cvxb3FnRDFd3J7wK3L5nv5R/?format=pdf⟨=en
https://www.scielo.br/j/ni/a/Cvxb3FnRDFd...
; Covain et al., 2012Covain R, Fisch-Müller S, Montoya-Burgos JI, Mol JH, Le Bail PY, Dray S. The Harttiini (Siluriformes, Loricariidae) from the Guianas: a multi-table approach to assess their diversity, evolution, and distribution. Cybium. 2012; 36(1):115–61.; Oyakawa et al., 2018Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
; our data; L. Rapp Py-Daniel, 2022, pers. comm.). The smaller environments, apparently preferred by these southeastern Harttia species, are now significantly altered (Oliveira, Oyakawa, 2019Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
), which seems to have contributed to the restriction of their distribution into reduced areas.

The pattern of plates in the body ventral region is very important in discriminating Harttia species, comprising the preanal plates, thoracic plates, abdominal plates, and canal plate, as already noted by Oyakawa, (1993)Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. , Rapp Py-Daniel, Oliveira, (2001)Rapp Py-Daniel LH, Oliveira EC. Seven New Species of Harttia from the Amazonian-Guyana Region (Siluriformes: Loricariidae). Ichthyol Explor Freshw. 2001; 12(1):79–96. , Langeani et al., (2001)Langeani F, Oyakawa OT, Montoya-Burgos JI. New species of Harttia (Loricariidae, Loricariinae) from the rio São Francisco basin. Copeia. 2001(1):136–42. https://doi.org/10.1643/0045-8511(2001)001[0136:NSOHLL]2.0.CO;2
https://doi.org/10.1643/0045-8511(2001)0...
, Provenzano et al., (2005)Provenzano F, Machado-Allison A, Chernoff B, Willink P, Petry P. Harttia merevari, a new species of catfish (Siluriformes: Loricariidae) from Venezuela. Neotrop Ichthyol. 2005; 3(4):519–24. Available from: https://www.scielo.br/j/ni/a/Cvxb3FnRDFd3J7wK3L5nv5R/?format=pdf⟨=en
https://www.scielo.br/j/ni/a/Cvxb3FnRDFd...
, Oyakawa et al., (2013Oyakawa OT, Fichberg I, Langeani F. Harttia absaberi, a new species of loricariid catifish (Siluriformes: Loricariidae: Loricariinae) from the upper rio Paraná basin, Brazil. Neotrop Ichthyol. 2013; 11(4):779–86. https://doi.org/10.1590/S1679-62252013000400005
https://doi.org/10.1590/S1679-6225201300...
, 2018Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
), and Oliveira, Oyakawa, (2019)Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
. However, the definition of each of these plates can be confusing to non-experts in the group and non-taxonomists. Thus, we present a clearer delimitation of these plates based on our interpretation of previously published description articles. The preanal plates are those strictly close to the anus and the urogenital pore (Fig. 3, pink). The thoracic plates are the ones on the sides of the body, between the pectoral and pelvic girdles (Fig. 3, blue). The abdominal plates correspond to all the other plates that appear in the ventral region, between the pectoral girdle and the preanal plates (Fig. 3, yellow). Finally, the canal plate is an approximately triangular plate located ventrally to the preopercle that bears the final portion of the preopercular ramus of the laterosensory canal (Schaefer, 1987Schaefer SA. Osteology of Hypostomus plecostomus (Linnaeus), with a phylogenetic analysis of the loricariid subfamilies (Pisces: Siluroidei). Contr Sci Nat His Mus Los Angeles. 1987; 394:1–31. , 1991Schaefer SA. Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae (Pisces: Siluroidei: Loricariidae), with comments on generic diagnoses and geographic distribution. Zool J Linn Soc. 1991; 102(1):1–41. https://doi.org/10.1111/j.1096-3642.1991.tb01535.x
https://doi.org/10.1111/j.1096-3642.1991...
; Armbruster, 2004Armbruster JW. Phylogenetic relationships of the suckermouth armoured catfishes (Loricariidae) with emphasis on the Hypostominae and the Ancistrinae. Zool J Linn Soc. 2004; 141(1):1–80. https://doi.org/10.1111/j.1096-3642.2004.00109.x
https://doi.org/10.1111/j.1096-3642.2004...
; Pereira, Reis, 2017Pereira EHL, Reis RE. Morphology-based phylogeny of the suckermouth armored catfishes, with emphasis on the Neoplecostominae (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2017; 4264 (1):1–104. https://doi.org/10.11646/zootaxa.4264.1.1
https://doi.org/10.11646/zootaxa.4264.1....
) (Fig. 3, purple).

