Two new species of Astyanax Baird & Girard (Characiformes: Characidae) from the upper rio Paraguaçu basin, Chapada Diamantina, Bahia, Brazil

Abstract

Astyanax lorien and A. rupestris are described from tributaries of upper rio Paraguaçu basin, Bahia, Brazil. Both species shares the majority of the characters currently used to define the Astyanax scabripinnis species complex, with highest body depth approximately at vertical through pectoral-fin midlength and a low number of branched anal-fin rays (15–21). Astyanax lorien is distinguished from its congeners by a combination of characters, including presence of small bony hooks on all fins of mature males, a unique vertically elongate humeral blotch, and ventral margin of the third infraorbital close to the horizontal limb of preopercle. Astyanax rupestris is distinguished from its congeners also by a combination of characters, including ventral margin of third infraorbital distinctly separated from horizontal limb of preopercle, leaving a broad area without superficial bones, absence of midlateral stripe on body, absence of blotch on caudal peduncle, and presence of a vertically elongated conspicuous dark humeral blotch, not bordered anteriorly and posteriorly by distinct unpigmented areas. Both species are apparently endemic to the upper portion of the Paraguaçu river basin, within the Chapada Diamantina domain.

Full text

Accepted by F. Lima: 23 Apr. 2018; published: 22 Jun. 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2018 Magnolia Press
Zootaxa 4438 (3): 471
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http://www.mapress.com/j/zt/
Article
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https://doi.org/10.11646/zootaxa.4438.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:59A7C325-12E7-4BF4-891A-59796401CA49
Two new species of Astyanax Baird & Girard (Characiformes: Characidae)
from the upper rio Paraguaçu basin, Chapada Diamantina, Bahia, Brazil
ANGELA M. ZANATA
1,3
, RAFAEL BURGER
1
& PRISCILA CAMELIER
2
1
Instituto de Biologia, Universidade Federal da Bahia, Rua Barão de Geremoabo, 147, Ondina, 40170–290, Salvador, BA, Brazil.
E-mails: zanata.angela@gmail.com (AMZ); rafaelburger1984@gmail.com (RB)
2
Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04218–970, São Paulo, SP, Brazil.
E-mail: pricamelier@gmail.com
3
Corresponding author
Abstract
Astyanax lorien and A. rupestris are described from tributaries of upper rio Paraguaçu basin, Bahia, Brazil. Both species
shares the majority of the characters currently used to define the Astyanax scabripinnis species complex, with highest body
depth approximately at vertical through pectoral-fin midlength and a low number of branched anal-fin rays (15–21).
Astyanax lorien is distinguished from its congeners by a combination of characters, including presence of small bony
hooks on all fins of mature males, a unique vertically elongate humeral blotch, and ventral margin of the third infraorbital
close to the horizontal limb of preopercle. Astyanax rupestris is distinguished from its congeners also by a combination
of characters, including ventral margin of third infraorbital distinctly separated from horizontal limb of preopercle, leaving
a broad area without superficial bones, absence of midlateral stripe on body, absence of blotch on caudal peduncle, and
presence of a vertically elongated conspicuous dark humeral blotch, not bordered anteriorly and posteriorly by distinct
unpigmented areas. Both species are apparently endemic to the upper portion of the Paraguaçu river basin, within the Cha-
pada Diamantina domain.
Key words: Neotropical fishes, taxonomy, sexual dimorphism, lateral line
Resumo
Astyanax lorien e A. rupestris são descritas de tributários do alto curso da bacia do rio Paraguaçu, Chapada Diamantina,
Bahia, Brasil. Ambas compartilham a maioria dos caracteres utilizados atualmente para definir o complexo de espécies
Astyanax scabripinnis, com maior altura do corpo em torno da vertical que passa pela metade do comprimento da nada-
deira peitoral e baixo número de raios ramificados na nadadeira anal (15–21). Astyanax lorien difere dos seus congêneres
por vários caracteres, incluindo presença de pequenos ganchos ósseos em todas as nadadeiras de machos maduros, uma
única mancha umeral e margem ventral do terceiro infraorbital próxima ao ramo horizontal do pré-opérculo. Astyanax ru-
pestris difere dos seus congêneres por uma série de caracteres, incluindo margem ventral do terceiro infraorbital distante
do ramo horizontal do preopérculo, deixando uma ampla área sem ossos superficiais, ausência de faixa mediana lateral no
corpo, ausência de mancha escura no pedúnculo caudal e presença de uma mancha humeral alongada verticalmente e não
margeada anteriormente e posteriormente por áreas despigmentadas. Ambas as espécies são aparentemente endêmicas do
alto curso da bacia do rio Paraguaçu, dentro do domínio da Chapada Diamantina.
Palavras chave: Peixes neotropicais, taxonomia, dimorfismo sexual, linha lateral
Introduction
Astyanax Baird & Girard is one of the most speciose genus within Characidae, with about 150 species currently
considered as valid (Eschmeyer et al., 2018). It is also the more widespread genus of Neotropical characiforms,

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ranging from southeastern United States (Page & Burr, 2011) to Argentina (Almirón et al., 1997; Menni, 2004).
Astyanax has been proposed as a non-monophyletic genus based on both molecular and morphological
phylogenetic analyses (e.g., Mirande, 2010; Oliveira et al., 2011; Rossini et al., 2016) and its current definition is
still based on the pre-cladistic concept of Eigenmann (1917, 1921), who defined the genus based on a combination
of some of the most common character states in the Characidae (Zanata et al., 2017). Furthermore, it is one of the
most taxonomically poorly known genus within Characidae (Rossini et al., 2016). Many species belonging to the
genus are difficult to diagnose and several populations of Astyanax can currently only be identified at the generic
level.
The rio Paraguaçu is one of the most important drainage of the Northeastern Mata Atlântica freshwater
ecoregion (NMAF, ecoregion 328 sensu Abell et al., 2008). The basin is included into the North Group, a group of
eight drainages situated in the northernmost portion of the NMAF ecoregion proposed by Camelier & Zanata
(2014a), based on their shared fish species. Although sharing some fish species, the endemic component of the
ichthyofauna of the rio Paraguaçu basin is notable. It includes the endemic subfamily Copionodontinae de Pinna,
the monotypic genera Kalyptodoras Higuchi, Britski & Garavello and Myxiops Zanata & Akama, plus 22 endemic
species (see Camelier & Zanata, 2014a; Pereira & Zanata, 2014; Mattos et al., 2015; Zanata & Camelier, 2015;
Melo & Espíndola, 2016; Zanata et al., 2017). Consequently, the rio Paraguaçu basin possesses the highest number
of endemic species within Northeastern Mata Atlântica freshwater ecoregion. In fact, most of the species described
recently from this ecoregion are endemic to tributaries of the upper rio Paraguaçu basin (e.g., Astyanax brucutu
Zanata, Lima, Di Dario & Gerhard, A. hamatilis Camelier & Zanata, Characidium clistenesi Melo & Espíndola, C.
deludens Zanata & Camelier, Lepidocharax diamantina Ferreira, Menezes & Quagio-Grassioto, and Rhamdiopsis
krugi Bockmann & Castro).
More than ten species of Astyanax are reported to occur in the rio Paraguaçu basin but most of them cannot be
identified at the specific level and probably represent undescribed species (Santos & Caramaschi, 2007, 2011;
Camelier & Zanata, 2014b). Three Astyanax species, endemic to tributaries of the upper Paraguaçu, were recently
described: Astyanax brucutu, A. epiagos Zanata & Camelier, and A. hamatilis. Two additional species, also
apparently endemic to tributaries of the upper Paraguaçu basin, both sharing the majority of the characters listed by
Bertaco & Lucena (2006) to define the Astyanax scabripinnis species complex, as highest body depth
approximately at vertical through pectoral-fin midlength, a low number of branched anal-fin rays, presence of one
or two humeral spots, and body depth smaller than 41% of SL, are described herein.
Material and methods
Counts and measurements were taken according to Fink & Weitzman (1974) and Menezes & Weitzman (1990). In
Tables 1-2, standard length (SL) is expressed in mm and all other measurements are expressed as a percentage of
SL, except subunits of the head that are expressed as percentages of head length (HL). Meristic data are provided in
the description, an asterisk indicates counts of the holotype, and the frequency of each count is given in
parentheses. The number of horizontal scale rows between dorsal-fin origin and lateral line does not include scale
of predorsal series situated just anterior to first dorsal-fin ray or the short series of small scales just below dorsal-fin
base. Counts of vertebrae, supraneurals, procurrent caudal-fin rays, branchiostegal rays, gill rakers, and dentary
teeth were taken only from cleared and stained specimens (c&s), prepared according to Taylor & Van Dyke (1985).
Precaudal vertebrae include the Weberian apparatus and the vertebrae associated with ribs or haemal arches
without haemal spines. Caudal vertebrae are vertebrae associated with haemal spines. The fused first preural and
first ural centrum of the caudal region are counted as a single element. The pattern of circuli and radii was
examined on scales sampled from the region between the lateral line and the the dorsal-fin origin. The sex of
specimens was confirmed by dissection. Institutional abbreviations follow Fricke & Eschmeyer (2017).
Comparisons with species not cited in the ‘Comparative material examined’ were based on the literature
information (e.g., Eigenmann, 1921, 1927; Azpelicueta & Garcia, 2000; Bertaco & Malabarba, 2001; Melo, 2001;
Azpelicueta et al., 2002a; Azpelicueta et al., 2002b; Almirón et al., 2002; Haluch & Abilhoa, 2005; Bertaco &
Lucena, 2006; Triques et al., 2003; Abilhoa & Duboc, 2007; Alcaraz et al., 2009; Azpelicueta & Loureiro, 2009;
Bertaco & Lucena, 2010; Bertaco et al., 2010; Garavello & Sampaio, 2010; Tagliacollo et al., 2011; Oliveira et al.,
2013, 2017; Lucena et al., 2013; Ingenito & Duboc, 2014; Lucena et al., 2017).

