A new species of Astyanax (Characiformes: Characidae) from Costa Rica and Panama, with a key to the lower Central American species of the genus

Abstract

Astyanax anai, a new species of characid fish, is described from the Sixaola River basin, eastern Costa Rica–western Panama, Central America. The new species can be distinguished from all other congeners by the following combination of characters: premaxillary teeth 4–5 at the inner series and 4–6 at the outer series; maxillary teeth tricuspid, 2–4; predorsal scale series irregular and incomplete, with an unscaled space behind tip of supraoccipital process and 12–14 scales; lateral line scales 34–39; humeral region with a conspicuous black and rounded to horizontally ovate spot and two diffuse brown and vertically elongate bars (the first through the rounded to horizontally ovate spot, the second 2–4 scales behind the first); body depth 36.6–42.3% of standard length (LS); midlateral stripe formed by a series of 10–14 anteriorly‐directed dermal herringbone, or chevron‐shaped, marks, most apparent in juveniles and in preserved specimens, extending above the lateral line from the black humeral spot or just behind it (from the second vertical bar) to the caudal peduncle; scale rows from lateral line to base of first dorsal‐fin ray 8–9; scale rows from lateral line to base of pelvic fin 7–8; pre‐anal distance 53.9–61.9% of LS; total anal‐fin elements 29–33; caudal spot elongated, rhomboid or rectangular, with its anterior margin surpassing the middle of the caudal peduncle, usually reaching the anal‐fin insertion, posteriorly covering 4–7 principal caudal‐fin rays and not extending onto the ventral and dorsal margins of the caudal peduncle, covering 3–5 horizontal scale rows. In order to test the phylogenetic relationships of the new taxon in relation to the other North and Central American species of the genus, a new phylogenetic hypothesis based on a reanalysis of the morphological matrix by Schmitter‐Soto (2016) is proposed. A key to the lower Central American (southern Nicaragua to eastern Panama) species of Astyanax is also provided.

Full text

Journal of Fish Biology (2018) 92, 1866–1887
doi:10.1111/jfb.13626, available online at wileyonlinelibrary.com
A new species of Astyanax (Characiformes: Characidae)
from Costa Rica and Panama, with a key to the lower
Central American species of the genus
A. A*†‡, A. C. S†, M. L*, F. L†
C. D. M§
*Museo de Zoología and Centro de Investigación en Ciencias del Mar y Limnologia
(CIMAR), Universidad de Costa Rica, 11501– 2060, San Pedro de Montes de Oca, San José,
Costa Rica, †UNESP, Universidade Estadual Paulista ‘Júlio de Mesquita Filho’, Laboratório
de Ictiologia, Departamento de Zoologia e Botânica, Rua Cristóvão Colombo, 2265,
15054–000 São José do Rio Preto, SP, Brazil and §The Field Museum of Natural History,
1400 S, Lake Shore Drive, Chicago, Illinois 60605, U.S.A.
urn:lsid:zoobank.org:pub:345E76FF-C6CE-459F-BDD8-26C56634022A
(Received 6 December 2017, Accepted 27 March 2018)
Astyanax anai, a new species of characid sh, is described from the Sixaola River basin, eastern Costa
Rica– western Panama, Central America. The new species can be distinguished from all other con-
geners by the following combination of characters: premaxillary teeth 4– 5 at the inner series and 4–6
at the outer series; maxillary teeth tricuspid, 2– 4; predorsal scale series irregular and incomplete, with
an unscaled space behind tip of supraoccipital process and 12–14 scales; lateral line scales 34 –39;
humeral region with a conspicuous black and rounded to horizontally ovate spot and two diffuse brown
and vertically elongate bars (the rst through the rounded to horizontally ovate spot, the second 2–4
scales behind the rst); body depth 36·6–42·3% of standard length (LS); midlateral stripe formed by
a series of 10– 14 anteriorly-directed dermal herringbone, or chevron-shaped, marks, most apparent
in juveniles and in preserved specimens, extending above the lateral line from the black humeral spot
or just behind it (from the second vertical bar) to the caudal peduncle; scale rows from lateral line to
base of rst dorsal-n ray 8– 9; scale rows from lateral line to base of pelvic n 7–8; pre-anal distance
53·9–61·9% of LS; total anal-n elements 29– 33; caudal spot elongated, rhomboid or rectangular, with
its anterior margin surpassing the middle of the caudal peduncle, usually reaching the anal-n inser-
tion, posteriorly covering 4– 7 principal caudal-n rays and not extending onto the ventral and dorsal
margins of the caudal peduncle, covering 3–5 horizontal scale rows. In order to test the phylogenetic
relationships of the new taxon in relation to the other North and Central American species of the genus,
a new phylogenetic hypothesis based on a reanalysis of the morphological matrix by Schmitter-Soto
(2016) is proposed. A key to the lower Central American (southern Nicaragua to eastern Panama)
species of Astyanax is also provided.
© 2018 The Fisheries Society of the British Isles
Key words: Astyanax orthodus; lower Central America; Neotropical sh; taxonomy; Teleostei.
INTRODUCTION
Astyanax Baird & Girard 1854, is one of the most species-rich and widespread genera
of the Characidae (Pisces: Characiformes). Currently, it is composed of c. 150 valid
‡Author to whom correspondence should be addressed. Tel.: +506 25115446; email: arturo.angs@gmail.com
1866
© 2018 The Fisheries Society of the British Isles

A NEW ASTYANAX FROM CENTRAL AMERICA 1867
species distributed from the southern U.S.A. to central Argentina (Lucena & Soares,
2016). As could be expected from a large and widespread sh assemblage, and as typ-
ical of other genera in the family, the systematics of Astyanax is very complex and
still needs to be more thoroughly examined (Marinho & Lima, 2009). According to
recent phylogenies (Weitzman & Malabarba, 1998; Calcagnotto et al., 2005; Javonillo
et al., 2010; Mirande, 2010; Oliveira et al., 2011) the genus does not represent a mono-
phyletic group. On the contrary, Astyanax is currently dened by a combination of
morphological characters, proposed nearly a century ago (Eigenmann, 1921, 1927),
which are broadly distributed in many other genera of the Characidae (e.g. Hemigram-
mus Gill 1858; Hyphessobrycon Durbin 1908; Moenkhausia Eigenmann 1903).
In lower Central America (southern Nicaragua to eastern Panama), Loftin (1965),
Bussing (1987, 1998, 2008), Smith & Bermingham (2005) and Angulo et al. (2013)
listed a total of seven (valid) species of Astyanax:Astyanax aeneus (Günther 1860)
(Pacic and Atlantic Ocean, from southern Mexico to Panama), Astyanax cocibolca
Bussing 2008 (Atlantic Ocean, southern Nicaragua and northern Costa Rica), Astyanax
kompi Hildebrand, 1938 (Pacic Ocean, western Panama), Astyanax nasutus Meek
1907 (Atlantic Ocean, southern Nicaragua), Astyanax orthodus Eigenmann, 1907
(Atlantic Ocean, from eastern Costa Rica to western Colombia), Astyanax panamen-
sis (Günther 1864) (Pacic and Atlantic Ocean, central and western Panama) and
Astyanax ruberrimus Eigenmann 1913 (=A. panamensis; Pacic and Atlantic Ocean,
from central Panama to eastern Colombia). In addition, Loftin (1965) and Bussing
(1987, 1998) recognized several morphological variants and noted some taxonomic
and nomenclatural problems highlighting the need for further studies at the regional
scale.
Schmitter-Soto (2016, 2017) reviewed the species of the genus occurring from
the U.S.A. to Panama and listed a total of six species occurring south of Nicaragua,
including three (A. cocibolca,A. nasutus and A. panamensis) of those listed by
previous authors (see above) but omitting two of them [A. kompi and A. orthodus
(sensu Bussing, 1987, 1998)]. In addition to these species, Schmitter-Soto (2016,
2017), based on his own research and supported by molecular evidence published by
Ornelas-Garcia et al. (2008), proposed revalidating Astyanax nicaraguensis Eigen-
mann & Ogle, 1907 (Atlantic Ocean, from central Nicaragua to western Costa Rica)
and Astyanax orstedii (Lütken 1875) (Pacic and Atlantic Ocean, from southern
Nicaragua to western Panama), considered by previous authors as junior synonyms
of A. aeneus; synonymizing Bramocharax Gill 1877 with Astyanax, adding Astyanax
bransfordii (Gill 1877) (from southern Nicaragua and north-eastern Costa Rica) to the
list of valid species of the genus; synonymizing A. ruberrimus with A. panamensis.
