Content uploaded by Juan J Schmitter-Soto
Author content
All content in this area was uploaded by Juan J Schmitter-Soto on Jun 24, 2022
Content may be subject to copyright.
Accepted by M.R. de Carvalho: 29 Feb. 2016; published: 6 May 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4109 (2): 101
–
130
http://www.mapress.com/j/zt/
Article
101
http://doi.org/10.11646/zootaxa.4109.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:A084F919-17D2-41E1-90EB-0B6AF576C272
A phylogeny of Astyanax (Characiformes: Characidae)
in Central and North America
JUAN J. SCHMITTER-SOTO
El Colegio de la Frontera Sur (ECOSUR), A.P. 424, MX-77000 Chetumal, QR, Mexico. E-mail: jschmitt@ecosur.mx
Abstract
A phylogeny is presented for 34 species of Astyanax, 27 of them once included within A. aeneus or A. fasciatus in Central
America and Mexico, based on 52 morphological characters (mostly osteological, but also pigmentation and meristics),
with three outgroups. Monophyly is not supported for A. aeneus s. lat., as Brazilian species such as A. fasciatus s. str. and
others occur also within that clade. There were only five resolved clades, three of them including both Brazilian and Cen-
tral American species, one purely Nicaraguan, and one for central-northern Mexico and Texas. Coincidence with previous
cladistic hypotheses is only partial. The genus Bramocharax Gill is not recovered, and thus confirmed as a synonym of
Astyanax Baird & Girard. The findings point at a more complex biogeographic history of the region than usually recog-
nized.
Key words: Osteology, cladistics, Bramocharax, characids, tetragonopterines, Middle America
Resumen
Se presenta una filogenia para 34 especies de Astyanax, 27 de ellas incluidas alguna vez en A. aeneus o A. fasciatus
en Centroamérica y México, con base en 52 caracteres morfológicos (sobre todo osteológicos, pero también de
pigmentación y merística), con tres grupos externos. No se apoya la monofilia de A. aeneus s. lat., dado que
especies brasileñas, como A. fasciatus s. str. y otras, aparecen dentro de tal clado. Hubo sólo cinco clados definidos,
tres de ellos con especies de Brasil y de Centroamérica, uno puramente nicaragüense, y uno del centro-norte de
México y Texas; la coincidencia con hipótesis cladísticas previas es sólo parcial. El género Bramocharax Gill no se
recupera, de modo que se confirma como sinónimo de Astyanax Baird & Girard. Según los hallazgos, la historia
biogeográfica de la región es más compleja de lo que se generalmente se reconoce.
Introduction
Few authors have attempted to provide a cladistic hypothesis for Central American and Mexican Astyanax, perhaps
because the alpha taxonomy is not clear to begin with. The prevailing views have been that virtually all Astyanax
forms in the study area belong either to A. fasciatus s. lat. (restricted to Rio São Francisco, Brazil, by Melo 2005),
to A. aeneus s. lat. and, in northern Mexico and Petén (Miller et al. 2009), to A. mexicanus.
Populations identifiable as Astyanax aeneus sensu lato (sometimes as A. fasciatus s. lat. or A. mexicanus s. lat.)
in Central America and Mexico were molecularly analyzed by Ornelas-García et al. (2008), who recommended
several species to be recognized. This suggestion is being examined by Schmitter-Soto (unpubl. data), who
describes nine new species and resurrects nine more. The present contribution is a phylogenetic study of these
species.
Valdez-Moreno (1997) used 30 cranial characters for 23 Mexican populations and found osteological
synapomorphies for the northern Mexican populations that validated their earlier recognition (e.g. Miller 1986) as
A. mexicanus (de Filippi 1853), but she was unable to diagnose the southern Mexican – northern Middle American
SCHMITTER-SOTO
102
·
Zootaxa 4109 (2) © 2016 Magnolia Press
populations, provisionally called A. aeneus (Günther 1860, non Hensel 1870) by recent authors (e.g. Greenfield &
Thomerson 1997; Bussing 1998; Schmitter-Soto et al. 2008; Miller et al. 2009). She also suggested that the
population at Tamazulapan was a new species. Later on, Valdez-Moreno (2005) erected a phylogeny of 19 characid
species, among which only two Astyanax, A. bimaculatus and A. mexicanus, and found the genus Bramocharax to
be monophyletic, a finding contested by Ornelas-García et al. (2008) and by the present work.