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TWO NEW ASTYANAX FROM CHAPADA DIAMANTINA
Astyanax lorien, new species
(Figs. 1–2)
Holotype. MZUSP 123398, 66.4 mm SL, Brazil, Bahia State, Lençóis, Chapada Diamantina, stream tributary of
rio Mucugezinho, itself a tributary of rio Santo Antônio, rio Paraguaçu basin, 12°28’02’’S 41°24’56’’W, 734 m
above sea level (a.s.l.), 3 Mar 2017, A.M. Zanata, M.C.C. de Pinna, V. Abraão & V. Carvalho.
Paratypes. All from Brazil, Bahia State, Chapada Diamantina, rio Paraguaçu basin, rio Santo Antônio sub-
basin. Rio Santo Antônio, riacho Mãe Inácio: UFBA 8109, 10, 38.1–56.1 mm SL, Palmeiras, Arcado, near
Morro do Pai Inácio, 12º27’24”S 41º27’56”W, 863 m a.s.l., A.C.A. Santos. UFBA 8309, 22, 24.3–41.6 mm SL,
Palmeiras, Arcado, near Morro do Pai Inácio, 12º27’04”S 41º28’02”W, 826 m a.s.l., 28 Feb 2017, R. Burger.
UFBA 8346, 31, 22.8–38.6 mm SL, 1c&s, 40.5 mm SL, Palmeiras, Arcado, near Morro do Pai Inácio, 12º27’19”S
41º27’58”W, 831 m a.s.l., 3 Mar 2017, A.M. Zanata, M.C.C. de Pinna, V. Abraão & V. Carvalho. Rio
Mucugezinho: UFBA 8359, 12, 25.5–75.0 mm SL, 1 c&s, 63.5 mm SL; MZUSP 123399, 10, 34.7–71.8 mm SL;
MCP 53155, 10, 36.9–64.1 mm SL, same data as holotype. UFBA 6952, 6, 18.7–64.2 mm SL, Lençóis, near BR-
242, 12º27’42”S 41º25’02”W, 720 m a.s.l., 7 Jul 2011, A.M. Zanata, P. Camelier, J.O. Birindelli, R. Burger & B.
Sardeiro. UFBA 8114, 9, 46.7–63.5 mm SL, 11 Apr 2016, R. Burger; UFBA 8160, 12, 29.4–60.2 mm SL, 1 c&s,
50.6 mm SL, 26 Jun 2016, A.M. Zanata, R. Burger, P. Camelier & R. Freire; UFBA 8436, 4, 42.4–48.3 mm SL,
Palmeiras, Vale do Capão, riacho Águas Claras near Morrão, tributary of rio Mucugezinho, 12º31’18”S
41º28’52”W, 917 m a.s.l., 24 Jun 2017, A.M. Zanata & R. Burger. UFBA 8348, 5, 23.6–57.7 mm SL, Lençóis, near
BR 242, 12º27’57”S 41º26’13”W, 730 m a.s.l., 3 Mar 2017, A.M. Zanata, M.C.C. de Pinna, V. Abraão & V.
Carvalho. UFBA 8356, 16, 21.9–61.7 mm SL, 1 c&s, 51.1 mm SL; MZUSP 123400, 8, 39.4–58.0 mm SL,
Lençóis, near Morro do Pai Inácio, 12º28’29”S 41º27’18”W, 840 m a.s.l., 3 Mar 2017, A.M. Zanata, M.C.C. de
Pinna, V. Abraão & V. Carvalho. Rio São José and tributaries: MZUSP 49849, 2, 49.7–58.5 mm SL, Lençóis, rio
Lençóis, 3 km upstream of Lençóis city, above Cachoeira do Serrano, 12º34’S 41º22’W, 22 Feb 1995, A. Akama.
UFBA 7365, 23, 36.0–50.8 mm SL, 1 c&s, 42.6 mm SL, Lençóis, córrego Laranjeira, tributary of rio Lençóis,
12º33’48”S 41º24’06”W, 487 m a.s.l., 4 May 2013, A.M. Zanata, R. Burger, L. Oliveira & R. Abreu. UFBA 7780,
40, 32.9–57.2 mm SL, 1 c&s, 53.2 mm SL, Lençóis, rio Lençóis, 12º33’52”S 41º23’56”W, 465 m a.s.l., 30 Jan
2014, A.M. Zanata, R. Burger, L. Sales & R. Abreu. Rio Preto and tributaries: UFBA 5393, 2, 40.3–41.2 mm SL,
1 c&s, 42.1 mm SL, 20 Jun 2009, A.M. Zanata & R. Burger; UFBA 6931, 1, 57.8 mm SL, Palmeiras, Vale do
Capão, rio Riachinho, above Cachoeira do Riachinho, 12°34’20’’S 41°30’52’’W, 935 m a.s.l., 24 Jun 2011, A.M.
Zanata, P. Camelier & R. Burger. UFBA 5397, 18, 28.6–54.7 mm SL, 1 c&s, 34.3 mm SL; MZUSP 123401,10,
34.0–44.7 mm SL, Palmeiras, Vale do Capão, rio Preto above Cachoeira do rio Preto, 12º36’12”S 41º31’29”W, 800
m a.s.l., 25 Jun 2009, A.M. Zanata & R. Burger. UFBA 6691, 20, 15.6–43.9 mm SL, Palmeiras, Vale do Capão,
Córrego dos Gatos, tributary of rio do Capão, 12º36’44”S 41°29’35”W, 913 m a.s.l., 27 Jun 2010, A.M. Zanata &
R. Burger. UFBA 8161, 24, 19.9–48.5 mm SL, Palmeiras, Vale do Capão, stream tributary of rio Conceição dos
Gatos, 12º32’58”S 41º29’04”W, 960 m a.s.l., 26 Jun 2016, A.M. Zanata, R. Burger, P. Camelier & R. Freire. UFBA
8162, 18, 17.4–53.7 mm SL; NUP 19985, 8, 30.5– 53.5 mm SL, Palmeiras, Vale do Capão, stream tributary of rio
Conceição dos Gatos, 12º32’18”S 41º29’04"W, 960 m a.s.l., 26 Jun 2016, A.M. Zanata, R. Burger, P. Camelier &
R. Freire. UFBA 8351, 11, 28.9– 49.7 mm SL, Palmeiras, Vale do Capão, stream tributary of rio Conceição dos
Gatos, 12º31’52”S 41º29’58”W, 946 m a.s.l., 6 Mar 2017, A.M. Zanata, M.C.C. de Pinna, V. Abraão & V.
Carvalho.
Diagnosis. Astyanax lorien can be distinguished from most congeners, except A. brachypterygium Bertaco &
Malabarba, A. burgerai Zanata & Camelier, A. courensis Bertaco, Carvalho & Jerep, A. cremnobates Bertaco &
Malabarba, A. epiagos, A. eremus Ingenito & Duboc, A. goyanensis (Miranda-Ribeiro), A. guaricana Oliveira,
Abilhoa & Pavanelli, A. gymnogenys Eigenmann, A. jacobinae Zanata & Camelier, A. jenynsii (Steindachner), A.
joaovitori Oliveira, Pavanelli & Bertaco, A. jordanensis Alcaraz, Pavanelli & Bertaco, A. laticeps (Cope), A.
leonidas Azpelicueta, Casciotta & Almirón, A. microschemos Bertaco & Lucena, A. obscurus (Hensel), A. ojiara
Azpelicueta & Garcia, A. paranae Eigenmann, A. paris Azpelicueta, Almirón & Casciotta, A. pirabitira Lucena,
Bertaco & Berbigier, A. pirapuan Tagliacollo, Britzke, Silva & Benine, A. rivularis (Lütken), A. scabripinnis
(Jenyns), A. serratus Garavello & Sampaio, A. taurorum Lucena, Zaluski & Lucena, A. totae Haluch & Abilhoa, A.
troya Azpelicueta, Casciotta & Almirón, A. turmalinensis Triques, Vono & Caiafa and A. varzeae Abilhoa &
Duboc, by having a low number of branched anal-fin rays (16–21, mode =17) and highest body depth