Based on these proposed changes, Astyanax would be represented in lower Central
America by a total of eight valid species; i.e. A. bransfordii,A. cocibolca,A. kompi,
A. nasutus,A. nicaraguensis,A. orstedii, A. orthodus and A. panamensis.
In a previous review of the Costa Rican species of Astyanax, Bussing (1987, 1998)
classied some specimens collected in the Sixaola River basin (Costa Rica– Panama)
and deposited at Universidad de Costa Rica, Escuela de Biología, Museo de Zoología,
Departemento de Biología, San Jose, Costa Rica (UCR) and the Natural History
Museum of Los Angeles County, Los Angeles, U.S.A. (LACM) as A. orthodus.
These specimens presented a combination of characters unique among other Central
American congeners [i.e. body deep, about 40·0% of standard length (LS); presence of
a conspicuous black and horizontally ovate humeral spot and two diffuse brown and
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1868 A. ANGULO ET AL.
vertically elongate spots (or bars) in the humeral area (the rst through the humeral
spot, the second 2–4 scales behind the rst); as well as a series of dermal herringbone
or chevron-shaped marks on the sides, most apparent in juveniles and in preserved
specimens], which match well with the diagnosis of A. orthodus, originally described
from western Colombia (Eigenmann & Ogle, 1907). With this new record for Central
American waters, Bussing (1987, 1998)) extended the known distributional range
of A. orthodus and proposed a continuous distribution for the species through lower
Central America. However, earlier revisions (Hildebrand, 1938; Loftin, 1965), as well
as recent papers (Smith & Bermingham, 2005) and expeditions assessing part or the
totality of the freshwater sh fauna of Panama, fail to list A. orthodus eastward to
the Sixaola River basin, contradicting Bussing’s (1987, 1998) conclusion. Moreover,
other authors (Lima et al., 2003; Ruiz-C et al., 2011; Maldonado-Ocampo et al., 2012;
Ruiz-C et al., 2018), in accordance with the foregoing, have restricted A. orthodus to
western Colombia (i.e. Atrato River basin), generating some doubt about the identity
of the specimens listed as A. orthodus by Bussing (1987, 1998) and other authors, for
Costa Rica and Panama.
A detailed examination of museum specimens catalogued as A. orthodus, both from
Costa Rica–Panama and Colombia, including type material from Colombia as well
as other sympatric and related species, revealed that specimens from Costa Rica and
Panama represent a previously unknown species (different from A. orthodus sensu
lato as well as from the other lower Central American species of the genus) which
is described here. Additionally, in order to test the phylogenetic relationships of the
new taxon in relation to the other North and Central American species of the genus, a
new phylogenetic hypothesis, based on a re-analysis of the morphological matrix by
Schmitter-Soto (2016), is proposed and presented. Finally, a key to the identication of
the lower Central American species of Astyanax, adapted from Schmitter-Soto (2017),
also is provided.
MATERIAL AND METHODS
Counts (except those for supraneurals, vertebrae and caudal-n procurrent rays) and mea-
surements (Table I) were taken on the left side of specimens and followed Fink & Weitzman
(1974) and Melo & Buckup (2006). Counts of supraneurals, vertebrae and caudal-n procur-
rent rays were taken from seven paratypes that were cleared and stained (c&s) following Tay-
lor & Dyke (1985). Osteological terminology and descriptions followed Weitzman (1962) and
Schmitter-Soto (2016, 2017). In the description, numbers between parentheses after each count
indicate the percentage of each count in examined material (58 specimens in total). An aster-
isk indicates counts for the holotype. Colour in life description was based on photos of freshly
collected specimens and direct observations of live specimens in the eld.
Morphological data cited in the diagnosis from other species of Astyanax were based on
original descriptions, additional literary resources (Eigenmann & Ogle, 1907; Eigenmann,
1921; Hildebrand, 1938; Garutti & Britski, 1997; Bussing, 1998, 2008; Garutti, 1998; Garutti
& Langeani, 2009; Dagosta, 2011; Maldonado-Ocampo et al., 2012; Torres-Mejia et al., 2012;
Marinho & Birindelli, 2013; Marinho & Ohara, 2013; Lucena & Soares, 2016; Schmitter-Soto,
2016 and Ruiz-C et al., 2018) and additional analysed specimens. Institutional abbreviations
followed Sabaj (2016): CAS, Chinese Academy of Sciences, Institute of Zoology, Beijing,
PDR China; DZSJRP, Departmento de Zoologia e Botânica, Universidade Estadual Paulista
Júlio de Mesquita Filho (UNESP), Campus de São José do Rio Preto, São Paulo, Brazil;
FMNH, Field Museum of Natural History, Geology Department (slide collection), Chicago, IL,
U.S.A.; MZUSP, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil; USNM,
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1869
T I. Range mean, mode and .. (including the holotype) morphometric and meristic char-
acteristics of Astyanax anai (n=58) expressed as percentages of standard length
Characteristic Holotype Range Mean
(Mode) ..
Morphometrics
Head length 25·56 25·56–33·01 29·17 1·58
Predorsal distance 48·30 42·88–52·26 49·19 1·71
Prepelvic distance 38·95 38·23–46·66 41·83 1·76
Prepectoral distance 23·55 19·06–28·85 24·57 2·12
Pre-anal distance 55·02 53·88–61·92 57·64 1·86
Depth at dorsal-n origin 39·22 36·57–42·31 39·71 1·23
Dorsal-n base length 15·21 12·62–15·32 14·25 0·63
Dorsal-n height 25·62 24·24 –30·41 26·64 1·33
Posterior terminus of dorsal n
to adipose n length
22·43 22·03–24·30 23·17 0·64
Posterior terminus of dorsal n
to hypural joint length
37·01 36·60–42·02 38·98 1·42
Anal-n base length 35·21 30·58– 39·91 34·22 2·21
Pectoral-n length 21·28 19·16–24·35 21·65 1·09
Pelvic-n length 15·75 14·43–23·02 16·52 1·39
Caudal-peduncle depth 11·93 9·75–12·90 11·62 0·67
Caudal-peduncle length 10·40 9·00–14·19 11·70 1·08
Snout length 6·62 6·46–9·91 7·65 0·86
Upper jaw length 9·89 7·68–17·51 10·64 1·61
Maxillary length 13·36 8·42–16·04 11·22 1·57
Eye diameter 9·98 7·21 –13·29 9·37 1·33
Interorbital width 8·21 9·07–11·61 10·03 0·50
Post-orbital head length 10·71 7·98– 15·34 12·37 1·52
Meristics
Dorsal-n elements 12 12 12·00 (12)0·00
Anal-n elements 31 29–33 30·79 (31)0·85
Pectoral-n elements 13 12–13 12·81 (13)0·40
Pelvic-n elements 8 8 8.00 (8)0·00
Caudal-n elements 19 18–20 19·14 (19)0·44
Predorsal scales 12 12–14 12·81 (13)0·51
Lateral-line scales 38 34–39 36·62 (36)1·24
Scale between dorsal-n origin
and lateral line
98–98
·81 (9)0·40
Scale rows between lateral line
and pelvic-n origin
77–87·84 (8)0·37
Scale rows between lateral line
and anal-n origin
98–98·84 (9)0·37
Scales around caudal peduncle 15 15 –19 16·76 (17)1·17
Total gill-rakers on rst gill arch 19 22 21·10 (22)0·83
National Museum of Natural History, Smithsonian Institution, Department of Vertebrate
Zoology, Washington D.C., U.S.A.
Phylogenetic placement was tested by including the new taxon in the data matrix of
Schmitter-Soto (2016). The phylogenetic analysis of the new dataset (the character codication
for A. orthodus and A. anai is presented in Table II) was performed using the software TNT
1·1 (Goloboff et al., 2008). All characters (52 in total) were considered as unordered, binary or
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1870 A. ANGULO ET AL.
T II. Character states of Astyanax anai and A. orthodus for the examined matrix of
Schmitter-Soto (2016) using TNT software
Astyanax anai
0000010200 ?002?1121? 1120001101 1100022111 01?111?110 00
Astyanax orthodus
0000010200 ?002?11211 1120001101 1100022101 01?1010110 00
Polymorphic condition is represented as ‘?’.
multistate, non-additive and received equal weights. The most parsimonious trees were obtained
through a heuristic search using new technologies (Goloboff et al., 2008) under the following
parameters: ratchet =200 iterations; drift =50 iterations, random sequence additions =1000
replicates; maximum retention =10 000 trees. From the equally parsimonious trees a strict
consensus tree was constructed using the respective function in TNT. Bremer support was
calculated in TNT using suboptimal trees with up to 15 steps more than the fundamental trees.