The results of Strecker et al. (2004) support Valdez-Moreno’s (1997), inasmuch as they found only one
mtDNA lineage in northern Mexico, but at least five in the south. On the other hand, in their molecular cladogram
for 147 populations, Ornelas-García et al. (2008) found two lineages in northern Mexico and several in Middle
America, many of these considered as possible new species. As stated by Hausdorf et al. (2011), “…further
investigations are necessary to show which […] clades classified by Ornelas-García et al. (2008) as separate
species represent independently evolving entities.” However, Hausdorf et al. (2011) found only one slightly mixed
population of A. mexicanus, with 4% individuals having nuclear genes from A. aeneus s. lat.
Mirande (2010) included one Central American species of Bramocharax and one North American species of
Astyanax in his osteology-based phylogenetic hypothesis of the Characidae. He found them to lie in quite distant
clades, contradicting all previous work.
The present phylogenetic reconstruction does not recover monophyly of A. aeneus s. lat. However,
autapomorphies support the diagnosis of several new and resurrected species (Schmitter-Soto, unpubl. data) and
the phylogeny, although not completely resolved, suggests an interesting, complex biogeographic history of the
region, given the tendency for South American forms to be nested within Middle American clades.
The objective of this contribution is to propose a phylogeny for the former species complex of A. aeneus s. lat.
and close species in the same genus in Characidae, based on morphology (mostly osteology), and to compare it to
other hypotheses, especially the molecular phylogeny by Ornelas-García et al. (2008) and the cladogram for
Mexican populations based on cranial osteology by Valdez-Moreno (1997). All species of genus Bramocharax are
included, as this taxon has been proven to be polyphyletic (Ornelas-García et al. 2008).
Material and methods
The specimens examined (Appendix 1) include representatives from all nominal species that have at various times
been synonymized with Astyanax aeneus s. lat. (Schmitter-Soto, unpubl. data), as well as comparative material of
other species in the genus and all species of Bramocharax. Brycon guatemalensis, Hyphessobrycon compressus,
and Roeboides guatemalensis were chosen as outgroups, following previous authors (Valdez-Moreno 1997, 2005;
Ornelas-García et al. 2008), in addition to three species of Astyanax never determined as A. fasciatus s. lat.: A.
bimaculatus, A. atratoensis, and A. orthodus All specimens were observed by the author, except for H. compressus
and R. guatemalensis, coded based on the detailed illustrations by Valdez-Moreno (2005).
Osteological methodology (i.e., clearing and staining) followed Taylor and van Dyke (1985), with the
following minor modifications (W. Fink, pers. comm.): before dehydration, specimens were bleached in 50% H
2
O
2
for ca. 3 h; after alcian blue staining and neutralization in a saturated borax solution, clearing proceeded by
changing the enzyme buffer solution (30% borax solution, with trypsin) every 3 days, with the specimen over a
neon-light box, then staining with alizarin red, returning to the clearing regime, rinsing, and equilibrating in
glycerin. Radiographs (of type specimens) and skeletonized material were also used.
Cladistic analyses were based on maximum parsimony and reconstructions were performed with program
PARS-discrete character parsimony in package PHYLIP, version 3.69 (Felsenstein 2009); character states were
considered as unordered, multistate, or binary, and all received the same weight. Meristic characters were used only
if satisfactorily amenable to statistical coding: their states were determined by means of an ANOVA followed by a
Duncan a posteriori test (“homogeneous subset coding”, Simon 1983), with differences significant at α < 0.05
marking different character states.