Additionally, to better understand the three types of abdominal plates coverage proposed by Oyakawa et al., (2018)Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
, it is necessary to observe the area between the pectoral and pelvic girdles. The abdomen completely covered by plates represents the condition where there are abdominal plates along the entire area between the two girdles (Fig. 3C). In the abdomen partially covered, there are regions with and without abdominal plates between the two girdles, the proportion of each vary depending on the species (Fig. 3B). In the last condition, e.g., the abdomen naked, there are no abdominal plates between the two girdles (Fig. 3A,D,E,F), although in some cases abdominal ones might be present around the preanal plates, after the pelvic girdle (Fig. 3G). Therefore, Harttia canastra is readily distinguished from species of the genus, including its congeners of the rio São Francisco basin, due to the following plate pattern combination: two to four large and trapezoidal preanal plates (sometimes bordered by small abdominal plates), abdomen naked (no abdominal plates between the two girdles), and thoracic and canal plates present. In this respect, it is worth mentioning that among the comparative material from the rio São Francisco basin we have observed specimens of H. longipinna (MCNIP 1584 and 1588) with and without abdominal plates, and specimens here identified as H. cf. novalimensis (DZSJRP 11612, 11620, and 2016) with a canal plate, features not mentioned until now for those two species (Oyakawa, 1993Oyakawa OT. Cinco espécies novas de Harttia Steindachner, 1876 da região Sudeste do Brasil, e comentários sobre o gênero (Teleostei, Siluriformes, Loricariidae). Comun Mus Ciênc Tecnol PUCRS, Sér Zool.1993; 7:3–27. ; Langeani et al., 2001Langeani F, Oyakawa OT, Montoya-Burgos JI. New species of Harttia (Loricariidae, Loricariinae) from the rio São Francisco basin. Copeia. 2001(1):136–42. https://doi.org/10.1643/0045-8511(2001)001[0136:NSOHLL]2.0.CO;2
https://doi.org/10.1643/0045-8511(2001)0...
; Oyakawa et al., 2013Oyakawa OT, Fichberg I, Langeani F. Harttia absaberi, a new species of loricariid catifish (Siluriformes: Loricariidae: Loricariinae) from the upper rio Paraná basin, Brazil. Neotrop Ichthyol. 2013; 11(4):779–86. https://doi.org/10.1590/S1679-62252013000400005
https://doi.org/10.1590/S1679-6225201300...
, 2018Oyakawa OT, Fichberg I, Rapp Py-Daniel LH. Three new species of Harttia (Loricariidae: Loricariinae) from Serra do Cachimbo, rio Xingu basin, Pará, Northern Brazil. Zootaxa. 2018; 4387(1):75–90. https://doi.org/10.11646/zootaxa.4387.1.3
https://doi.org/10.11646/zootaxa.4387.1....
; Oliveira, Oyakawa, 2019Oliveira CJ, Oyakawa OT. New loricariid fishes from headwaters on Serra da Mantiqueira and Complexo do Espinhaço, Minas Gerais State, Brazil (Teleostei: Siluriformes: Loricariidae). Zootaxa. 2019; 4586(3):401–24. https://doi.org/10.11646/zootaxa.4586.3.1
https://doi.org/10.11646/zootaxa.4586.3....
); this intraspecific variation should be better analyzed by a future taxonomic revision of Harttia from the rio São Francisco basin.