RESULTS
ASTYANAX ANAI SP. NOV. [Figs. 1 and 3 and Table I].
urn:lsid:zoobank.org:act:38F43231-3856-41B0-95CE-25F809BA205C
Astyanax orthodus (non Eigenmann 1907) Bussing, 1987: 71 [brief description with
notes on their ecology and distribution]; Bussing, 1998: 86 [brief description with notes
on their ecology and geographical distribution]; Smith & Bermingham, 2005: 1839
[listed for Panama; geographical distribution and biogeographic afnity]; McLarney
et al., 2008: 8 [habitat preference and some other ecological aspects]; Ornelas-Garcia
et al., 2008: 5 [phylogenetic relationships]; Dagosta, 2011: 36 [in part; listed for Costa
Rica and Panama; geographical and vertical distribution; morphological characteriza-
tion]; Angulo et al., 2013: 991 [listed for Costa Rica; geographical and vertical distri-
bution].
HOLOTYPE
UCR 1298-06·1, 81·9mmLS, stream 1·6 km south-east of Shiroles near Suretka, on
Bratsi–Shiroles road, province of Limón, Costa Rica, 9∘34′50′′ N; 82∘56′44′′ W;
60 m, coll. W.A. Bussing, 6 October 1979 (Fig. 1).
PARATYPES
DZSJRP 21087 (ex UCR 1825-04), 4 (1 c&s), 33·30–72·96 mm LS, Mata de Limón
and San Miguel, Crique Azul, between Finca Lomas (ANAI) and the Finca of Marcos
Corea, province of Limón, Costa Rica, 9∘33′29′′ N; 82∘38′9′′ W; 40 m, collectors
W. McLarney and S. Ervin; 27 February 1986; DZSJRP 21088 (ex UCR 1141-03), 2
c&s, 47·70–52·20 mm LS, stream 0·6 km south-east of Shiroles, on Bratsi–Shiroles
road, province of Limón, Costa Rica, 9∘34′59′′ N; 82∘57′9′′ W; 60 m, collectors
W.A. Bussing, K. Gocke and P. Siegfried, 13 November 1977; DZSJRP 21089 (ex
UCR 1298-06·2), 2 c&s, 53·10–56·80 mm LS, collected with the holotype; FMNH
131444 (ex UCR 1141-03), 2, 79·16–87·58 mm LS, stream 0·6 km south-east of Shi-
roles, on Bratsi–Shiroles road, province of Limón, Costa Rica, 9∘34′59′′ N; 82∘
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1871
(a)
(b)
(c)
(d)
F. 1. Astyanax anai sp. nov., Sixaola River, Costa Rica; (a) UCR 1298-06·1, holotype, 81·9 mm standard length
(LS); (b) FMNH 131445, paratype, 92·3mmLS; (c) and (d) fresh-caught non-preserved specimens, 89·5mm
LSand 70·9mmLS, respectively.
57′9′′ W; 60m, collectors W.A. Bussing, K. Gocke and P. Siegfried, 13 Novem-
ber 1977; FMNH 131445 (ex UCR 1298-06·2),1,92·28 mm LS, collected with the
holotype; UCR 1140-07, 21 (2 c&s), 33·41–76·26 mm LS, stream 200 m south-east of
Shiroles, on Bratsi–Shiroles road, province of Limón, Costa Rica, 9∘35′5′′ N; 82∘
57′9′′ W; 60m, collectors W.A. Bussing, K. Gocke and P. Siegfried, 13 November
1977; UCR 1142-02, 3, 68·84–100·43 mm LS, stream 1·6 km south-east of Shiroles,
on Bratsi–Shiroles road (at Suretka), province of Limón, Costa Rica, 9∘34′44′′ N;
82∘56′39′′ W; 60m, collectors W.A. Bussing, K. Gocke and P. Siegfried, 13 Novem-
ber 1977; UCR 1299-13, 6, 84·41–112·66 mm LS, Río Cocolis, 3·5 km south-east of
Shiroles, on Bratsi–Shiroles road, province of Limón, Costa Rica, 9∘34′4′′ N; 82∘56′
4′′ W; 40m, coll. W.A. Bussing, 6 October 1979; UCR 1531-06, 5, 37·64–50·09 mm
LS, Quebrada 0·5 km south-east of Shiroles, province of Limón, Costa Rica, 9∘35′5′′
N; 82∘57′9′′ W; collectors W.A. Bussing, F. Muñoz and J. Pérez, 7 October 1983;
UCR 1532-02, 3, 46·06–57·59 mm LS, stream near Laguna Don, on road to Sixaola,
province of Limón, Costa Rica, 9∘35′29′′ N; 82∘44′39′′ W; collectors W.A. Bussing,
F. Muñoz and J. Pérez, 8 October 1992; UCR 1881-07, 8, 33·41–67·09 mm LS,San
Miguel, Crique Azul on Finca Los Laureles of Anai, province of Limón, Costa Rica,
9∘33′49′′ N; 82∘40′39′′ W; 15m, coll. W. McLarney, 31 March 1981.
NON-TYPE SPECIMENS
CAS 68769, 4, 57·6–87·1mmLS, stream into tributary of San-San River below rail-
way bridge, Panama, 9∘27′15′′ N; 82∘37′17′′ W; collectors Behre & Chambers, 6
February 1923.
DIAGNOSIS
Astyanax anai differs from all other congeners, except those belonging to the
A. bimaculatus species complex (Garutti & Britski, 1997; Garutti, 1998; Garutti &
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1872 A. ANGULO ET AL.
Langeani, 2009; Lucena & Soares, 2016) by the presence of a black and rounded
to horizontally ovate humeral spot (covering 3–4 scales horizontally and 2–3 scale
series vertically) and two very diffuse brown and vertically elongate spots (or bars)
in the humeral area (the rst through the black humeral spot, the second 2–4 scales
behind the rst), being more evident in preserved specimens (v. not as described
above in other congeners). Astyanax anai differs from all species of the A. bimacu-
latus species complex, except from Astyanax boliviensis Ruiz-C., Román-Valencia,
Taphorn, Buckup & Ortega 2018 (from the upper Madeira River drainage, Amazon
River basin, Bolivia–Peru), Astyanax bopiensis Ruiz-C., Román-Valencia, Taphorn,
Buckup & Ortega 2018 (=Astyanax multidens Pearson 1924) (from the upper Madeira
River drainage, Bolivia–Peru), Astyanax embera Ruiz-C., Román-Valencia, Taphorn,
Buckup & Ortega 2018 (from the Patía River basin, Colombia), Astyanax moorii
(Boulenger 1892) from the upper Paraguay River basin, southern Brazil), A. orthodus
(now restricted to the Atrato River Basin, Colombia), Astyanax superbus Myers
1942 (from the Orinoco River basin, Colombia– Venezuela) and Astyanax yariguies
(Torres-Mejia et al., 2012) (from the Magdalena River basin, Colombia), i.e. the
A. orthodus species-group sensu Ruiz-C et al. (2018), by the presence of a midlateral
stripe formed by a series of usually 10–14 anteriorly directed dermal herringbone or
chevron-shaped marks most apparent in juveniles and in preserved specimens, which
extends above the lateral line from the humeral spot or just behind it (from the second
vertical bar) to the caudal peduncle (v. absent or not as described above in the other
species of the group), as well as by the presence of a black caudal spot of variable size
and shape that continues over the median caudal-n rays (v. absent or not as described
above in the other species of the group).
Furthermore, A. anai differs numerous other Astyanax spp., as follows. Astyanax
boliviensis by the following combination of characters: pre-anal distance 53·9–61·9%
of LS, mean 57·6% (v.60·8–65·9% of LS, mean 63·6%); lateral line scales 34– 39, usu-
ally 38 (v. 39–42, usually 40); chevrons without distal extensions (v. present; Ruiz-C
et al., 2018); caudal spot restricted to the caudal n and adjacent caudal peduncle, not
extending anteriorly as a continuous dark line or stripe on side of body (v. not restricted
to the caudal n and adjacent caudal peduncle, extending anteriorly as a continuous
dark line or stripe on side of body). Astyanax bopiensis by the following combination
of characters: maxillary length 34·3–45·7% of LH, mean 42·2% (v.46·7–58·9% of LH,
mean 50·6%); maxillary teeth 2–4 (v. 7–13); post-orbital head length 39·4–48·0% of
LH, mean 44·7% (v.20·8–30·4% of LH, mean 25·2%); lateral line scales 34–39, usu-
ally 38 (v. 39–42, usually 40); absence of a reticulated pattern on the sides of the body
formed by dark pigment on the mediodistal margins of scales (v. present); caudal spot
restricted to the caudal n and adjacent caudal peduncle, not extending anteriorly as
a continuous dark line or stripe on side of body (v. not restricted to the caudal n and
adjacent caudal peduncle, extending anteriorly as a continuous dark line or stripe on
side of body). Astyanax. embera by the following combination of characters: maxillary
length 34·3–45·7% of LH, mean 42·2% (v.27·2–33·3% of LH, mean 30·5%); pre-anal
distance 53·9–61·9% of LS, mean 57·6% (v.63·1–67·2% of LS, mean 64·6%); caudal
spot not extending onto the ventral and dorsal margins of the caudal peduncle, cov-
ering 3–5 horizontal scale rows (v. extending onto the ventral and dorsal margins of
the caudal peduncle, covering 5– 7 horizontal scale rows). Astyanax moorii by the fol-
lowing combination of characters: nasal with lamellae (v. without lamellae; Dagosta,
2011); interorbital width 30·0–41·9% of LH, mean 35·25% (v.27·4–31·4% of LH, mean
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A NEW ASTYANAX FROM CENTRAL AMERICA 1873