The strict consensus tree was obtained and optimized using the package Mesquite, version 2.75 (Maddison &
Maddison 2011), to arrive at a phylogenetic hypothesis. Optimization followed the ACCTRAN option in order not
to lose any putative synapomorphy (Kitching et al. 1998). Homoplasy levels were calculated with Mesquite, as
consistency and retention indices, for the tree (CI, RI) and also for every character (ci, ri) (Archie 1989, Farris
1989).
Zootaxa 4109 (2) © 2016 Magnolia Press
·
103
PHYLOGENY OF ASTYANAX
Results
Character description and analysis. The following list includes 52 characters (see matrix, Table 1), selected out
of 90 screened traits mostly because they were amenable to reasonably objective coding. Some of them have been
used before, but all relevant anatomical material was reevaluated; I compare my interpretations mainly against
those of Valdez-Moreno (1997, 2005) and Mirande (2010). Many characters (22) are binary, but most are
multistate, non-additive, unordered. I did not exclude autapomorphies, because they provide special support for
species diagnoses (Schmitter-Soto, unpubl. data); to avoid bias, I did not leave out polymorphic and other highly
homoplastic traits either.
Gill arches, characters 1–6
1. Upper pharyngeal tooth plates, contour
[0] oval, fusiform (Fig. 1a); [1] crescent-shaped (Fig. 1b); [2] S-shaped (Fig. 1c)
Fifteen steps, ci=0.20, ri=0.14. State 1, homoplastically synapomorphic for A. cf. fasciatus “Ceará” + A.
panamensis; state 2, homoplastically synapomorphic for the northern Mexican-Texan clade. Several
polymorphisms and homoplastic autapomorphies.
This is Valdez-Moreno’s (1997) character 20, where she distinguished two states, similar to the ones
established here: 0, oval; 1, S-shaped; apparently she conflated my states 0 and 1 into her state 0.
FIGURE 1. Upper pharyngeal tooth plates: (a) oval, fusiform (Astyanax aeneus, UMMZ 191719); (b) crescent-shaped (A.
panamensis, GCRL 13409); (c) S-shaped (A. “Quiché”, UMMZ 193886). Bars are 0.2 mm long.
2. Lower pharyngeal tooth plates
[0] single (Fig. 2a); [1] double (Fig. 2b)
State 1 occurs only in A. cocibolca and A. “Costa Rica”, convergent autapomorphies; two steps, ci=0.50,
ri=0.00.
FIGURE 2. Lower pharyngeal tooth plates: (a) single (Astyanax aeneus, UMMZ 178568), illustrating also caudal side
concave; (b) double (A. “Costa Rica”, UMMZ 243884), illustrating also caudal side straight.
SCHMITTER-SOTO
104
·
Zootaxa 4109 (2) © 2016 Magnolia Press
3. Lower pharyngeal tooth plates, caudal side
[0] concave (Fig. 2a); [1] straight (Fig. 2b)
Synapomorphic homoplastically for A. cf. fasciatus “das Velhas” + A. cocibolca and for B. bransfordii + A.
nasutus; also in A. “Costa Rica”. Three steps, ci=0.33, ri=0.50.
4. Epibranchial III, insertion of uncinate process
[0] widely open, “hyperbolic” (Fig. 3a); [1] rather closed, semicircular (Fig. 3b)
State 1 is synapomorphic for “A. fasciatus s. lat.” (i.e. all Central American and Mexican Astyanax species,
plus A. fasciatus s. str. and other Brazilian material close to A. fasciatus). Nine steps; rather homoplastic (ci=0.22),
but providing some structure (ri=0.50). A synapomorphic reversion to state 0 for the clade A. nicaraguensis + (A.
cf. fasciatus “das Velhas” + A. cocibolca); also autapomorphically in A. “Petén”, A. aeneus, A. “Tehuacán”, and A.
mexicanus. Some species polymorphic.
FIGURE 3. Epibranchial III, insertion of uncinate process: (a) widely open; distal segment straight (Astyanax “Texas”, UMMZ
170107); (b) rather closed; distal segment curved (A. “Acatlán”, UMMZ 191698). Bars are 0.5 mm long.