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approximatelly at vertical through pectoral-fin midlength (vs. 21 or more branched anal-fin rays and highest body
depth just anterior to the dorsal-fin origin). Astyanax lorien can be distinguished from the aforementioned
congeners, except A. ojiara and A. troya, by having small bony hooks on all fins of mature males (vs. bony hooks
absent or not present on all fins of mature males; see item Discussion). The new species differs from A. ojiara, A.
troya, and further differs from aforementioned congeners (except A. courensis, A. epiagos, A. eremus, A.
goyanensis, A. gymnogenys, A. jacobinae, A. jenynsii, A. jordanensis, A. microschemos, A. paranae, A. rivularis, A.
totae, and A. varzeae) by having a unique vertically elongate humeral blotch, without horizontally elongate portion
(vs. two humeral blotches or one vertically elongate blotch with its central portion horizontally elongate). Astyanax
lorien further differs from A. brachypterygium, A. cremnobates, A. epiagos, A. eremus, A. gymnogenys, A. totae,
and A. varzeae by having ventral margin of the third infraorbital close to the horizontal limb of preopercle, with
narrow or no space between the two bones (vs. ventral margin of third infraorbital distinctly separated from
horizontal limb of preopercle, leaving a broad area without superficial bones), from A. courensis and A. jenynsii by
having usually 17–21 branched anal-fin rays, a single specimen with 16 (vs. 13–16 in A. courensis and 13-15 in A.
jenynsii), from A. goyanensis by having seven to eight branched pelvic-fin rays (vs. six), from A. jacobinae by
presence of a dark midlateral stripe (vs. absence), from A. jordanensis by the absence of bony hooks on fin-rays of
females (vs. presence), from A. microschemos by its deeper body (31.2–37.8 vs. 26.9–29.7% of SL), from A.
paranae by having 32–36 pored lateral-line scales (vs. 37–40), from A. rivularis by having humeral blotch roughly
ovate and almost completely located dorsally to the lateral line, with or without a short vertical line ventral to it (vs.
humeral blotch equally distributed above and below lateral line or, at least, with large portion below it).
FIGURE 1. Astyanax lorien: A. holotype, MZUSP 123398, 66.4 mm SL, Brazil, Bahia, Chapada Diamantina, Lençóis, stream
tributary of rio Mucugezinho, rio Paraguaçu basin; B. paratype, UFBA 8436, 48.3 mm SL, Brazil, Bahia, Palmeiras, Vale do
Capão, riacho Águas Claras, tributary of rio Mucugezinho, photographed alive.
Description. Morphometric data of holotype and paratypes in Table 1. Body moderately compressed and
elongated. Greatest body depth in the region approximatelly at vertical through midlength of pectoral fin. Dorsal
profile of head convex from upper lip to vertical through posterior nostrils; straight to slightly convex from latter
point to tip of supraoccipital spine, and slightly convex from this point to dorsal-fin origin. Body profile straight

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TWO NEW ASTYANAX FROM CHAPADA DIAMANTINA
and posteroventrally slanted from dorsal-fin origin to adipose fin and slightly concave to straight along caudal
peduncle. Ventral profile of head and body profile slightly convex from lower lip to anal-fin origin and straight and
posterodorsally slanted along anal-fin base. Ventral profile of caudal peduncle nearly straight to slightly concave.
Mouth terminal; upper jaw similar in length anteriorly or slightly longer than lower jaw. Posterior tip of
maxilla extending beyond anterior margin of orbit. Infraorbital series well developed, ventral margin of the third
infraorbital close to the horizontal limb of preopercle, with narrow or no space between the two bones. Number of
infraobitals variable from five to seven ossifications in six c&s specimens examined; one specimen with seven
elements on both sides of head (UFBA 7365, 42.6 mm SL), apparently due to split of the infraorbital four, two
specimens with six elements (UFBA 5393, 42.1 mm SL, and UFBA 8160, 50.6 mm SL), and three specimens with
five elements (UFBA 7780, 53.2 mm SL, UFBA 8346, 40.5 mm SL, and UFBA 8359, 63.5 mm SL) apparently
with infraorbitals three and four fused. Supraorbital absent.
Premaxillary teeth arranged in two rows (Fig. 2). Outer premaxillary row with three (25), four* (46) or five (2)
teeth bearing three or five cusps; inner row with four (44) or five* (39) teeth on both contralateral bones, bearing
three to seven cusps; symphyseal tooth of inner series asymmetrical, with one cusp on medial side, one larger
central cusp and two or three smaller on lateral side; remaining teeth asymmetrical or symmetrical; second teeth the
largest, with five to seven cusps; last tooth of inner premaxillary series with three to five cusps. Maxilla with one*
(55) or two (24) teeth bearing three or five cusps; anteriormost tooth largest. Dentary with seven (1), nine (2), 10
(2) or 11(1) teeth; anterior four or five teeth larger, usually symmetrical, with five or six cusps, followed by small
teeth with three or five cusps and posterior teeth with one or three cusps. Dentary teeth decreasing abruptly in size
posteriorly.
TABLE 1. Morphometric data for Astyanax lorien, new species; n= 43, range includes holotype. SD= standard
deviation.
holotype range mean SD
Standard length (mm) 66.4 32.1–75.4 52.3 -
Percentages of standard length
Depth at dorsal-fin origin 34.5 31.2–37.8 34.0 1.9
Snout to dorsal fin origin 54.5 50.7–56.1 53.8 1.3
Snout to pectoral-fin origin 25.6 24.3–30.3 27.3 1.3
Snout to pelvic-fin origin 46.7 46.0–50.5 48.1 1.3
Snout to anal-fin origin 63.3 62.1–66.8 64.7 1.3
Caudal peduncle depth 12.8 10.9–13.2 12.1 0.6
Caudal peduncle length 16.7 13.1–17.8 15.6 1.0
Pectoral-fin length 22.1 19.1–22.6 20.8 0.9
Pelvic-fin length 17.5 15.1–17.9 16.5 0.8
Dorsal-fin base length 12.8 11.9–14.5 13.2 0.7
Dorsal-fin heigth 25.5 23.9–28.3 25.7 1.1
Anal-fin base length 27.1 24.0–27.3 25.9 1.1
Anal-fin lobe length 16.1 16.1–20.2 18.1 1.2
Eye to dorsal-fin origin 40.5 37.6–42.1 40.2 1.1
Dorsal-fin origin to caudal-fin base 53.3 50.1–54.6 52.6 1.3
Head length 27.4 24.9–29.3 27.2 0.9
Percentages of head length
Horizontal eye diameter 30.2 28.2–34.7 31.8 1.6
Snout length 23.6 23.0–26.9 25.1 1.2
Least interorbital width 33.5 28.0–33.5 30.7 1.4
Upper jaw length 44.5 40.3–44.5 42.6 1.4

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FIGURE 2. Upper and lower left jaws of Astyanax lorien (UFBA 8356, 51.1 mm SL, paratype) in lateral view.
Scales cycloid, circulli absent on exposed area of scales, with various parallel or slightly divergent radii
extending to posterior margin of scales. Lateral line slightly decurved ventrally. Scales with pores on lateral line
series variably; 36 specimens with completely pored lateral line, having with 32*(1), 33 (3), 34 (16), 35 (14) or 36
(2) pored scales; remaining specimens (27) with pored scales interspersed with non-pored ones (discontinuous
lateral line) of variable pattern. Horizontal scale rows between dorsal-fin origin and lateral line five* (50) or six (8).
Horizontal scale rows between lateral line and pelvic-fin insertion four* (40) or five (13). Scales along middorsal
line between tip of supraoccipital process and origin of dorsal fin 10* (16), 11(27) or unordered (5). Horizontal
scale rows around caudal peduncle 13 (1), 14* (49) or 15 (3). Base of anteriormost anal-fin rays with single scale
row composed of three to seven scales.
Dorsal-fin rays ii* (56), eigth (3) or nine* (53). Distal margin of dorsal fin straight or slightly rounded. Dorsal-
fin origin slightly anterior or approximately at middle of SL. Base of last dorsal-fin ray aligned with base of
anteriormost branched anal-fin rays. First dorsal-fin pterygiophore inserting behind neural spine of ninth (4) or
tenth (2) vertebra. Adipose fin present. Anal-fin rays iv* (53), 16 (1), 17 (9), 18 (12), 19* (14), 20 (14) or 21 (3).
Distal margin of anal fin slightly concave to nearly straight. First anal-fin pterygiophore inserting behind haemal
spine of 17
th
(5) or 19
th
(1) vertebra. Pectoral-fin rays i* (54), 11 (13), 12* (26) or 13 (15). Tip of pectoral fin falling
slightly short or slightly surpassing vertical through pelvic-fin insertion. Pelvic-fin rays i* (57), 7* (55) or 8 (2).
Caudal fin forked, lobes rounded, similar in size. Caudal-fin rays i9+8i (4), i9+7i (1) or i8+8i (1). Eight (2), 9 (1) or
10 (1) dorsal procurrent caudal-fin rays and 7 (1) or 8 (4) ventral procurrent caudal-fin rays.
Total number of vertebrae 33 (4), 34 (1) or 35 (1); precaudal vertebrae 14 (2) or 15 (4); caudal vertebrae 18 (2),
19 (3) or 20 (4). Supraneural bones four (1), five (4) or six (1). A single uroneural (6). Branchiostegal rays four (6),
three connected to anterior ceratohyal, one connected to posterior ceratohyal or region between anterior and
posterior ceratohyal. First gill arch with 2 (6) gill rakers on hypobranchial, 8 (6) rakers on ceratobranchial, one (6)
raker on intermediate cartilage, and 5 (2), 6 (3) or 7 (1) rakers on epibranchial.