F. 2. Astyanax orthodus, Truando, Colombia, USNM 55655, holotype, 90·2 mm standard length.
29·2%); pre-anal distance 53·9–61·9% of LS, mean 57·6% (v.63·3–64·8% of LS, mean
63·8%); rhinosphenoid present and ossied (v. absent or cartilaginous; Dagosta, 2011);
posterior portion of the coracoid without a conspicuous projection (v. with a conspic-
uous projection; Dagosta, 2011); caudal spot conspicuous and restricted to the caudal
n and adjacent caudal peduncle, not extending anteriorly as a continuous dark line
or stripe on side of body (v. usually inconspicuous and not restricted to the caudal
n and adjacent caudal peduncle, extending anteriorly as a continuous dark line or
stripe on side of body). Astyanax orthodus by the following combination of charac-
ters: maxillary length 34·3–45·7% of LH, mean 42·2% (v.27·8–34·4% of LH, mean
30·6%); scale rows between lateral line and pelvic n origin 8, rarely 7 (v. 6, rarely
7); pre-anal distance 53·9–61·9% of LS, mean 57·6% (v.62·7–65·2% of LS, mean
63·9%); post-anal element longer (v. shorter; see Fig. 2 in Schmitter-Soto, 2016); cau-
dal spot elongated, rhomboid or rectangular (v. not horizontally elongated and usually
teardrop shaped, square or pentagonal; Fig. 2), with its anterior margin surpassing, as
a stripe, the middle of the caudal peduncle, usually reaching the anal-n insertion (v.
not surpassing the middle of the caudal peduncle; Fig. 2) and posteriorly covering 4–7
principal caudal-n rays (v. covering 0– 3 principal caudal-n rays; Fig. 2). Astyanax
superbus by the following combination of characters: maxillary length 34·3–45·7%
of LH, mean 42·2% (v.30·1–35·5% of LH, mean 32·9%); total gill-rakers on rst gill
arch 19–22, usually 22 (v. 22–25, usually 23–24); predorsal scales 12– 14, usually
13 (v. 13–16, usually 15); bony hooks present on both pelvic and anal-n rays on
mature males (v. absent); absence of dark brown zig-zag longitudinal bands (or wavy
streaks) on the sides of the body formed by the concentration of chromatophores in
the upper and lower portions of each scale (v. present). Astyanax yariguies by the fol-
lowing combination of characters: snout length 21·8–31·1% of LH, mean 27·10% (v.
14·9–25·3% of LH, mean 20·4%); maxillary length 34·3–45·7% of LH, mean 42·2%
(v.26·9–34·7% of LH, mean 30·8%); maxillary teeth 2–4 (v. 6); predorsal scale series
irregular and incomplete, with an unscaled space behind the tip of the supraoccipi-
tal process (v. regular and complete, without an unscaled space behind the tip of the
supraoccipital process).
Also, in addition to its coloration pattern A. anai can be easily distinguished from
all other lower Central American congeners by the following combination of external
characters. Premaxillary teeth 3–5 in the outer series, 4–6 in the inner series (v.8
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1874 A. ANGULO ET AL.
at each series, inner and outer, in A. kompi); premaxillary teeth tricuspid in the outer
series (v. conical in A. cocibolca); maxillary teeth 2–4 (v. 9–13 in A. bransfordii);
predorsal scale series irregular and incomplete, with an unscaled space behind tip of
the supraoccipital process (v. regular and complete, without an unscaled space behind
tip of supraoccipital process in all other species; Eigenmann, 1921; Schmitter-Soto,
2016); predorsal scales 12–14, usually 13 (v. 8–14, usually 10–12, in all other species;
Schmitter-Soto, 2016); body depth 36·6–42·3% of LS(v.27·0–37·0% in A. bransfordii,
A. cocibolca, A. kompi and A. nasutus); scale rows from lateral line to base of rst
dorsal-n ray 8–9 (v.5–6inA. cocibolca and A. kompi); scale rows from lateral line to
base of pelvic n 7–8 (v.5–6inA. bransfordii,A. cocibolca, A. kompi and A. nasutus);
total pectoral-n elements 12–13 (v.10inA. nicaraguensis;11inA. nasutus); total
anal-n elements 29–33, usually 31 (v. 28 or less in A. bransfordii,A. cocibolca,A.
nasutus and A. panamensis; 22–32, usually 25–27 in A. nicaraguensis and A. orstedii;
27–30, usually 28–29 in A. kompi).
DESCRIPTION
Morphometric and meristic data for examined specimens in Table I. Body deep, mod-
erately elongate and robust; greatest body depth immediately anterior to dorsal-n
origin; largest specimen examined, 100·4mmLS. Dorsal body prole convex from
upper lip to vertical through middle of eye; concave to nearly straight from the latter
point to the tip of the supraoccipital; convex from the supraoccipital to insertion of
the last dorsal-n ray; slightly convex from the latter point to the adipose-n insertion.
Ventral prole of head and abdomen convex, almost straight along anal-n base. Dorsal
and ventral proles of caudal peduncle slightly concave.
Mouth terminal, not upturned and rounded from the margin of upper lip to vertical
through the anterior nostrils. Snout length 21·8–31·1% of LH. Dentigerous arm of pre-
maxilla relatively long (see g. 15a in Schmitter-Soto, 2016), with two rows of teeth:
the outer row with three (2%), four* (90%) or ve (9%) tricuspid teeth with central
cusp longest; the inner row with four (3%), ve* (95%) or six (2%) pentacuspid teeth,
decreasing in length posteriorly with last tooth distinctly smaller and in all cases with
central cusp longest; toothless portion of the dentigerous arm larger than toothed por-
tion (Fig. 2). Maxilla extending posteriorly to vertical through anterior half of the orbit
and posterior to a vertical through the lateral ethmoid, slightly curved and aligned at
an angle of c.45
∘in relation to the longitudinal body axis; its length 34·3–45·7% of
LH; maxillary teeth two (3%), three (33%) or four* (64%), tricuspid, with central cusp
longest and aligned in a single row (Fig. 3). Dentary with four (43%) or ve* (57%)
large, pentacuspid teeth, decreasing in length with last teeth distinctly smaller and with
central cusp longest, followed by a series of six (7%), seven* (22%), eight (35%), nine
(19%) or 10 (17%) small tricuspid teeth (Fig. 2). Contour of the upper pharyngeal tooth
plates, oval, fusiform [see Fig. 1(a) in Schmitter-Soto, 2016]; lower pharyngeal tooth
plates single [see Fig. 2(a) in Schmitter-Soto, 2016], with the caudal side concave [see
Fig. 2(a) in Schmitter-Soto, 2016]. Eye diameter 25·3–36·6% of LH; interorbital width
30·0–41·9% of LH; post-orbital head length 39·4–48·0% of LH.
Anterior fontanel of neurocranium medium sized, with anterior sides straight and
rostral tip blunt (see g. 6b in Schmitter-Soto, 2016). Nasal with lamellae (see g.
83a in Dagosta, 2011). Rhinosphenoid present and ossied. Orbitosphenoid with
a prominent posteroventral process (see g. 22b in Dagosta, 2011). Caudal side of
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1875
F. 3. Jaws and dentition (lateral view, left side) of Astyanax anai sp. nov., DZSJRP 21089, paratype, 53·1mm
standard length. Scale bar 2 mm.
supraoccipital convex in lateral view (see g. 7c in Schmitter-Soto, 2016); caudal
process of supraoccipital short and usually broad-based in dorsal view (short-based
in some specimens; see g. 8b in Schmitter-Soto, 2016). Infraorbital II triangular
and convex, with the base short (see g. 10b in Schmitter-Soto, 2016); posteroventral
side of infraorbital III semicircular (see g. 11b in Schmitter-Soto, 2016); contact
between infraorbital II and III wide (Schmitter-Soto, 2016); infraorbital IV usually
square, approximately rectangular in some specimens, with a rostroventral projection
(see g. 12b in Schmitter-Soto, 2016). Metapterygoid with two dorsorostral projec-
tions (see g. 13c in Schmitter-Soto, 2016); rostral arm somewhat longer than ventral
one (see g. 13a in Schmitter-Soto, 2016). Dorsal process of quadrate distally expanded
(see g. 14a in Schmitter-Soto, 2016). Sides of the dorsal half of the opercle approx-
imately parallel at its middle (see g. 16a in Schmitter-Soto, 2016). Interopercle
relatively short (Schmitter-Soto, 2016), with the posterior edge convex and without
any angle (see g. 17b in Schmitter-Soto, 2016). Dorsorostral projection of the
subopercle present (Schmitter-Soto, 2016). Ventral rim of the preopercle straight,
at least anteriorly; anterodorsal edge of the preopercle straight to concave (see g.