5. Epibranchial III, main body, distal segment
[0] straight (Fig. 3a); [1] curved (Fig. 3b)
Seven steps, ci=0.29, ri=0.17. A synapomorphy for the northern Mexican-Texan clade, autapomorphical in A.
“Veracruz”, polymorphic in several species.
6. Gill rakers, first arch, total number
[0] 26 or more; [1] 19–26, mean 22–25; [2] 18–23, mean 20–21; [3] 16–20, mean 17–19
Fourteen steps, ci=0.29, ri=0.17. State 1 is synapomorphic for the studied Astyanax; state 2, synapomorphic for
the northern Mexican-Texan clade, as convergent autapomorphies in Hyphessobrycon and several Astyanax. State
3 is an autapomorphy for A. cocibolca, occurring also in Roeboides.
This was one of the classic diagnostic characters used to key out “A. mexicanus” sensu Miller et al. (2009), i.e.
including A. “Quiché”, with state 2, vs. “A. aeneus” s. lat., supposedly with state 1 (actually polymorphic for the
trait).
Hyoid series, characters 7–10
7. Urohyal, ventrorostral edge
[0] convex, uniform (Fig. 4a); [1] angled (Fig. 4b); [2] spiny (Fig. 4c)
Three steps, ci=1.00, ri=0.00; states 1 and 2 are strict autapomorphies for A. cf. fasciatus “Ceará” and A.
“Tehuacán”, respectively. Astyanax “Quiché” is polymorphic for the trait.
8. Urohyal, ventral apex
[0] almost at caudal end (Fig. 4d); [1] about equidistant between rostral and caudal ends (Fig. 4a); [2] somewhat
closer to caudal end (Fig. 4b)
Seventeen steps, ci=0.18, ri=0.13. In addition to convergent autapomorphies for several Astyanax, state 1 is
homoplastically synapomorphic for A. cf. fasciatus “Ceará” + A. panamensis; also in Roeboides. State 2 is a
synapomorphy for all the studied Astyanax, with a reversion to state 1 in A. atratoensis. A reversion to state 0 is
autapomorphic for B. dorioni and for A. “Tamazulapan”. Astyanax “Costa Rica”, A. “Petén”, and A. nicaraguensis
are polymorphic for the trait.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
105
PHYLOGENY OF ASTYANAX
Valdez-Moreno (1997) found all characteristics of the urohyal to be variable, except the rostral tip, her
character 17, which she coded as “round”, state 0, vs. “pointed”, state 1. The bone was more useful at the generic
level: Valdez-Moreno (2005) used its relative length and thickness as her characters 69 and 70, respectively.
FIGURE 4. Urohyal: (a) ventrorostral edge convex, uniform; ventral apex about equidistant between rostral and caudal ends
(Astyanax mexicanus, UMMZ 169835); (b) ventrorostral edge angled; ventral apex closer to caudal end (A. “Quiché”, UMMZ
161769); (c) ventrorostral edge with a spine (arrow) (A. “Tehuacán”, UMMZ 198853); (d) ventrorostral edge convex; ventral
apex almost at caudal end (A. “Tamazulapan”, UMMZ 234194). Bars are 1 mm long.
9. Ceratohyal, foramen
[0] oval (Fig. 5a); [1] drop-shaped (Fig. 5b); [2] circular; [3] comet-shaped (Fig. 5c); [4] absent
Fifteen steps, ci=0.33, ri=0.09. State 1 is homoplastically autapomorphic for A. atratoensis, A. “Campeche”,
and B. dorioni. State 2 is a strict autapomorphy for A. cf. fasciatus “das Velhas”. State 3 occurs convergently in two
Brazilian species and in A. nasutus; state 4, in B. bransfordii, A. “Macal”, and A. “Texas”. Three out of seven
species in the Atlantic southern Mexican-Guatemalan clade are polymorphic for the trait.