Furthermore, Cherobim, (2022)Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
performed a morphological and molecular phylogenetic analysis of the genus Harttia and included H. canastra. It is referred to as Harttia sp. São Roque in the morphological analysis, and Harttia sp. São Francisco in the molecular analysis. In the morphological phylogeny, Cherobim, (2022)Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
recovered a monophyletic group composed of: (H. canastra, São Francisco basin + (H. carvalhoi, Paraíba do Sul basin + (H. intermontana, Doce basin + H. torrenticola, São Francisco basin))). In the molecular phylogeny, another monophyletic group was recovered with the same species, but in a different topology: (Harttia carvalhoi + (H. intermontana + (H. canastra + (H. torrenticola + H. canastra)))). This unexpected result in the molecular analysis, with a paraphyletic H. canastra, led us to check those previous identifications. Some samples previously identified in Cherobim, (2022)Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
as Harttia canastra (DZSJRP 16126), which resulted as sister to H. torrenticola as in the relationships mentioned above, are in fact belonging to H. torrenticola. Therefore, the molecular hypothesis of Cherobim, (2022)Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
would actually be: (Harttia carvalhoi + (H. intermontana + (H. canastra + (H. torrenticola + H. torrenticola)))), resulting in H. canastra as sister to H. torrenticola. According to Cherobim, (2022)Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
, the establishment of a correlation between the phylogenetic relationships among these species and their geographic distribution pattern, does not result in a simple explanation, demonstrating the complexity of this area, and its biogeographic discontinuity as previously highlighted by Ribeiro, (2006)Ribeiro AC. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop Ichthyol. 2006; 4(2):225–46. https://doi.org/10.1590/S1679-62252006000200009
https://doi.org/10.1590/S1679-6225200600...
and Buckup, (2011)Buckup PA. The Eastern Brazilian Shield. In: Albert JS, Reis RE, editors. Historical biogeography of Neotropical freshwater fishes. Los Angeles: University of California Press; 2011. p.203–10.. The Brazilian Shield is a region characterized by many ancient faults from the time of Gondwana breakup, and also recent tectonic stresses and differential erosions that finally lead to stream captures. So, the “exchange of fauna among these basins was probably due to the continuity of these processes throughout history” (Cherobim, 2022Cherobim AM. Phylogenetic analysis of Harttia Steindachner, 1877 (Siluriformes: Loricariidae: Loricariinae). [PhD Tesis]. São José do Rio Preto: Universidade Estadual Paulista; 2022. Available from: https://repositorio.unesp.br/handle/11449/216922
https://repositorio.unesp.br/handle/1144...
). Besides, an increase in the diversity of the genus is expected in the rio São Francisco basin, as well as in the area of distribution of the species as the drainage becomes better sampled.