18a in Schmitter-Soto, 2016); preopercular canal with two posterior branches. Edge
of the epuric plate on the last neural spine convex, not indented (see g. 19b in
Schmitter-Soto, 2016). Posterior portion of the coracoid without a conspicuous pro-
jection (see g. 88a in Dagosta, 2011). Cleithrum sutured to coracoid with 2–3 inter-
digitations (see g. 24b in Schmitter-Soto, 2016). Distance between postcleithrum one
and two shorter than height of postcleithrum one; caudal process of postcleithrum two
concave-sided dorsally, almost straight ventrally (see g. 25b in Schmitter-Soto, 2016).
Pectoral-n elements i, 11 (19%) or i, 12* (81%); tip of longest ray extending
beyond the pelvic-n origin. Pelvic-n elements i, 7 (100%); its origin anterior to
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1876 A. ANGULO ET AL.
vertical through the dorsal-n origin; longest pelvic ray usually reaching anal-n ori-
gin; proximal edge of pelvic bone curved to irregular (see g. 26a in Schmitter-Soto,
2016). Dorsal-n elements iii, 9 (100%), rst simple ray small, easily visible only in
c&s specimens, detectable only with dissecting needle in non-c&s specimens; rst
unbranched ray approximately 50% of length of second unbranched ray; dorsal-n
origin located just anterior to middle of the body; rostral edge of rst dorsal-n
pterygiophore usually with a spine (see g. 23b in Schmitter-Soto, 2016). Adipose-n
origin located dorsal to insertion of the last seven or eight anal-n rays. Anal-n
elements iv, 25 (5%), iv, 26 (22%), iii, 27 (5%), iv, 27* (55%), iv, 28 (7%), iv, 29 (3%),
iii, 30 (2%), rst simple ray small, easily visible only in c&s specimens, detectable
only with dissecting needle in non-c&s specimens; its origin posterior to vertical
through last two or three dorsal-n rays; rst anal-n pterygiophore anterodorsally
directed (see g. 87 in Dagosta, 2011); rst branched anal-n ray noticeably wider
than others (see g. 21b in Schmitter-Soto, 2016); post-anal element relatively long
(see g. 22b in Schmitter-Soto, 2016). Caudal n forked and unscaled, with both lobes
similar in size or lower one slightly larger, with i, 16, i (3%), i, 17, i* (79%) or i, 18,
i (17%) elements. Dorsal and ventral procurrent rays 10 or 11.
Scales cycloid, moderately large. Lateral line regular and complete, with 35 (12%),
36 (29%), 37 (22%), 38* (35%) or 39 (2%) scales. Predorsal scale series irregular and
incomplete, with an unscaled space behind tip of supraoccipital process (see g. 20a
in Schmitter-Soto, 2016) and with 12* (24%), 13 (71%) or 14 (5%) scales. Scale rows
between dorsal-n origin and lateral line 8 (19%) or 9* (81%); scale rows between lat-
eral line and pelvic-n origin 7 (16%) or 8* (85%); scale rows between lateral line and
anal-n origin 8 (16%) or 9* (85%). Axillary scale on pelvic-n insertion posteriorly
extended and covering 1·5 (29%) or 2* (71%) scales. Scale sheath along the anal-n
base simple and long (see Schmitter-Soto, 2016), composed by 12 (29%), 13* (31%),
14 (24%) or 15 (16%) scales, arranged in a single series and covering the base of the
corresponding 11–16 anteriormost n rays. Scales around caudal peduncle 15* (21%),
16 (16%), 17 (36%), 18 (22%), or 19 (5%).
Total gill-rakers on rst gill arch 19 (2%), 20 (24%), 21 (36%) or 22* (38%);
8 (47%), 9* (45%) or 10 (8%) on the epibranchial, one (100%) at the symphysis,
between the epibranchial and ceratobranchial and 10 (2%), 11 (48%) or 12* (50%)
on the ceratobranchial. Insertion of uncinate process of epibranchial III widely
open, hyperbolic [see Fig. 3(a) in Schmitter-Soto, 2016]; main body (distal segment)
straight (Schmitter-Soto, 2016). Anteroventral edge of the urohyal convex and uniform
[see Fig. 4(a) in Schmitter-Soto, 2016]; ventral apex somewhat closer to the caudal
end [see Fig. 4(b) in Schmitter-Soto, 2016]. Rostral vertices of ceratohyal rounded
[see Fig. 5(d) in Schmitter-Soto, 2016]; ceratohyal foramen oval [see Fig. 5(a) in
Schmitter-Soto, 2016]. Total vertebrae, including those of the Weberian apparatus, 36
or 37; precaudal vertebrae 18 and caudal vertebrae 18 or 19 (the terminal centrum was
counted as a single element). Supraneurals 5.
SEXUAL DIMORPHISM
Mature males have small retrorse bony hooks [similar to those described for
A. lacustris by Vieira et al., 2016] on both pelvic and anal-n rays, one pair per ray
segment, dispersed in the distal portion of the longest unbranched ray and in the
posterior branches of each subsequent branched ray. Juveniles and mature females do
not have n hooks.
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1877
85·5°
10·5°
(a)
(b)
9·0°
9·6°
Telire
Negro
San San
Yorkin
Sixaola
9·4°
9·6°
9·4°
10.5°
9.0°
50 100 150 200 km0
50
510 15 20 km0
5
84·0° 82·5°
82·9° 82·8°82·6°
82·9° 82·8°82·6°
81·0°
85·5° 84·0° 82·5° 81·0°
N
F. 4. Map of (a) south-eastern Costa Rica and western Panama, Central America and (b) detail of the Sixaola
River basin, showing collection localities of Astyanax anai sp. nov. ( ). Some symbols represent more than
one sample or locality. , The species’ type locality. Data from the UCR sh collection and Asociación
ANAI (collection localities, unpubl. data).
COLOUR IN ALCOHOL
General body colour pale, yellowish-brown [Fig. 1(a), (b)]. Dorsal and dorso-
lateral portions of head and body brown. Infraorbital and opercular areas usually
with scattered dark brown chromatophores. Scales on lateral portion of body also
with dark brown chromatophores. A dark spot or band of variable size and shape is
present on the mid-posterior portion of the eye, which, in most specimens, contin-
ues posteriorly covering the skin above the dorsal portion of the infraorbital series
(i.e. infraorbitals III and IV) and part of the opercle. Humeral spot conspicuous, black
and rounded to horizontally ovate, covering 2–4 scales in the horizontal axis and
2–3 in the vertical axis. Two diffuse brown and vertically elongate spots (or bars)
also are present in the humeral area, each one covering 1– 1·5 scales in the horizontal
axis and 2–2·5 in the vertical axis, the rst passing through the black humeral spot,
the second 2–4 scales behind the rst; in most specimens, the region just posterior
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1878 A. ANGULO ET AL.
Brycon guatemalensis
Roeboides guatemalensis
Hyphessobrycon compressus
Astyanax anai
Astyanax orthodus
Astyanax bimaculatus
Other North and Central
American species of
Astyanax, plus a few South
American species
F. 5. Strict consensus topology of 133 equally parsimonious trees (consistency index =0·26; retention
index =0·41) for the North and Central American species of Astyanax (summarized) from the analysis
of Schmitter-Soto (2016).
to the humeral spot is pale. A very thin black line of chromatophores originating
behind the second humeral spot and extending to the caudal-n base is also present.
Midlateral stripe formed by a pattern of 10–14 anteriorly directed herringbone or
chevron-shaped marks formed by deep pigmentation between myomeres, decreasing
in size and denition toward caudal peduncle and separated from each other by two to
three scales; these marks are most apparent in juveniles and in preserved specimens.
Caudal spot black, thick, elongated and rhomboid or rectangular in shape, with its
anterior margin surpassing the middle of the caudal peduncle, usually reaching the
anal-n insertion, not extending anteriorly as a continuous dark line or stripe on side
of body and posteriorly covering four to seven principal caudal-n rays; caudal spot
not extending onto the ventral and dorsal margins of the caudal peduncle and covering
three to ve horizontal scale rows. All ns (except the adipose n) with few dispersed
dark chromatophores, usually being more densely distributed at the membranes
between rays (principally in the dorsal, anal and caudal ns and on the mid-portion of
the anal n, but usually absent in their anterior, basal and distal portions).