FIGURE 5. Ceratohyal: first three specimens, rostral vertices angled: (a) foramen oval (Astyanax mexicanus, UMMZ 169835);
(b) foramen drop-shaped (A. “Campeche”, UMMZ 196571); (c) foramen comet-shaped (A. cf. fasciatus “Alto São Francisco”,
UMMZ 216281); (d) foramen oval, rostral vertices round (A. “Bacalar”, UMMZ 196540). Bars are 0.5 mm long.
SCHMITTER-SOTO
106
·
Zootaxa 4109 (2) © 2016 Magnolia Press
10. Ceratohyal, rostral vertices
[0] round (Fig. 5d); [1] angled (Fig. 5a,b,c)
Ten steps, ci=0.20, ri=0.33. State 1 is synapomorphic for the subgenus Astyanax; several reversions and
polymorphisms lower the consistency for this character.
Neurocranium, characters 11–18
11. Anterior fontanel, length
[0] longer (Fig. 6a); [1] shorter (Fig. 6b)
Nine steps, ci=0.22, ri=0.42. State 1 is homoplastically synapomorphic for Astyanax cf. fasciatus “Ceará” + A.
panamensis, for A. “Rioverde” + A. “Texas”, and for the Atlantic southern Mexican-Guatemalan clade, with a
reversion to state 0 for A. “Campeche” + A. “Ocotal” and polymorphisms in A. “Petén” and A. “Quiché”.
FIGURE 6. Anterior fontanel: (a) longer, sides convex at mid-length (arrow), tip sharp (Astyanax “Texas”, UMMZ 170107);
(b) shorter, sides straight at mid-length (arrow), tip blunt (A. “Veracruz”, UMMZ 97336).
12. Anterior fontanel, sides
[0] straight (Fig. 6b); [1] convex (Fig. 6a)
Five steps, ci=0.40, ri=0.50. State 1 is homoplastically synapomorphic for Astyanax cf. fasciatus “Alto São
Francisco” + A. “Macal” + A. altior, and A. “Rioverde” + A. “Texas”, with two other autapomorphies. Polymorphic
for A. “Costa Rica”.
13. Anterior fontanel, rostral tip
[0] blunt (Fig. 6b); [1] sharp (Fig. 6a)
Fifteen steps, ci=0.20, ri=0.20. State 1 is synapomorphic for the two Nicaraguan clades and also for A.
“Rioverde” + A. “Texas”, with the strictly autapomorphic state 2 for A. cf. fasciatus “das Velhas”. Several
polymorphisms.
14. Caudal side of supraoccipital, lateral view
[0] undulate to straight (Fig. 7a); [1] slightly concave (Fig. 7b); [2] convex (Fig. 7c); [3] angled (Fig. 7d); [4]
strongly concave.
Nineteen steps, ci=0.26, ri=0.26. State 1 is homoplastically synapomorphic for A. atratoensis + the large clade
that includes A. fasciatus s.s. and A. aeneus. State 2 is a strict autapomorphy for A. orthodus; state 3 occurs in A.
nasutus, A. nicaraguensis, A. “Veracruz”, and A. “Tehuacán”, with state 4 a convergent autapomorphy for B.
bransfordii and for A. “Tamazulapan”. Polymorphisms and reversions to state 0 in the sister group to A.
“Veracruz”.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
107
PHYLOGENY OF ASTYANAX
FIGURE 7. Supraoccipital, caudal side: (a) undulate to straight (Brycon guatemalensis, UMMZ 190656); (b) slightly concave
(Astyanax mexicanus, UMMZ 169835); (c) convex (A. orthodus, UMMZ 160225); (d) angled (A. “Veracruz”, UMMZ 191727)
. Bars are 1 mm long.
15. Supraoccipital, caudal process, dorsal view
[0] longer, narrow-based (Fig. 8a); [1] shorter, wide-based (Fig. 8b)
Nine steps, ci=0.11, ri=0.20. State 1 is a synapomorphy for the studied species of Astyanax + Hyphessobrycon,
with several reversions, including one for B. bransfordii + A. nasutus.