Comparative material examined. All from Brazil except otherwise noticed.Harttia absaberi: Mato Grosso do Sul State, upper rio Paraná basin, MZUSP 85806, 5 of 34 paratypes, (1 of 2 c&s), 56.4–42.2 mm SL. Harttia carvalhoi: Rio de Janeiro State, rio Paraíba do Sul, DZSJRP 20358, 3, 40.0–45.7 mm SL. São Paulo State, Pindamonhangaba, rio Paraíba do Sul; MZUSP 44505, 1 c&s; MZUSP 79367, 2, 68.8–75.9 mm SL. Harttia dissidens: Pará State, Itaituba municipality, INPA 7046, 2 of 22, (1 c&s), 108.6–126.5 mm SL. Harttia duriventris: Pará State, Marabá municipality, rio Itacaiúnas, MZUSP 34226, 4 of 26, 72.2–100.3 mm SL; MZUSP 106199, 2, 99.5–114.9 mm SL. Harttia fowleri: Amapá State, Porto Marcirio municipality, rio Araguari, INPA 7845, 2 of 49, (1 c&s), 105.4–142.3 mm SL. Harttia garavelloi: Minas Gerais State, rio Jequitinhonha, MZUSP 42696, 1 of 2 paratypes, c&s; MCP 49359, 18, 30.4–63.3 mm SL; MZUSP 94430, 3, 53.4–75.0 mm SL. Harttia gracilis: Minas Gerais State, Passos and Itaú de Minas municipalities, rio Grande, DZSJRP 17746, 1, 101.3 mm SL. São Paulo State, São Bento do Sapucaí municipality, rio Grande, MZUSP 99678, 7 of 29, (1 c&s), 38.5–83.1 mm SL. Harttia guianensis: French Guyana, Saut. Deux Roros, INPA 14985, 1 of 4, (1 c&s), 63.8-86.3 mm SL. Harttia intermontana: Minas Gerais State, rio Doce, DZSJRP 16150, 23, 43.5–46.0 mm SL; DZSJRP 19145, 2, 27.6–45.5 mm SL; DZSJRP 20069, 38, (1 c&s), 26.8–51.3 mm SL; DZSJRP 20978, 47: 33.0–53.6 mm SL; DZSJRP 22636, 5, 14.4–72.2 mm SL. Harttia kronei: São Paulo State, rio Ribeira de Iguape, MZUSP 10602, 2, 70.3–77.6 mm SL; MZUSP 42694, 1 c&s; MZUSP 53811, 9, 71.3–94.1 mm SL; DZSJRP 13697, 3, 51.9 mm SL. Harttia leiopleura: Minas Gerais State, Caeté, rio São Francisco, MZUSP 109373, 3, (1 c&s), 38.4–48.3 mm SL. Minas Gerais state, Ouro Preto, rio São Francisco, MZUSP 109426, 2, 43.2–45.2 mm SL. Harttia longipinna: Bahia State, São Desidério, rio São Francisco, DZSJRP 3666, 6 paratypes, (2 c&s), 48.6–84.7 mm SL. Minas Gerais state, rio São Francisco, DZSJRP 2819, 1 paratype, 84.0 mm SL; MCNIP 1584, 19, 29,6–90.1 mm SL; MCNIP 1588, 2, 56.7–98.5 mm SL; MZUSP 39145, 5, 64.0–80.0 mm SL; MZUSP 73692, 1, 108.6 mm SL. Harttia loricariformis: Minas Gerais State, Muriaé municipality, rio Paraíba do Sul, MZUSP 80954, 4, 61.8–117.9 mm SL. Rio de Janeiro State, São Fidélis municipality, rio Paraíba do Sul, DZSJRP 21103, 2, 130.3–131.4 mm SL. São Paulo State, rio Paraíba do Sul, MZUSP 2163, 1 c&s. Harttia novalimensis: Minas Gerais State, Nova Lima municipality, rio São Francisco, MZUSP 37147, 1 paratype, c&s, 55.4 mm SL; MZUSP 94481, 3 of 28, 44.5–49.7 mm SL. Harttia cf. novalimensis: Minas Gerais State, São Roque de Minas municipality, rio São Francisco, DZSJRP 11612, (2, 55.3–61.9 mm SL), Córrego Lavra, surroundings of Parque Nacional Serra da Canastra, property of Mr. Vitor, upstream bridge, 20o18’36”S 46o25’59”W, 9 Aug 2008; DZSJRP 11620, 4, 22.3–72.7 mm SL (3, 48.4–72.7 mm SL), Córrego Luciano, surroundings of Parque Nacional Serra da Canastra, park entrance under bridge, 20o18’46”S 46o31’46”W, 16 Jun 2008; DZSJRP 20166, 2, 47.9–72.3 mm SL (1, 71.2 mm SL), creek on the Vicinal on the right, unpaved road, São Roque de Minas to São José do Barreiro, before the Vargem Bonita, 20o19’19”S 46o23’27”W, 18 Nov 2014. Harttia panara: Pará State, rio Curuá, MZUSP 97088, 1 of 3 paratypes, c&s; MZUSP 118551, 1 of 2 paratypes, 89.9 mm SL. Harttia punctata: Goiás State, rio Tocantins, DZSJRP 22659, 1, 67.3 mm SL; MCP 15873, 2, 59.9–60.0 mm SL; MZUSP 15873, 2, 58.3–60.5 mm SL; MZUSP 96172, 5, (1 c&s), 65.8–92,7 mm SL. Maranhão State, rio Tocantins, DZSJRP 20764, 1, 72.2 mm SL; DZSJRP 20702, 1, 98.3 mm SL. Harttia rondoni: Pará State, Altamira municipality, rio Curuá, MZUSP 97083, 1 of 9 paratypes, c&s; MZUSP 97085, 9 paratypes, 90.3–109.7 mm SL. Harttia surinamensis: Amapá State, rio Cupixi, MZUSP 34224, 3 of 60, (1 c&s), 104.7–132.3 mm SL. Harttia torrenticola: Minas Gerais State, Belo Vale municipality, rio São Francisco, MZUSP 94482, 2, 69.4–85.5 mm SL. Cristiano Otoni municipality, rio São Francisco, DZSJRP 5725, 1 c&s, 51.8 mm SL; DZSJRP 16126, 21, 25.2–35.1 mm SL; DZSJRP 20970, 2, 20.0–30.8 mm SL; DZSJRP 20090, 9, 29.3–35.7 mm SL. Harttia trombetensis: Pará State, Oriximiná municipality, rio Trombetas, INPA 3011, 3 (1 c&s), 79.4–90.7 mm SL. Harttia uatumensis: Amazonas State, INPA 12954, 2, 74.0–87.7 mm SL; INPA 3239, 1, 105.8 mm SL. Suriname, MZUSP 107732, 2 of 11, (1 c&s), 131.8–132.5 mm SL. Harttia villasboas: Pará State, Altamira municipality, rio Curuá, MZUSP 115485, 2 of 58 paratypes, (1 of 6 c&s), 87.4–93.5 mm SL. Harttia sp. 1: Minas Gerais State, Delfim Moreira municipality, rio Grande, DZSJRP 20587, 1, 28.2 mm SL; DZSJRP 20588, 2, 110.0–116.6 mm SL. Harttia sp. 2: Pará State, rio Braço Norte, MZUSP 116594, 2, 89.6–89.9 mm SL; MZUSP 97079, 2 of 81, 100.2–106.2 mm SL.