COLOUR IN LIFE
Description based on live specimens photographed in eld aquaria just after capture.
General body colour silvery [Fig. 1(c), (d)]. Dorsal and dorsolateral portion of head and
body brown or greenish-brown. Midline of body silvery or metalic bluish to greenish.
Ventral portion of head and abdomen silvery to white. Infraorbital and opercular areas
usually covered with scattered, dark chromatophores. Scales on lateral portion of body
with dark brown chromatophores. A dark spot or band of variable size and shape
present on the posteriomedial portion of the eye, which, in most specimens, continues
posteriorly covering the skin above the dorsal portion of the infraorbital series (i.e.
infraorbitals III and IV) and part of the opercle. Humeral spot usually inconspicuous,
grey to black and rounded to horizontally ovate, covering two to four scales in the hori-
zontal axis and two to three in the vertical axis. Caudal spot black, thick, elongated and
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1879
rhomboid or rectangular in shape, with its anterior margin surpassing the middle of the
caudal peduncle, usually reaching the anal-n insertion and posteriorly covering four
to seven principal caudal-n rays; not extending onto the ventral and dorsal margins
of the caudal peduncle and covering three to ve horizontal scale rows. Dorsal, adi-
pose and caudal ns are yellowish to yellowish-brown; pectoral, pelvic and anal ns
are usually transparent at the base with some pigmentation (yellow to orange) distally
and in the case of the anal n at the base; all ns (except adipose n) with few dis-
persed dark chromatophores, usually being more densely distributed at the membranes
between rays (principally in the dorsal, anal and caudal ns and on the mid-portion of
the anal n, but usually absent in their anterior, basal and distal portions).
ETYMOLOGY
Astyanax anai is named in honour of the Asociación Anai (www.anaicr.org), given
their exceptional work in support of the knowledge and conservation of aquatic envi-
ronments and in the promotion and execution of sustainable development initiatives in
the Talamanca region of eastern Costa Rica– western Panama. The name of this new
species also represents a tribute to Ana R. Ramírez Coghi for her dedicated service,
since the year 2000, as the assistant collection manager of the UCR sh collection.
COMMON AND TECHNICAL NAMES
Largespot tetra (English); sardina, sardina blanca (Angulo, 2013), sardina anai, sar-
dina de Ana (Spanish).
GEOGRAPHIC DISTRIBUTION
This species appears to be endemic to the Sixaola River basin, eastern Costa
Rica–western Panama, Central America (Fig. 4).
ECOLOGICAL NOTES
Astyanax anai inhabits streams and ponds between 1 and 100 m above sea level,
in waters ranging from stagnant to swift current at temperatures ranging from 21
to 32∘C (Bussing, 1998, Asociación Anai, unpubl. data); in the Watsi River, Costa
Rica, this species has been found at temperatures reaching 36·3∘C (Asociación
Anai, unpubl. data). Astyanax anai is usually found in pools and backwaters, where
usually hides among roots, or in caves and undercuts (McLarney et al., 2008). This
species co-occurs with A. nicaraguensis, being relatively uncommon and usually
less abundant than the former, except in well-preserved environments, with calm and
shallow waters, where A. anai reaches greater relative abundances (McLarney et al.,
2008, Asociación Anai, unpubl. data). Other sh species that usually co-occur with
A. anai are: Eretmobrycon scleroparius (Regan 1908), Hyphessobrycon panamensis
Durbin 1908 (Characidae); Amatitlania kanna Schmitter-Soto 2007, Amatitlania
myrnae (Loiselle 1997), Cribroheros bussingi (Loiselle 1997), Parachromis loisellei
(Bussing 1989) (Cichlidae); Rhamdia guatemalensis (Günther 1864) (Heptapteri-
dae); Brachyhypopomus occidentalis (Regan 1914) (Hypopomidae); Alfaro cultratus
(Regan 1908), Phallichthys amates (Miller 1907), Poecilia gillii (Kner 1863) and
Priapichthys annectens (Regan 1907) (Poeciliidae).
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1880 A. ANGULO ET AL.
Astyanax anai is an omnivorous sh that feeds mainly on seeds, as well as terres-
trial and aquatic insects and reaches at least 110 mm in LSand 132 mm in total length
(Asociación Anai, unpubl.). Based on examined stomach contents of four specimens,
large quantities of terrestrial [mainly ants (Formicidae), beetles (Scarabaeidae and Cur-
culionidae) and small ies] as well as aquatic [mainly beetles (Elmidae)] insects were
identied; in addition, small seeds (about 1 mm in diameter) and other plant compo-
nents (parts of fruits, owers and leaves) were detected. Dagosta (2011), on the other
hand, reported small shes (members of the Characidae) in stomach contents of some
Panamanian specimens; this nding, however, as that author explains, could be an arte-
fact of the sampling method (i.e. rotenone).
PHYLOGENETIC RECONSTRUCTION
A total of 132 equally parsimonious trees were recovered. The consensus tree
obtained was 360 steps long, with consistency index IC=0·26 and retention index
IR=0·41 (Fig. 5). The phylogenetic reconstruction indicated that A. anai is more
closely related to A. orthodus (from Colombia) than to the other North and Central
American congeners. The clade composed of these two species was recovered as the
sister group of A. bimaculatus plus the clade formed by all remaining species of the
genus reviewed by Schmitter-Soto (2016) and also considered in this study.
Synapomorphies for the clade A. anai plus A. orthodus are: character 14, state
2=caudal side of supraoccipital convex in lateral view (see g. 7c in Schmitter-Soto,
2016); character 21: state 1 =contact between infraorbitals II and III narrower
(see Schmitter-Soto, 2016); character 25: state 0 =quadrate dorsal process distally
expanded (see g. 14a in Schmitter-Soto, 2016); character 49, state 1 =caudal process
of postcleithrum concave-sided dorsally, almost straight ventrally (see g. 25b in
Schmitter-Soto, 2016). Autapomorphies for A. anai are: character 39, state 1 =mean
7 or 8 scale rows between lateral line and pelvic-n origin; character 45, state
1=post-anal element longer (see g. 22b in Schmitter-Soto, 2016).
REMARKS
Astyanax anai and A. orthodus share a combination of characters unique among
all North and Central American congeners (i.e. predorsal scale series irregular and
incomplete, with an unscaled space behind tip of supraoccipital process; humeral
region with a conspicuous, black and rounded to horizontally ovate spot and two
diffuse vertical humeral bars; caudal peduncle with a thick and horizontally oval
black spot that continues over the median caudal-n rays), which is only present in
other South American congeners such as Astyanax bimaculatus (L. 1758), Astyanax
lacustris (Lütken 1875) and A. superbus, among others (Lucena & Soares, 2016;
Ruiz-C et al., 2018). The colouration pattern described above for such species was
used by several authors, including Garutti & Britski (1997), Garutti (1998) and Garutti
& Langeani (2009), to diagnose the ‘A. bimaculatus caudal peduncle spot’ complex,
a group that includes another 10–14 species, in addition to those already mentioned,
with essentially cis-Andean distributions [see Lucena & Soares, 2016 and Ruiz-C
et al., 2018 for an overview]. Based on the above and following the criteria of previous
authors, the placement of A. anai within the A. bimaculatus caudal-peduncle-spot
group seems logical.
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A NEW ASTYANAX FROM CENTRAL AMERICA 1881
Our phylogenetic reconstruction (despite not being very inclusive in taxonomic
terms) recovered the A. bimaculatus caudal-peduncle-spot complex as paraphyletic
(i.e. the clade formed by A. anai plus A. orthodus was recovered as the sister group
of A. bimaculatus plus the clade containing all remaining congeners). This result
agrees with an alternative hypothesis proposed by Dagosta (2011), Torres-Mejia et al.
(2012) and Lucena & Soares (2016) and supported partially by Mirande (2010),
which consider A. orthodus (as well as A. anai) and A. superbus more related to
Astyanacinus Eigenmann 1907 [current status =Astyanax; Eschmeyer et al. (2018)]
than to Astyanax. In fact, A. orthodus and A. superbus (as well as A. anai) share several
of the synapomorphies listed by Dagosta (2011) and Torres-Mejia et al. (2012) for
Astyanacinus [i.e. toothless portion of the premaxilla larger than its toothed portion;
posterior margin of the maxilla posterior to vertical through the wing of the lateral
ethmoid; lower jaw larger than eye-orbit; midlateral stripe formed by a pattern of
10–14 anteriorly directed herringbone or chevron-shaped marks; caudal peduncle
with a thick and horizontally oval caudal black spot that continues over the median
caudal-n rays; anal-n rays 25–36; dorsal-n rays ii, 9; pectoral-n rays i, 11–13];
however, other characters listed by Dagosta (2011) and Torres-Mejia et al. (2012) as
diagnostic for the genus (i.e. three maxillary teeth; four large dentary teeth; distance
between the postcleithrum one and two shorter than height of postcleithrum one; rst
anal-n pterygiophore dorsally directed; lack of hooks in ns) do not necessarily
match with the description of A. anai (and in the case of the rst two, they also are
variable in A. orthodus and A. superbus) (also see Ruiz-C et al., 2018).