This is Valdez-Moreno’s (1997) character 26, same coding and polarity. We do, however, differ in particular
evaluations: e.g., she found A. “Veracruz”, which she considered a series of intergrades between southern and
northern forms, to be polymorphic for the trait (nine populations examined), while state 1 is found here to be
constant for the species (eight populations examined).
Mirande (2010) separated the length of the supraoccipital spine in two binary characters, comparing it to the
length of the neural complex of the Weberian apparatus: in state 0 of his character 52, the spine is as long as the
complex; state 1 of character 52, to one half the extent of the complex; state 0 of character 53 includes both states
of character 52 (“spine extends to at least middle length of complex”); state 1 of character 53, only to anterior limit
of complex. Apparently, we coincide in the polarity found, and he faced several polymorphisms as well.
FIGURE 8. Supraoccipital, dorsal view: (a) longer, narrow-based (Astyanax aeneus, UMMZ 178568); (b) shorter, wide-based
(A. “Quiché”, UMMZ 173731) . Bars are 0.5 mm long.
16. Interorbital width
[0] mean 8% SL or less; [1] mean 9% SL or more.
Eleven steps, ci=0.18, ri=0.47. State 1, synapomorphic for Astyanax; reversions to state 0 for Panamanian and
Nicaraguan clades. Several polymorphisms.
The character was considered suitable for cladistic analysis because, unlike an osteologically composite
measurement (like head length), interorbital width involves only the frontal bones. For coding, see Methods
(ANOVA results available from the author on request).
SCHMITTER-SOTO
108
·
Zootaxa 4109 (2) © 2016 Magnolia Press
17. Supracephalic profile
[0] convex (Fig. 9a); [1] concave (Fig. 9b)
Seventeen steps, ci=0.12, ri=0.21. State 1, synapomorphic for the studied Astyanax (parallel in Roeboides); a
reversion to state 0 is synapomorphic for the clade A. mexicanus + (A. “Rioverde” + A. “Texas”). The clade B.
bransfordii + A. nasutus is polymorphic, as are 50% of the species in the Atlantic southern Mexican-Guatemalan
the clade and elsewhere.
As in character 16, this externally observable trait involves only the frontal bones.
FIGURE 9. Supracephalic profile: (a) convex (Astyanax mexicanus, UMMZ 169835); (b) concave (Astyanax aeneus, UMMZ
178568).
Infraorbitals, characters 18–21
18. Infraorbital II
[0] rectangular; [1] triangular, base short, angled (Fig. 10a); [2] triangular, base short, convex (Fig. 10b); [3] an
elongated triangle, base angled (Fig. 10c); [4] triangular, base with two angles (Fig. 10d); [5] triangular, base long,
convex (Fig. 10e)
Thirteen steps, ci=0.46, ri=0.13. State 1, synapomorphic for the studied species of Astyanax. State 2 appears in
parallel in Hyphessobrycon, A. orthodus, and others. State 3 is a strict autapomorphy for A. cf. fasciatus “das
Velhas”; state 4, for B. bransfordii. State 5 evolved frequently in the A. mexicanus clade, or else may be interpreted
as synapomorphic, with a reversion to state 1 in A. “Rioverde”; parallel in A. “Tamazulapan”. Trait polymorphic in
A. “Belize”.
This is Valdez-Moreno’s (1997) character 1. State 0 is the same, but she discerns just two more states: 1,
inferoposterior edge (“base”) short, and 2, long; I concur with her in finding a longer base in the northern form (A.
“Texas”). Valdez-Moreno (2005) distinguished two characters for the bone: anterior shape (square vs. triangular)
and posterior shape (square, semicircular, globose, or triangular).
19. Infraorbital III, infraposterior side
[0] angled, hyperbolic (Fig. 11a); [1] semicircular (Fig. 11b)
Eleven steps, ci=0.18, ri=0.10. State 1, synapomorphic for Astyanax + Hyphessobrycon; several reversions and
polymorphisms.