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Color in alcohol. Overall ground color tan, dorsal portion of body darker (Fig. 1a). Guanine pigmentation
present on infraorbitals of recently fixed specimens. Dark chromatophores densely concentrated on dorsal surface
of head from upper lip to supraoccipital spine. Middorsal series of scales dark. Small dark chromatophores present
on anterior third of maxilla, proximal area of orbit, preopercle, and opercle. Large dark chromatophores sparsely
distributed under skin of third infraorbital. Ventral portion of head less pigmented, with scattered small dark
chromatophores more concentrated anteriorly and on lateral borders of head. Scales of lateral portion of body with
dark chromatophores concentrated along its posterior margin, forming a regular dark border and resulting in an
overall reticulate pattern. Central portions of scales poorly pigmented, with smaller chromatophores sparsely
distributed. Humeral region with an inconspicuous, slightly vertically-elongated blotch; blotch almost completely
dorsal to the lateral line and occupying up to two scales wide and up to two scales deep; most specimens without
ventral elongation of the blotch, some specimens with a narrow and short ventral elongation, approximatelly one
scale long. Humeral blotch usually not bordered anteriorly or posteriorly by distinct clear areas. Midlateral dark
stripe present, usually poorly visible on anterior half of body, widening at caudal peduncle, forming a conspicuous
dark horizontally elongate blotch. Caudal peduncle blotch extending from vertical through adipose-fin origin to
distal margin of median caudal-fin rays. Humeral and caudal blotches formed by deeply-lying dark pigmentation.
All fins darkened; dorsal fin with membranes on proximal portion of rays distinctly darker; pectoral, pelvic,
and caudal fins with melanophores mainly on borders of rays; anal-fin with melanophores throughout interradial
membranes, with distal border darker in some specimens. Adipose fin with sparse small dark chromatophores.
Color in life. Freshly collected specimens with overall golden coloration; a silvery hue on infraorbitals,
opercle, and on sides of body (Fig. 1b). Dorsal portion of head and body somewhat darker; belly whitish. Iris
yellow. Humeral blotch poorly visible or not visible at all. Inconspicuous horizontally elongated dark blotch on
caudal peduncle, continuing or not up to the end of middle caudal-fin rays. Anal and caudal fins with reddish-
orange overall coloration. Caudal fin with yellowish rays bordering the central dark area. Dorsal fin similarly
colored to anal and caudal fins, although more yellowish on its proximal half. Pectoral fins yellowish and pelvic
fins orange- to reddish. Adipose fin yellow- or orangish.
Sexual dimorphism. Bony hooks on all fins were observed in mature males of Astyanax lorien. Pectoral fin
with hooks on the dorsal margin from the unbranched up to the tenth branched ray, distributed usually on distal half
of ray, but in some rays along its entire length. Up to 24 hooks per ray, distributed on both branches of rays but
more commonly on the inner side of rays. Pelvic fins with very small hooks present in the ventral surface of all
branched pelvic-fin rays of mature males, from the area near the branching points to their tips. Up to 15 hooks
present per ray, distributed on both main branches of bifurcation. Up to 20 well developed hooks present from the
last unbranched up to the 19
th
branched rays of the anal fin. Two lateral-dorsally directed bony hooks present per
segment, distributed from near the first bifurcation of fin rays to their distal tips, on both branches or just on the
posterior one; hooks more numerous on anteriormost rays, decreasing in number posteriorly. Very small hooks also
occur in the longer unbranched and from the first to the eighth branched dorsal-fin ray, from near the first
bifurcation of fin rays to their distal tips, mainly on posterior border of rays. Very small hooks are also present
laterally, in the centralmost four rays on the caudal fin of some mature males. The presence of bony hooks on the
pectoral and anal fins of the species is the most common condition observed. Gill glands (Burns & Weitzman,
1996) were not found on first gill arch of both sexes.
Distribution. Astyanax lorien is known from tributaries of the upper rio Paraguaçu, particularly from the rio
Santo Antônio sub-basin (Fig. 3), within the Chapada Diamantina domain, Bahia, Brazil.
Habitat and ecological notes. Astyanax lorien inhabits transparent and blackwater tributaries of the rio
Paraguaçu basin (Fig. 4). These tributaries are perennial and torrential in the rainy season, with headwaters around
1,000 m a.s.l., running on the eastern slopes of the Sincorá Range, partially within the Parque Nacional da Chapada
Diamantina (PNCD), in areas of “Campo Rupestre”, a herbaceous-shrub vegetation that is present at elevations of
about 900 m on sandstones and quartzite, usually under moister weather conditions than in the surrounding
Caatinga vegetation (Queiroz, 2006). The species was collected in both pools and lotic stretches running over
rocky beds, alternate shallow rapids and waterfalls in altitudes of 465–960 m a.s.l. The pools sampled have variable
dimensions and depth, mainly rocky bottoms and, less frequently, pebbles and sand (at the shallow borders of
pools). Astyanax lorien occurs sintopically with Copionodon sp., Hyphessobrycon negodagua Lima & Gerhard,
Myxiops aphos Zanata & Akama, Hoplerythrinus unitaeniatus (Spix & Agassiz), and Trichomycterus sp.

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FIGURE 3. Map of the upper rio Paraguaçu basin, Brazil, Bahia, showing the collection sites and type locality of Astyanax
lorien (yellow and red circles, respectively), and of Astyanax rupestris (white and black circles, respectively).
FIGURE 4. Sampling locality of Astyanax lorien, Brazil, Bahia, Chapada Diamantina, Palmeiras, Vale do Capão, riacho Águas
Claras near Morrão, rio Paraguaçu basin.