ACKNOWLEDGEMENTS

This work was carried out with the support of the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, Código de Financiamento 001 to LC) and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq proc. 305756/2018–4, FL). To Profa. Dra. Lilian Casatti, to Profa. Dra. Lucia H. Rapp Py-Daniel and to Dr. Osvaldo T. Oyakawa for their suggestions in the early version of the article. To the two anonymous reviewers for their suggestions. To the curators of the Instituto Nacional de Pesquisas da Amazônia (INPA); Laboratório de Ictiologia de Ribeirão Preto (LIRP); Museu de Ciências Naturais da Pontifícia Universidade Católica de Minas Gerais (MCNIP); Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul (MCP); Museum d’Histoire Naturelle, Département d’Herpétologie et Ichthyologie, Ville de Genève (MHNG); Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ); Museu de Zoologia da Universidade de São Paulo (MZUSP) by borrowing specimens under their care.

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HOW TO CITE THIS ARTICLE

  • Caldas L, Cherobim AM, Langeani F. A new species of Harttia from the rio São Francisco basin (Siluriformes: Loricariidae). Neotrop Ichthyol. 2022; 20(4):e220051. https://doi.org/10.1590/1982-0224-2022-0051

Edited-by

Paulo Lucinda

Publication Dates

  • Publication in this collection
    21 Nov 2022
  • Date of issue
    2022

History

  • Received
    14 June 2022
  • Accepted
    27 Sept 2022
Sociedade Brasileira de Ictiologia Neotropical Ichthyology, Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá., Av. Colombo, 5790, 87020-900, Phone number: +55 44-3011-4632 - Maringá - PR - Brazil
E-mail: neoichth@nupelia.uem.br