Based on the aforementioned and considering that most of the synapomorphies listed
for Astyanacinus by Dagosta (2011) and previous authors (e.g. toothless portion of the
premaxilla larger than its toothed portion; lower jaw larger than orbit; caudal peduncle
with thick and horizontally oval caudal black spot that continues over median caudal-n
rays; anal-n rays 25–36; dorsal-n rays ii, 9; pectoral-n rays i, 11–13), are shared
with members of Astyanax; the uncertain relationship between species of these two
genera, we decided to keep the species herein described as new in Astyanax (and outside
the A. bimaculatus species group) until a detailed taxonomic study assessing these
taxonomic problems is carried out.
IDENTIFICATION KEY TO THE LOWER CENTRAL AMERICAN
(SOUTHERN NICARAGUA, COSTA RICA AND PANAMA) SPECIES OF
ASTYANAX, ADAPTED FROM SCHMITTER-SOTO (2017)
1a. Predorsal scale series irregular and incomplete, with an unscaled space behind
the tip of the supraoccipital process and 12–14, usually 13, scales; humeral spot
rounded to horizontally ovate; midlateral stripe formed by a series of 10– 14 anteri-
orly directed dermal herringbone or chevron-shaped marks; total anal-n elements
29–33 [Sixaola River basin; Costa Rica and Panama (Atlantic)]. . . . . . . . . . . . . . . .
..................................................... Astyanax anai sp. nov.
1b. Predorsal scale series regular and complete, without an unscaled space behind the
tip of the supraoccipital process and with 8–14, usually 10–12, scales; humeral
spot square, rhomboid, p-shaped or indistinct (oval in some specimens of A. brans-
fordii and A. cocibolca); midlateral stripe absent or not as described above; total
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1882 A. ANGULO ET AL.
anal-n elements usually 28 or less (22–32, usually 25–27 in A. nicaraguensis
and A. orstedii; 27–30, usually 28–29 in A. kompi) .........................2.
2a. In adults (>60 mm LS), premaxillary teeth unicuspid and enlarged at front of very
elongated jaws, juveniles usually with multicuspid teeth with central cusp long and
slender; maxillary teeth 9–13; gill-rakers on rst gill arch 26–27; total pectoral-n
elements 13–16 [Lake Nicaragua and Río Frío, San Juan, San Carlos, Sarapiquí,
Tortuguero and Parismina River basins; Nicaragua and Costa Rica (Atlantic)] . . .
. ......................................... Astyanax bransfordii (Gill 1877).
2b. In both adults and juveniles, premaxillary teeth usually tri or quin-cuspid (uni-
cuspid in A. cocibolca) with the central cusp robust and enlarged but usually not
elongated and the jaws not very elongated; maxillary teeth 0– 8 (9 in some speci-
mens of A. nicaraguensis); gill-rakers on rst gill arch usually 17– 24 (21–26 in A.
nicaraguensis, 23–34 in A. cocibolca); total pectoral-n elements usually 10–13
(rarely 14 or 15 in A. cocibolca and A. orstedii) ............................ 3.
3a. Mouth upturned; teeth at the outer row of premaxilla unicuspid; gill-rakers on
rst gill arch 23–34, usually 27–29; scale rows from lateral line to base of rst
dorsal-n ray 5–6, rarely 7 [Lake Nicaragua and Río Frío, San Juan, San Carlos,
Sarapiquí and Tortuguero River basins; Nicaragua and Costa Rica (Atlantic)] . . .
. ........................................ Astyanax cocibolca Bussing, 2008.
3b. Mouth terminal, rarely upturned; teeth at the outer row of premaxilla usually tri-
cuspid; gill-rakers on rst gill arch 17–26, usually 19–24; scale rows from lateral
line to base of rst dorsal-n ray usually 7–9 (6 in A. kompi) ................4.
4a. Premaxillary teeth 8 at the outer row and 8 at the inner row; scale rows from lateral
line to base of rst dorsal-n ray 6; body depth at dorsal-n origin 27·0–35·3%
LS, usually 29·1–33·3% [apparently endemic to the Grande (Gulnar) Lagoon,
Chririquí Viejo River basin; Panama (Pacic)] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. ........................................ Astyanax kompi Hildebrand, 1938.
4b. Premaxillary teeth 0–5, usually 4 or 5 at the outer row, 4–6, usually 4 or 5, at
the inner row; scale rows from lateral line to base of rst dorsal-n ray 7–9; body
depth, at dorsal-n origin 30·0–44·0% LS, usually 34·0–40·0% LS...........5.
5a. Head prole somewhat concave; caudal spot only on peduncle, not extending onto
n rays; lower lobe of caudal n slightly longer [from the Chiriquí to the Tuira
River basins; Panama (Pacic and Atlantic)] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
....................................... Astyanax panamensis (Günther 1864).
5b. Head prole usually straight (concave in some specimens of A. nicaraguensis);
caudal spot on peduncle and on n rays; lobes of caudal n subequal . . . . . . . . . 6.
6a. Maxillary relatively long, 9·7–13·0% LS, usually 10·2–11·9%, with 4– 6 teeth;
supraoccipital, long and narrow-based in dorsal view [apparently endemic to Lake
Managua; Nicaragua (Atlantic)] . . . . . . . . . . . . . . . Astyanax nasutus, Meek 1907.
6b. Maxillary relatively short, 5·4–10·8% LS, usually 6·5–9·7%, with 2 or 3 teeth [4 or
more (up to 9) only in a few specimens of A. nicaraguensis]; supraoccipital short
and wide-based in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7.
7a. Upper lip usually protruding; maxillary teeth 1 –9, usually more than 3; metaptery-
goid with a single posterodorsal projection; lower pharyngeal plate single, with its
caudal side concave; infraorbital III semicircular; proximal edge of the pelvic bone
straight [from the Matagalpa, including the Lake Nicaragua, to the Sixaola River
basins; Nicaragua and Costa Rica (Atlantic)] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
........................... Astyanax nicaraguensis Eigenmann & Ogle, 1907.
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A NEW ASTYANAX FROM CENTRAL AMERICA 1883
7b. Upper and lower lip about equal, upper lip usually not protuding; maxillary teeth
1–5, usually 2; metapterygoid with two posterodorsal projections; lower pharyn-
geal plate double, with its caudal side straight; infraorbital III angled; proximal
edge of the pelvic bone convex [From the San Juan to the Chiriquí River basins;
Nicaragua (Atlantic), Costa Rica (Pacic and Atlantic) and Panama (Pacic)] . . .
........................................... Astyanax orstedii (Krøyer 1875).
DISCUSSION
The new species described herein was assigned to Astyanax in accordance with the
denition of the genus by Eigenmann, 1921, which is still in use (Reis et al., 2003;
Camelier & Zanata, 2014; Bertaco & Vigo, 2015); i.e. two rows of premaxillary teeth;
ve teeth in the inner premaxillary series; lateral line complete; adipose n present;
caudal n naked. Within the genus, the new species ts the denition of the Astyanax
bimaculatus species complex, sensu Garutti & Britski (1997), Garutti (1998), Garutti &
Langeani (2009) and Ruiz-C et al. (2018); however, their inclusion in this group could
be problematic if A. anai (and other similar species) is more related to Astyanacinus,
sensu Dagosta (2011), than Astyanax, as suggested by Mirande (2010), Torres-Mejia
et al. (2012) and Lucena & Soares (2016). In any case, the species herein described
as new shows a combination of characters unique among members of both Astyanax
(e.g. colouration pattern and other morphological, morphometric and meristic, charac-
ters; see Diagnosis) and most Astyanacinus,sensu Dagosta (2011) and Torres-Mejia
et al. (2012) [e.g. maxillary teeth usually 4 (v. usually 2, 3 or more than 4); body
depth 36·6–42·3% of LS(v. usually 29·0–39·0% of LS); presence of hooks in the ns
(v. absence); see Diagnosis], which warrant its recognition as a valid species.