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The analysis of the stomach contents of three specimens of A. lorien (UFBA 7780, UFBA 8114, and UFBA
8353) revealed the presence of allochthonous and autochthonous items, composed of filamentous algae, fragments
of vascular plants and seeds, organic debris, Trichoptera larvae, fragments of adults of terrestrial insects
(Hymenoptera: Formicidae and other unidentified orders).
Etymology. The name lorien, from the Quenya [ˈkʷwɛnja] language meaning “Dream Land”, is used in
allusion to the beautiful areas on the Chapada Diamantina inhabited by the species. The Quenya is a fictional
language devised by J. R. R. Tolkien and spoken by the Elves. A name in apposition.
Astyanax rupestris, new species
(Figs. 5–6)
Astyanax sp. Zanata & Primitivo, 2013: 209 [citation; co-occurence with Copionodon pecten].
Holotype. MZUSP 89567, 85.2 mm SL, Brazil, Bahia State, Chapada Diamantina, Andaraí, Igatu, rio Coisa Boa,
tributary of rio Piabas, Rio Paraguaçu basin, 12°53’55’’S, 41°19’01’’W, 710 m a.s.l., 9 May 2009, A.M. Zanata, R.
Burger & C. Primitivo.
Paratypes. All from Brazil, Bahia State, Chapada Diamantina, rio Paraguaçu basin. Rio Cumbuca sub-basin:
MZUSP 38537, 72, 30.3–63.7 mm SL, 4 c&s, 49.8–65.2 mm SL, Mucugê, rio Mucugê, 26 Sep 1987, M.T.
Rodrigues. UFBA 2789, 19, 24.3–66.0 mm SL, Mucugê, Parque Municipal de Mucugê, rio Cumbuca, 13°01’56’’S
41°20’29’’W, 1,028 m a.s.l., 3 Dec 2005, A.M. Zanata, M.P. Geraldes & P. Camelier. UFBA 2790, 19, 39.9–80.1
mm SL, 2 c&s, 55.5–57.4 mm SL, Mucugê, rio Cumbuca, 13°01’36’’S 41°20’32’’W, 1,008 m a.s.l., 22 Aug 2005,
A.M. Zanata, P. Camelier, T. Chinelli, J.G.B. Lima & M.M. Nogueira. UFBA 2791, 1, 67.0 mm SL, rio Mucugê,
12°59’36’’S 41°21’01’’W, 903 m a.s.l., 24 Aug 2005, A.M. Zanata, P. Cameier, T. Chinelli, J.G.B. Lima & M.M.
Nogueira. UFBA 8112, 3, 42.6–55.7 mm SL, Mucugê, rio Cumbuca, near Cachoeira Andorinhas, 13°01’08’’S
41°20’34’’W, 970 m a.s.l., 2 Jan 2007, M.C.C. de Pinna & A.A. Oliveira. Rio Piabas sub-basin: UFBA 5288, 25,
37.9–81.0 mm SL, 2 c&s, 61.6–64.7 mm SL; NUP 19986, 8, 43.5– 68.6 mm SL; MCP 53156, 8, 42.1–73.7 mm
SL, same data as holotype. MNRJ 21985, 26, 30.3–102.3 mm SL, 16 Mar 1999; MNRJ 22698, 9, 46.6–80.3 mm
SL, Mucugê, rio Piabas, 12°57’00’’S 41°16’39’’W, 812 m a.s.l., 11 Nov 1999, A.C.A. Santos.
Diagnosis. Astyanax rupestris can be distinguished from most congeners, except A. brachypterygium, A.
burgerai, A. courensis, A. cremnobates, A. epiagos, A. eremus, A. goyanensis, A. guaricana, A. gymnogenys, A.
jacobinae, A. jenynsii, A. joaovitori, A. jordanensis, A. laticeps, A. leonidas, A. lorien, A. microschemos, A.
obscurus, A. ojiara, A. paranae, A. paris, A. pirabitira, A. pirapuan, A. rivularis, A. scabripinnis, A. serratus, A.
taurorum, A. totae, A. troya, A. turmalinensis and A. varzeae, by having a lower number of branched anal-fin rays
(15–20, mode=20) and highest body depth approximatelly at vertical through pectoral-fin midlength (vs. 21 or
more branched anal-fin rays and highest body depth just anterior to the dorsal-fin origin). The new species can be
distinguished from the aforementioned congeners, except A. brachypterygium, A. cremnobates, A. epiagos, A.
eremus, A. gymnogenys, A. totae, and A. varzeae, by having ventral margin of third infraorbital distinctly separated
from horizontal limb of preopercle, leaving a broad area without superficial bones (vs. ventral margin of the third
infraorbital close to the horizontal limb of preopercle, with narrow or no space between the two bones). Astyanax
rupestris can be distinguished from most aforementioned congeners, including A. brachypterygium,
A. cremnobates, A. epiagos, A. eremus, A. gymnogenys, A. totae and A. varzeae, by the absence of a midlateral dark
stripe on body or caudal-peduncle blotch (vs. presence). It can be further distinguished from A. eremus, A.
gymnogenys, and A. varzeae by having 11–13 circumpeduncular scales (vs. 16–19 in A. eremus and A.
gymnogenys, and 14–16 in A. varzeae), from A. brachypterygium and A. cremnobates by having only one humeral
blotch (vs. two), and from A. epiagos by having base of the anteriormost anal-fin rays covered by a row of scales
(vs. absence of row of scales on the base of the anal fin). The new species is further distinguished from A.
jacobinae, the single congener occurring in a adjacent river system (the rio Itapicuru) that also lacks a conspicuous
midlateral stripe, by the absence of a blotch over caudal peduncle (vs. presence of an inconspicuous caudal-
peduncle blotch extending to the middle caudal-fin rays), presence of a narrower humeral blotch horizontally
covering partially two or three horizontal scales and not surrounded by less pigmented areas (vs. humeral blotch
covering completely at least two horizontal scales and surrounded by clearer areas), and a smaller eye diameter
(21.7–34.1% vs. 36.8–40.3% of HL).

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Description. Morphometric data of holotype and paratypes in Table 2. Body moderately compressed and
elongated. Greatest body depth around vertical through midlength of pectoral fin. Dorsal profile of head convex
from upper lip to vertical through posterior nostrils; straight to slightly convex from latter point to tip of
supraoccipital spine, and slightly convex from this point to dorsal-fin origin. Body profile straight and
posteroventrally slanted from dorsal-fin origin to adipose fin and slightly concave to straight along caudal
peduncle. Ventral profile of head an body profile slightly convex from lower lip to anal-fin origin and straight and
posterodorsally slanted along anal-fin base. Ventral profile of caudal peduncle nearly straight to slightly concave.
Mouth terminal; upper jaw similar in length anteriorly or slightly longer than lower jaw. Posterior tip of
maxilla extending beyond anterior margin of orbit. Infraorbital series moderately developed; ventral margin of
third or third plus four infraorbitals falling distinctly short from horizontal limb of preopercle and leaving a
relatively broad area without superficial bones (Fig. 5). Number of infraobitals variable from four to six
ossifications in the c&s specimens examined; one specimen with four elements on both sides of head (UFBA 2790,
55.5 mm SL) apparently with infraorbitals one and two fused resulting in one element, three and four fused and
five and six independent; three specimens with five elements apparently due to the fusion of infraorbitals three and
four (UFBA 2790, 57.4 mm SL; UFBA 5288, 61.6 mm SL; MZUSP 38537, 49.9 mm SL), and four specimens with
six infraorbitals. Supraorbital absent.
Premaxillary teeth arranged in two rows (Fig. 6). Outer premaxillary row with two (2), three* (31) or four (22)
teeth bearing three or five cusps; inner row with four (2) or five* (52) teeth bearing four to seven cusps;
symphyseal tooth of inner series asymmetrical, with one cusp on medial side, one larger central cusp and two
smaller on lateral side; remaining teeth asymmetrical or symmetrical; second teeth the largest, with six or seven
cusps; last tooth of inner premaxillary series with four or five cusps, distinctly smaller than penultimate tooth in
some specimens. Maxilla with one (28), two* (23) or three (4) teeth bearing three or five cusps; anteriormost tooth
largest. Dentary with nine (1), 10 (1) or 11 (2) teeth; two anterior teeth larger, usually symmetrical, with six or
seven cusps, followed by teeth with three or five cusps and posterior teeth with one or three cusps. Dentary teeth
decreasing abruptly in size posteriorly.
Scales cycloid, circulli absent on exposed area of scales, with various parallel or slightly divergent radii
extending to posterior margin of scales. Lateral line slightly decurved ventrally, completely pored from
supracleithrum to base of caudal fin, with 35 (1), 36 (5), 37* (19), 38 (22) or 39 (1) pored scales. Horizontal scale
rows between dorsal-fin origin and lateral line six* (24) or seven (29), not including scale of predorsal series
situated just anterior to first dorsal-fin ray. Horizontal scale rows between lateral line and pelvic-fin insertion 5*
(44) or 6 (8). Scales along middorsal line between tip of supraoccipital process and origin of dorsal fin 11 (19), 12*
(24) or 13 (6). Horizontal scale rows around caudal peduncle 14* (23), 15 (23) or 16 (6). Base of anteriormost anal-
fin rays with single scale row composed of five to seven scales.
Dorsal-fin rays ii* (55), 8 (5) or 9* (50). Distal margin of dorsal fin straight or slightly rounded. Dorsal-fin
origin approximately at middle or slightly posterior of SL. Base of last dorsal-fin ray anterior to or aligned with
anal-fin origin. First dorsal-fin pterygiophore inserting behind neural spine of 9
th
(2) or 10
th
(4) vertebra. Adipose
fin present. Anal-fin rays iv* (54), 15 (1), 16 (5), 17 (21), 18* (19), 19 (8) or 20 (1). Distal margin of anal fin
slightly concave to nearly straight. First anal-fin pterygiophore inserting behind haemal spine of 17
th
(2) or 18
th
(4)
vertebra. Pectoral-fin rays i* (54), 11* (11), 12 (31) or 13 (12). Tip of pectoral fin falling slightly short of vertical
through pelvic-fin insertion. Pelvic-fin rays i* (55), 6 (2) or 7* (53). Caudal fin forked, lobes usually somewhat
pointed, similar in size. Caudal-fin rays i9+8i (6). Nine (2), 10 (1), 11 (1), 12 (1) or 13 (1) dorsal procurrent caudal-
fin rays, and 8 (1), 9 (2), 10 (1), 11 (1) or 12 (1) ventral procurrent caudal-fin rays.
Total number of vertebrae 34 (1), 36 (2) or 37 (3); precaudal vertebrae 15 (2), 16 (3) or 17 (1); caudal vertebrae
19 (1), 20 (3), 21 (1) or 22 (1). Supraneural bones 4 (2) or 5 (3). A single uroneural (6). Branchiostegal rays four
(6), three connected to anterior ceratohyal, one connected to posterior ceratohyal or region between anterior and
posterior ceratohyal. First gill arch with one (3) or two (3) gill rakers on hypobranchial, 10 (4) or 11(2) rakers on
ceratobranchial, one (6) rakers on intermediate cartilage, and seven (6) rakers on epibranchial.
Color in alcohol. Overall ground color tan, usually darker dorsally (Fig. 5a). Guanine pigmentation present on
infraorbitals of some specimens. Dark chromatophores densely concentrated on dorsal surface of head from upper
lip to supraoccipital spine. Middorsal series of scales dark. Small dark chromatophores present on maxilla, less
concentrated on its distal portion. Similar minute dark chromatophores concentrated on surface of infraorbitals, and
unossifed area between ventral border of infraorbitals and preopercle. Large dark chromatophores under skin of