As mentioned, Astyanax is one of the most diverse and taxonomically complex
genera in the Characidae; its taxonomy and species-level relationships are far from
complete. Comprising about 150 valid species (Lucena & Soares, 2016), unambiguous
identication of species of Astyanax can be difcult. Addressing these problems at
local–regional scales and through approaches, such as those by Lucena & Soares
(2016) and Schmitter-Soto (2016, 2017), among several other examples, could be,
in the absence of a broad taxon sampling, the best and most feasible option. In this
study some questions have not been resolved (mainly in reference to the phylogenetic
relationships of the new taxa); however we hope that our results, in a critical com-
parison with those obtained in previous (Ornelas-Garcia et al., 2008; Schmitter-Soto,
2016) and future studies will contribute to a better understanding of the diversity and
taxonomy of Astyanax in lower Central American waters.
COMPARATIVE MATERIAL EXAMINED
Astyanacinus (Astyanax) moorii: MZUSP 78752, 1 (c&s), 40·86 mm LS, Saloba River
on the road to Estação Ecológica de Serra das Araras, Porto Estrela, Mato Grosso,
Brazil, collectors H.A. Britski, F. A. Machado and O. Froehlich, 27 February 2002.
Astyanax ‘bimaculatus’: FMNH 58067, 10, 27·39–79·52 mm LS, Urubamba River,
Santa Ana, Peru, coll. E. Heller, 1915; FMNH 84165, 5, 45·15–76·33 mm LS, Ucayali
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

1884 A. ANGULO ET AL.
River, sand beach across from Pucallpa, Peru, collectors D. W. Greeneld, T. A. Green-
eld and G. S. Glodek, 27 July 1975; FMNH 97395, 1, 94·32 mm LS, Natentza Creek,
tributary of the Nangaritza River, Zamora-Chinchipe, Ecuador, collectors R. Barriga,
M. C. Ibarra and R. Lema, 16 march 1979; FMNH 113729, 3, 51·13–74·35 mm LS,
Muchientsa River from mouth to 300m above, Morona-Santiago, Ecuador, collectors
B. Chernoff, R. Leon and P. S. Jua, 22 July 1999. Astyanax kompi: FMNH 36648, 1
(Paratype), Laguna Gulnar or Grande, Volcán, Pacic slope, Panama, collector S.F.
Hildebrand, 7 February 1935; USNM 106510, 1 (Holotype) 51·3mmLS, Laguna Gul-
nar or Grande, Volcán, Pacic slope, Panama, coll. S.F. Hildebrand, 7 February 1935.
Astyanax lacustris: DZSJRP 3297, 2 (c&s), 48·95–51·58 mm LS, afuent stream of
Turvo River, Grande River drainage, Palestina, São Paulo, Brazil, collectors V. Garutti
et al., 01 January 2000. Astyanax nicaraguensis: UCR 0891-02, 16, 42·96– 82·57 mm
LS, stream between Tuba Creek and Cahuita, province of Limón, Costa Rica, 9∘45′
50′′ N; 82∘53′49′′ W; 10 m, collectors W. A. Bussing, M. Bussing and E. Bussing, 3
October 1975; UCR 1144-11, 5, 26·84–95·81 mm LS, Cocolis River, 3·5 km south-east
of Shiroles, on Bratsi–Shiroles road, province of Limón, Costa Rica, 9∘34′4′′ N; 82∘
56′4′′ W; 40 m., collectors W. Bussing, K. Gocke and P. Siegfried, 13 November 1977;
UCR 1350-22, 5 (1 c&s), 59·42–100·52 mm LS, Barbilla River, on Siquirres – Limón
road, province of Limón, Costa Rica, 15 m, collectors C. De Paco, R. Villegas and G.
Umaña, 25 September 1981; UCR 2494-03, 21, 38·28–62·51 mm LS, La Suerte River,
near Cariari, province of Limón, Costa Rica, 10∘27′59′′ N; 83∘46′19′′ W; collector
W. Raschi, 22 June 1996. Astyanax orstedii: UCR 0047-05, 3, 66·42 – 101·51 mm LS,
Cañaza River north-west side of Golto at end of landing eld, province of Puntare-
nas, Costa Rica, 8∘39′24′′ N; 83∘11′9′′ W; 20m, coll. S. Salas, 7 august 1966;
UCR 0152-01, 3, 59·08–69·43 mm LS, articial canal 7 km south-west of Finca Exper-
imental Jiménez at Finca Taboga, province of Guanacaste, Costa Rica, 10∘21′20′′ N;
85∘11′14′′ W; 10 m, collectors W. A. Bussing and T. Lodge, 13 July 1967; UCR
0172-01, 3, 62·05–82·93 mm LS, pool of Riyito River at end of street near Charcos,
30–50 m upstream, province of Puntarenas, Costa Rica, 8∘40′9′′ N; 83∘30′19′′ W;
10 m, collector K. Karlson, 28 February 1967; UCR 0757-01, 3, 45·99–81·58 mm LS,
Angelina Creek 0·5 km NO of Villa Briceño at Interamerican Highway, province of
Puntarenas, Costa Rica, 8∘43′29′′ N; 83∘10′4′′ W; 60m, collectors W. A. Bussing,
M. Bussing and E. Bussing, 1 March 1973; UCR 1370-02, 3, 36·46–69·03 mm LS,
Agua Mala Creek, 500 m north of Planta Palo Seco, 3·5 km east of Parrita, province
of Puntarenas, Costa Rica, 9∘32′9′′ N; 84∘17′29′′ W; collector W. A. Bussing, 26
march 1982; UCR 1465-01, 3, 36·70–48·13 mm LS, Machuea River, 3 km north-east
of San Mateo, Orotina road, province of Alajuela, Costa Rica, 9∘57′10′′ N; 84∘30′
4′′ W; 300m, collector W. A. Bussing, 15 April 1983. Astyanax orthodus: CAS 68770,
3, 51·7–102·6mmLS, Altacar Creek, a tributary to Río Telembi, at Barbacoas, Nar-
ino Colombia, 1∘40′32′′ N; 78∘8′21′′ W; collectors A. W. Henn and C. Wilson;
CAS 68771, 1, 66·2mmLS, Quibdo, at the junction of Rios Quito and Atrato, Chocó,
Colombia, 5∘41′3′′ N; 76∘40′1′′ W; collector C. H. Eigenmann; FMNH 10263, 7,
41·22–70·57 mm LS, Truando, Colombia, collector C. E. Wilson; FMNH 56605, 2,
91·32–106·74 mm LS, Barbacoas, Colombia, collector A. W. Henn, 17 January 1913;
FMNH 56578, 12, 46·59–75·80 mm LS, Truando, Colombia, collector C. E. Wilson,
1913; UCR 1958-01, 10 (2 c&s), 55·7–74·7mmLS, Truando River near Sucio River,
Colombia, collector C. E. Wilson; USNM 55655, 1 (holotype; X-ray), 90·20 mm LS,
Truando, Colombia, collector M. Schott. Astyanax nasutus: FMNH 5909, 1 (holotype),
© 2018 The Fisheries Society of the British Isles, Journal of Fish Biology 2018, 92, 1866–1887

A NEW ASTYANAX FROM CENTRAL AMERICA 1885
Lake Managua, Nicaragua, collector. S. E. Meek, March 1906. Astyanax panamensis:
UCR 0434-08, 12, 43·00–81·93 mm LS, López River at Transisthmian road, 9 miles S
Colón, Colón, Panama, collector E. Tyson, 28 February 1970. Astyanax superbus: CAS
36490, 1 (Holotype; X-ray), 81·52 mm LS, small brook tributary to Río Tamanaco at
Camoruco, 20 km northeast of San Carlos, Cojedes, Venezuela.
The authors are grateful to S. Mochel, K. Swagel, J. Bates (FMNH), C. Baldwin (USNM), A.
R. Ramírez (UCR), R. Silva Costa-Ferreira (UNESP) and the authorities and the administrative
staff of the FMNH, USNM, UCR and UNESP for encouragement and the use of facilities. The
authors are also grateful to A. M. Arias, M. Maa and W. McLarney (Asociación Anai) for kindly
providing photos (Fig. 1) and information on the ecology, distribution and live coloration pattern
of A. anai. J. Fong, D. Catania and E. Salas (CAS) kindly provided photos and information about
some specimens of A. anai,A. orthodus and A. superbus. The authors also are grateful to W.
McLarney, W. Matamoros (UNICACH) and several anonymous reviewers for the very useful
criticisms and the suggestions offered which improved substantially the content of this paper.
A.A. was supported by the Costa Rican Ministerio de Ciencia, Tecnología y Telecomunicaciones
(MICITT; PED-017-2015-1), by the Partnerships Program for Education and Training of the
Organization of American States and the Grupo Coimbra de Universidades Brasileiras (PAEC
OEA-GCUB; 2014) and by the Visiting Scholarships Program of the Field Museum of Natural
History (FMNH; 2015). F.L. is grateful to the Conselho Nacional de Desenvolvimento Cientíco
e Tecnológico (CNPq) for the scientic support.
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