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third infraorbital. Some populations with overall coloration darker and sides of head with higher concentration of
dark chromatophores. Ventral portion of head light brown, with scattered small dark chromatophores. Scales of
lateral portion of body with dark chromatophores concentrated along its posterior margin, resulting in an overall
reticulate pattern. Central portions of scales usually with smaller chromatophores sparsely distributed. Humeral
region with a vertically-elongated, relatively narrow blotch, approximatelly two scales wide and extending from
two or three scales above lateral line and two scales below it. Humeral blotch not bordered anteriorly or posteriorly
by clear areas. Midlateral dark stripe absent; some specimens with faint dark, deep-lying pigmentation on skin
under midlateral scales from vertical through adipose-fin insertion to caudal peduncle, but never forming a distinct
stripe or blotch on caudal peduncle. Some specimens, usually up to 28.0 mm SL (UFBA 5288), with sparse dark
superficial chromatophores on central portion of peduncle, but never forming a distinct blotch. Thin longitudinal
line formed by subjacent dark pigmentation along horizontal septum, from approximately the vertical through
middle portions of dorsal fin to caudal peduncle.
All fins darkened; dorsal, anal, and caudal fins with small dark chromatophores on interradial membranes,
mainly along borders of rays. Dorsal and anal fins with dark pigmentation distinctly more developed on distal half
of fin rays in most specimens; otherwise pigmentation of uniform intensity across fin. Caudal fin similarly
darkened; larger dark chromatophores along middle four to six caudal-fin rays and on border of adjacent rays of
most individuals; dark pigmentation resulting in an inconspicuous central dark “v” towards posterocentral portion
of fin. Adipose fin covered with small dark chromatophores.
Color in life. Freshly collected specimens had overall dark body coloration, with a plumbeous hue on sides of
body and at infraorbitals three and four (Fig. 5b). Borders of scales of some specimens and borders of lateral line
pores orangish. Infraorbital 1-2 and 6, region between infraorbitals and preopercle, opercle, maxilla, and ventral
portion of body tinged with yellow. Iris yellowish-orange. Humeral blotch visible. Fins usually with reddish-orange
overall coloration, lighter on borders, and with somewhat dark rays. Anal fin of some specimens with dark
chromatophores more concentrated distally on rays, forming a dark border; interradial membranes and distal half of
unbranched and first branched anal-fin rays of some specimens whitish. Middle caudal-fin rays darkened and
posterior border also usually dark.
Sexual dimorphism. Bony hooks on anal and pectoral-fin rays are present in mature males of Astyanax
rupestris, and, more rarely, on the pelvic-fin rays. Anal fin with hooks usually from the last unbranched to the tenth
branched rays in several specimens, up to 15th ray in one specimem (MZUSP 38537, 66.6 mm SL). Up to 15 hooks
are present on the lateral portion of the distal half of each ray, being more numerous on the anteriormost rays and
decreasing in number in the posteriormost rays. Pectoral fins with distal third of the three to five longest branched
pectoral-fin rays with up to 14 dorsally-projected hooks, usually situated on the medial side of rays and distributed
from area anterior to the first branching point to tip of rays. Pelvic fin of one specimen (MZUSP 38537, 66.6 mm
SL) with one to 11 hooks in all branched pelvic-fin rays; hooks situated on medial side of rays, posterior to the first
bifurcation. Remaining specimens examined with hooks on anal- and pectoral-fin rays lacking bony hooks on the
pelvic fin.Tiny hooks on three centralmost caudal-fin rays observed only on the aforementioned specimen. Gill
glands (Burns & Weitzman, 1996) were not found on first gill arch of both sexes.
Distribution. Astyanax rupestris is only known from the rio Cumbuca and rio Piaba sub-basins, upper rio
Paraguaçu basin, within the Chapada Diamantina domain, Bahia, Brazil (Fig. 3).
Habitat and ecological notes. Astyanax rupestris inhabits blackwater tributaries of the rio Paraguaçu basin
(Fig. 7), in environments similar to those described to A. lorien. Particularly, the rio Cumbuca is within the Parque
Municipal de Mucugê. Locally, Andaraí and Mucugê municipalities have a comparatively high precipitation,
around 1,000 mm/year, possibly due to the influence of the Serra do Sincorá as an orographic barrier (Azevedo &
Silva, 2000; Barreto, 2010). Astyanax rupestris occurs with the endemic trichomycterids Copionodon pecten de
Pinna and C. orthiocarinatus de Pinna (see Zanata & Primitivo, 2013). The presence of Hoplias malabaricus
(Bloch) was reported to occur in these sites by local dwellers and Hoplerythrinus unitaeniatus was sampled in rio
Mucugê, although not syntopycally with A. rupestris.
The analysis of the stomach contents of six c&s specimens of A. rupestris revealed the presence of
allochthonous and autochthonous items, composed of filamentous algae, fragments of Podostemataceae, fragments
of vascular plants and seeds, organic debris, terrestrial and aquatic Hemiptera (Cicadellidae and Notonectidae,
respectively), Trichoptera adults and larvae, Odonata nymphs, fragments of adults of terrestrial insects
(Hymenoptera: Formicidae and other unidentified orders) and of other unidentified arthropods.

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FIGURE 5. Astyanax rupestris: A. holotype, MZUSP 89567, 85.2 mm SL, Brazil, Bahia, Chapada Diamantina, Andaraí, Igatu,
rio Coisa Boa, tributary of rio Piabas, Poço da Madalena, rio Paraguaçu basin; B. paratype, UFBA 2789, 59.0 mm SL, Brazil,
Bahia, Mucugê, Parque Municipal de Mucugê, rio Cumbuca, rio Paraguaçu basin, photographed soon after fixation.
FIGURE 6. Upper and lower left jaws of Astyanax rupestris (MZUSP 38537, 55.3 mm SL, paratype) in lateral view.

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Etymology. The name rupestris, from the Latin, meaning rock-dwelling, refers to the rocky bottom rivers
inhabited by the species and rocky environment characteristic of the Chapada Diamantina.
Remarks. The rio Cumbuca sub-basin is composed by rio Cumbuca itself, which is a direct affluent of the rio
Paraguaçu, and its two tributaries, the rio Mucugê and the rio Piabinha. Except by their extensions, with the rio
Piabinha being distinctly shorter (8.6 km vs. 23.7 km of rio Cumbuca and 21.3 km of rio Mucugê), these rivers are
very similar to each other in physical characters, sharing the dark type of water, rocky substrate, and marginal
vegetation. A similar morphotype of Astyanax occurs in these three rivers. However, while populations of the rio
Cumbuca and rio Mucugê rivers are identified herein as A. rupestris, the one from the rio Piabinha is treated as A.
aff. rupestris (see ‘Comparative material examined’). These specimens are very similar morphologically to A.
rupestris but possess variations in some meristic characters that were not observed in specimens from the rio
Cumbuca and rio Mucugê. These include a high frequency of specimens with variable lateral-line perforation, four
premaxillary teeth in the inner row, and reduction in the number of branched dorsal- and pelvic-fin rays.
Morphometric data do not vary significantly among specimens from rio Piabinha when compared with specimens
identified herein as A. rupestris. Based on these differences, we prefer to consider the population from the rio
Piabinha as Astyanax aff.. rupestris. Further studies on this population using methods of integrative taxonomy,
including both molecular and morphological data, are currently under way by one of the authors (RB) and
hopefully will clarify the taxonomic status of this population.
Discussion
As mentioned previously, Astyanax lorien and A. rupestris shares the majority of the characters listed by Bertaco &
Lucena (2006) to define the Astyanax scabripinnis species complex. However, the dark midlateral body stripe
extending to the tip of the middle caudal-fin rays, listed by the authors as one of the features delimiting that species
complex, is absent in A. rupestris. In fact, a brief analysis of the list of species currently included in the Astyanax
scabripinnis species complex (Ingenito & Duboc, 2014) reveals that some of them lacks one or another of the
characters listed by Bertaco & Lucena (2006) to the group. According to Alcaraz et al. (2009), A. epiagos and A.
jacobinae does not fit all the characters delimited for the complex since they lack the midlateral dark body stripe.
Similarly, A. intermedius Eigenmann and A. ita Almirón, Azpelicueta & Casciotta, instead of having the highest
body depth approximatelly at vertical through pectoral-fin midlength, as expected to members of the
aforementioned species complex, possess the highest body depth at the dorsal-fin origin. As discussed by several
authors (e.g., Bertaco & Malabarba, 2001; Abilhoa & Duboc, 2007; Lucena et al., 2017 ), the A. scabripinnis
species complex lacks a phylogenetic diagnosis and possibly does not represent a monophyletic unit. However,
Lucena et al. (2017) remarked that, although possibly artificial, the recognition of a Astyanax scabripinnis species-
group facilitates discussions and comparisons within the species-rich genus Astyanax and because of that, we retain
it here as well. Future phylogenetic studies incorporating a larger sampling of the diversity of Astyanax are required
to provide a comprehensive hypothesis of relationships within the genus and, thus, a better delimitation of its
species groups.
Intraspecific variation in some morphological characters was observed in Astyanax lorien and A. rupestris.
Both species possess variations in the number of infraorbitals and, in the second species, in the degree of reduction
in size of the third infraorbital. In A. lorien, a reduction in the number of ossifications is observed, with five to
seven infraobitals in six c&s specimens examined. In A. rupestris, three of the six c&s specimens examined
(UFBA 2790, 57.4 mm SL; UFBA 5288, 61.6 mm SL; and MZUSP 38537, 49.8 mm SL) have only five
infraorbitals, in one or both sides of the head, with one large bony element in the position primitively occupied by
infraorbitals third and fourth. Another specimen (UFBA 2790, 55.5 mm SL) possess four of those ossifications on
both sides of head, possibly resulting from fusion of third and fourth infraorbital, similarly to the aforementioned
specimens, plus fusion of first and second infraorbitals. Reduction in number of infraorbitals are reported for
several other small characids (e.g., Zanata & Camelier, 2008; Ingenito & Duboc, 2014; Zanata et al., 2017). Fusion
of infraorbital bones in small characids from headwaters of Chapada Diamantina was previously described in the
congeners A. brucutu (Zanata et al., 2017), A. epiagos and A. jacobinae (Zanata & Camelier, 2008), and also to
Myxiops aphos (Zanata & Akama, 2004). The species of Astyanax described herein show similarities with the
infraorbital arrangements of Myxiops (see fig. 4b of Zanata & Akama, 2004), although in the last taxa the number

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of remaining infraorbitals ranges from three to five. Although sharing the fusion of infraorbital bones, the species
of Astyanax in question do not share the autapomorphies utilized to define Myxiops (Zanata & Akama, 2004: 46),
and are distinguished from A. epiagos and A. jacobinae by features already mentioned in the ‘Diagnosis’ sections.
Apart from the number of ossifications, the ventral margin of third infraorbital (or the ossification apparently
formed by the fusion of the third and fourth infraorbitals) is distinctly separated from the horizontal limb of the
preopercle in A. rupestris, leaving, consequently, a relatively broad area without superficial bones in the cheek
(Fig. 5). As mentioned in the diagnosis of the species, a similar condition also occurs in A. brachypterygium, A.
cremnobates, A. epiagos, A. eremus, A. gymnogenys, A. totae, and A. varzeae. In addition, a broad space between
the infraorbitals and the preopercle also occurs in the congeners A. brucutu and A. taeniatus (Jenyns). On the other
hand, in A. lorien the ventral margin of the third infraorbital is always close to the horizontal limb of preopercle,
with narrow or no space between the two bones.
In Astyanax lorien, 44 of 83 specimens examined present four inner premaxillary teeth on both contralateral
bones, instead of the five teeth typically found in Astyanax. Additionally, A. lorien presents a variable lateral-line
perforation, with the majority of the specimens having completely pored lateral lines (the pattern typically found to
Astyanax) and around 40% of the remaining specimens examined with pored scales interspersed with non-pored
ones. The presence of a variable lateral-line perforation within a species is an unusual or poorly documented
condition among the species of Characidae, known for some species of Hemigrammus Gill (Marinho et al., 2014)
and Moenkhausia Eigenmann (Marinho & Langeani, 2010). Among congeners, the presence of a variable lateral-
line perforation was observed in A. brachypterygium, A. cremnobates, and A. rivularis (C. Oliveira, pers. comm.).
Variation in the lateral-line perforation in Astyanax rupestris was restricted to the population identified herein as
Astyanax aff. rupestris from the rio Piabinha. In this population, the great majority of the specimens examined
present reductions in some morphological features, including lateral line incomplete or discontinuous, four
premaxillary teeth in the inner row, and reduction in the number of branched dorsal- and pelvic-fin rays (see
Remarks). According to Weitzman & Vari (1988), a series of morphological reductions is commonly found in
miniature species (i.e. adipose-fin loss, reduction of the degree of development of the laterosensory canal system,
bone losses and simplifications, among others). However, the morphological reductions observed in A. lorien and
Astyanax aff. rupestris are likely not related to miniaturization, since the examined specimens of the former species
do not fit Weitzman & Vari´s (1988) criteria of a maximum 26 mm SL to define miniature fishes (maximum
observed of 75.4 mm SL in A. lorien and 75.6 mm SL in A. rupestris).”
Camelier & Zanata (2014b) summarized the information on the occurrence of bony hooks in all fins of mature
males of Astyanax species, highlighting the derived nature of this condition as proposed by Casciotta & Almirón
(2004). Within this genus, presence of bony hooks on all fins of mature males was reported by Almirón et al.
(2010) to seven species from Argentina and/or Uruguay: A. chico Casciotta & Almirón, A. hermosus Miquelarena,
Protogino & Lòpez, A. ojiara, A. pynandi Casciotta, Almirón, Bechara, Roux & Ruiz Diaz, A. rutilus (Jenyns), A.
stenohalinus Messner, and A. troya. More recently, that condition was also reported for A. bagual Bertaco & Vigo,
A. douradilho Bertaco, and A. hamatilis (Bertaco, 2014; Camelier & Zanata, 2014b; Bertaco & Vigo, 2015). Apart
from A. ojiara and A. troya, already distinguished on the “Diagnosis” of A. lorien, the latter differs from the species
listed above by having a unique vertically-elongated humeral blotch (vs. two in A. bagual, A. chico, A. douradilho,
A. pynandi, and A. stenohalinus, and one small rounded or trapezoidal humeral blotch in A. aramburui), from A.
rutilus by having 13–15 horizontal scale rows around the caudal peduncle (vs. 16) and five to six series of scales on
the upper transversal line (vs. eight), and from A. hermosus by the absence of broad dark midlateral stripe (vs.
presence) and the terminal position of the mouth (vs. mouth slightly upturned).
Astyanax rupestris and some populations of A. lorien (e.g., UFBA 8114) have globular structures externally
visible as white dots over the body, mainly distributed over head and borders of scales of males and females (Fig.
5). The presence of similar epidermal structures, identified herein as breeding tubercles, has been reported to
various Characiformes within the families Characidae, Lebiasinidae and Parodontidae, where they occur mostly in
males (Wiley & Collette, 1970; Collette, 1977). Particularly in small characids, globular whitish structures were
cited for a few species [e.g., Astyanax aeneus (Günther), A. aramburui and Bryconamericus thomasi Fowler by
Protogino et al. (2006), A. epiagos by Zanata & Camelier (2008), A. eremus by Ingenito & Duboc (2014), A.
gymnodontus (Eigenmann), A. ojiara, A. parahybae (Eigenmann) and A. troya by Terán et al. (2017),
Eretmobrycon emperador (Eigenmann & Ogle) by Meek & Hildebrand (1916: 284), two species of
Monotocheirodon Eigenmann & Pearson by Collette (1977: 238), and Myxiops aphos by Zanata & Akama (2004,

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fig. 2)]. In some of these species (e.g., A. aeneus, A. aramburui, and B. thomasi) the globular structures were cited
to occur exclusively in mature males and identified as breeding tubercles. In others, as M. aphos, the structures
where not identified as breeding tubercles since they occur in in both males and females of all sizes. A histological
examination performed by Zanata & Akama (2004) revealed that the cited structures in M. aphos represent an
accumulation of epithelial cells. The latter authors questioned about the homology of the epidermal structures in
both sexes and juveniles of Myxiops versus the apparently true breeding tubercles cited previously to the families
Lebiasinidae and Parodontidae. Similarly to Myxiops, the whitish epidermal structures of A. lorien and A. rupestris
occur in both sexes. As mentioned by Zanata & Akama (2004), the exact homology of the epidermal structures
cited for various taxa is uncertain and future studies are necessary to ascertain their identity.
TABLE 2. Morphometric data for Astyanax rupestris, new species; n= 66, range includes holotype. SD= standard
deviation.
Both Astyanax lorien and Astyanax aff. rupestris, as well as some undescribed Astyanax species, are
apparently restricted to particular headwaters of the upper portion of the Paraguaçu river basin, not occurring in the
main channel of the river. Both Astyanax lorien and A. aff. rupestris present various morphological reductions,
including lateral line incomplete or discontinuous and four premaxillary teeth in the inner row. These reductions
are also found in some undescribed species of Astyanax from the same general region (AMZ and RB, pers. obs).
Particularly, the rio Piabinha, inhabited by the population with the greatest degree of morphological reductions
among the congeners examined herein, identified as Astyanax aff. rupestris herein, is somewhat isolated from the
main course of rio Cumbuca by a steep 600 meters long stretch, with a 35 meters high gradient. Similar to most of
the headwaters within the CDD inhabited by small populations of Astyanax, the rio Piabinha is included in the
“Polígono das Secas”, with long periods of drought and a critical low water period from April to October (Azevedo
& Silva, 2000; CPRM, 2005). Thus, these small fish populations are constantly exposed to critical environmental
holotype range mean SD
Standard length (mm) 85.2 28.8–86.8 56.6 -
Percentages of standard length
Depth at dorsal-fin origin 31.0 26.7–35.6 31.8 2.0
Snout to dorsal fin origin 50.1 49.4–54.8 52.1 1.4
Snout to pectoral-fin origin 26.1 25.0–30.8 27.6 1.3
Snout to pelvic-fin origin 48.8 46.4–52.2 49.6 1.7
Snout to anal-fin origin 65.8 61.5–67.8 65.3 1.7
Caudal peduncle depth 11.2 10.4–13.1 11.7 0.6
Caudal peduncle length 16.7 13.9–19.5 16.3 1.1
Pectoral-fin length 18.4 18.0–22.6 20.4 1.0
Pelvic-fin length 13.3 12.8–15.9 14.4 0.7
Dorsal-fin base length 12.7 10.5–14.3 12.2 0.9
Dorsal-fin heigth 20.9 20.9–27.4 23.3 1.2
Anal-fin base length 22.1 19.0–25.0 22.5 1.2
Anal-fin lobe length 14.4 14.4–21.2 16.3 1.1
Eye to dorsal-fin origin 38.0 35.7–41.0 38.5 1.3
Dorsal-fin origin to caudal-fin base 52.5 49.2–55.0 51.8 1.3
Head length 25.8 25.2–28.9 27.0 0.9
Percentages of head length
Horizontal eye diameter 25.0 25.0–30.9 28.2 1.7
Snout length 23.2 21.8–26.7 24.1 1.2
Least interorbital width 34.5 29.1–35.0 31.3 1.4
Upper jaw length 44.1 38.9–44.5 41.2 1.5

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factors as they have to cope with low water levels, restriction of inhabitable areas to small rocky bottomed pools,
with high temperatures and consequential oxygen and food limitations. Furthermore, small and isolated
populations also face threats from genetic drift and inbreeding, possibly leading to an increased frequency of
polymorphisms and weakly deleterious mutations (Lynch et al., 1995; Perrier et al., 2017; Romiguier et al., 2017).
The morphological reductions observed in Astyanax lorien and Astyanax aff. rupestris may be a consequence of
small isolated populations facing environmental stress and, consequently, under consequences of genetic drift and
inbreeding. Future studies examining whether intrinsic or extrinsic mechanisms are generating the observed
reductions in populations of distinct species of Astyanax in the upper Paraguaçu river are needed.
FIGURE 7. Sampling locality of Astyanax rupestris, Brazil, Bahia, Chapada Diamantina, Mucugê, rio Cumbuca above
Tiburtino, Parque Municipal Sempre Viva.
Comparative material examined. All from Brazil, except if otherwise noted. Astyanax burgerai: MZUSP
101245 (holotype, 53.2 mm SL); UFBA 4346 (paratypes, 15, 1 c&s, 19.5–54.4 mm SL), Bahia, rio Almada basin.
Astyanax epiagos: UFBA 2792 (paratypes, 344, 3 c&s, 14.0–52.6 mm SL); UFBA 2794 (paratypes, 6, 26.9–36.6
mm SL), Bahia, rio Paraguaçu basin. Astyanax fasciatus (Cuvier): UFBA 3972 (15, 1 c&s, 83.9–93.8 mm SL),
Alagoas, rio São Francisco basin. Astyanax hamatilis: MZUSP 49232 (137, two c&s, 17. 9–42.7 mm LS), Bahia,
rio Paraguaçu basin. Astyanax intermedius: MZUSP 79409 (7, 28.7–72.7 mm SL), São Paulo, rio Paraíba do Sul
basin. Astyanax jacobinae: UFBA 2793 (paratypes, 5, 1 c&s, 22.8–38.2 mm SL), Bahia, rio Itapicuru basin.
Astyanax lacustris (Lütken): UFBA 856 (23, 41,7–70,9 mm SL), Bahia, rio São Francisco basin. Astyanax ojiara:
MZUSP 40255 (paratypes, 5, 39.6–44.5 mm SL), Argentina, Misiones, rio Uruguay basin. Astyanax parahybae
Eigenmann: MZUSP 42870 (2 c&s, 67.3–70.6 mm SL), São Paulo, rio Paraíba do Sul basin. Astyanax paranae:
MZUSP 85864 (7, 29.2–68.2 mm SL), São Paulo, rio Tietê basin; MZUSP 55000, 3 c&s , 43.4–47.6 mm SL,
Paraná, rio Iguaçu basin. Astyanax pelecus Bertaco & Lucena: UFBA 5634 (16, 34.9–62.1 mm SL), Bahia, rio
Pardo basin. Astyanax rivularis: MZUSP 50734 (6, 76.7–99.1 mm SL), Minas Gerais, rio São Francisco basin;
UFBA 246 (111, 22.3–46.7 mm SL), Bahia, rio São Francisco basin. Astyanax scabripinnis: BMNH 1917.7.14:15
(holotype, 71.4 mm SL), Rio de Janeiro. Astyanax taeniatus: MZUSP 84624 (4, 54.3–84.6 mm SL), São Paulo, rio
Guaratuba. Astyanax turmalinensis: DZUFMG 8 (paratypes, 5, 28.7–56.3 mm SL), Minas Gerais, rio
Jequitinhonha basin; MZUSP 93807 (11, 17.6–26.1 mm SL), Minas Gerais, rio Pardo basin. Astyanax vermilion
Zanata & Camelier: MZUSP 101243 (holotype, 39.4 mm SL); MZUSP 101244 (paratypes, 10, 37.4–38.7 mm SL);

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UFBA 4342 (paratypes, 12, 3 c&s 26.9–39.2 mm SL), Bahia, rio Cachoeira basin; UFBA 4813 (24, 25.7–42.8 mm
SL), Bahia, rio Almada basin. Astyanax aff. rupestris: all from Bahia, rio Piabinha, tributary of rio Cumbuca,
tributary of rio Paraguaçu: UFBA 2784 (87, 20.4–71.9 mm SL); UFBA 5521 (23, 29.9–52.1 mm SL); UFBA 5522
(41, 29.0–65.5 mm SL); UFBA 5523 (117, 19.7–62.5 mm SL). UFBA 7708 (4, 53.0–68.5 mm SL); UFBA 8081
(11, 29.5–36.5 mm SL); UFBA 8085 (36, 26.3–72.0 mm SL).
Acknowledgements
Thanks to the Projeto Sempre Viva by the support and permission to proceed various samplings within the rio
Cumbuca and rio Piabinha. We are grateful to O.T. Oyakawa (MZUSP) and P.A. Buckup (MNRJ) for the loan of
specimens used in this study. Thanks also to the two anonymous reviewers for their careful reading of the
manuscript and useful comments and suggestions. We thank Carlos Oliveira (UEM) for information on lateral-line
perforation in some species of Astyanax. Funding for field activities to AMZ during which part of the specimens
were collected was provided by CNPq (grants # 476449/2007–3, # 562335/2010–2, # 476495/2010–5). Permission
for collecting was granted by IBAMA # 13754–1. RB was also supported by CNPq/CAPES (grant from Protax #
440534/2015–1) and PC by FAPESP (grant # 2016/03966–1).
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