Valdez-Moreno (1997) coded this bone as her character 2, with three states: 0, squarish, equivalent to what I
call “angled” or hyperbolic; 1, semicircular with dorsoposterior projection, and 2, semicircular without such
projection. I find the alleged projection to be very variable, even within populations; her states 1 and 2 appear in
populations of A. “Veracruz”, a species which I also find polymorphic, but for my states 0 and 1. On the other hand,
the polarity was found to be the same.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
109
PHYLOGENY OF ASTYANAX
FIGURE 10. Infraorbital II: (a) triangular, base short, angled (Astyanax “Rioverde”, UMMZ 192510); (b) triangular, base
shorter, convex (A. aeneus, UMMZ 178568); (c) an elongated triangle, base angled (A. cf. fasciatus “das Velhas”, UMMZ
216372); (d) triangular, base with two angles (B. bransfordii, FMNH 5919); (e) triangular, base longer, convex (A. mexicanus,
UMMZ 169835). Bars are 0.5 mm long.
FIGURE 11. Infraorbital III, infraposterior side: (a) angled, hyperbolic (Astyanax aeneus, UMMZ 144617); (b) semicircular
(A. mexicanus, UMMZ 169835).
20. Infraorbital IV
[0] square, no projections (Fig. 12a); [1] square, with a rostroventral projection (Fig. 12b); [2] pentagonal (Fig.
12c); [3] rectangular, no projections (Fig. 12d); [4] rectangular, with projections (Fig. 12e)
Fourteen steps, ci=0.36, ri=0.36. State 1, synapomorphic for the studied species of Astyanax. State 2, a strict
autapomorphy for A. bimaculatus; state 3, for A. cf. fasciatus “das Velhas”. State 4 is homoplastically
synapomorphic for the Atlantic southern Mexican-Guatemalan clade and for A. “Rioverde” + A. “Texas”. This
bone is triangular in Roeboides (Valdez-Moreno 2005).
This is Valdez-Moreno’s (1997) character 3, same polarity found, although she splits my state 1 in two: 1,
projection smaller; 2, projection larger, and she omits my states 3 and 4. Her states 1 and 2 occur polymorphically
SCHMITTER-SOTO
110
·
Zootaxa 4109 (2) © 2016 Magnolia Press
in A. “Veracruz”, A. “Tamiahua”, and A. “Texas”, which reinforces my view that they should be considered only
one state (with projection).
This is also Mirande’s (2010) character 67, but he conflates all of my states into his state 0, square or longer
longitudinally, and recognizes only one other state, longer dorsoventrally than longitudinally, which does not occur
in any of the species I studied. His character 68 deals with a posterior dorsoventral expansion which does not
correspond to any of my states, although it is reminiscent of what I describe as pentagonal, my state 2, a form that
he found in Oligosarcus.
FIGURE 12. Infraorbital IV: (a) square, no projections (Astyanax “Tamiahua”, UMMZ 167489); (b) square, with a
rostroventral projection (A. mexicanus, UMMZ 169835); (c) pentagonal (A. bimaculatus, UMMZ 206863); (d) rectangular, no
projections (A. cf. fasciatus “das Velhas”, UMMZ 216372); (e) rectangular, with a projection (A. “Texas”, UMMZ 170107).
Bars are 0.5 mm long, except in (a), which is 0.2 mm long.
21. Contact between infraorbital II and III
[0] wider; [1] narrower
Eight steps, ci=0.25, ri=0.00. State 1 is homoplastically autapomorphic for A. aeneus, A. nicaraguensis, and A.
“Tamazulapan”; several instances of polymorphism.
Mouth and jaws, characters 22–28
22. Mouth
[0] upper lip protruding; [1] not upturned, even; [2] upturned; [3] lower lip protruding; [4] elongated
Ten steps, ci=0.40, ri=0.14. State 1, synapomorphic for the studied species of Astyanax + Hyphessobrycon.
State 2, convergently autapomorphic for A. “Macal” and for A. cocibolca; same situation for state 3, for A.
nicaraguensis and for A. nasutus. State 4 arose independently in all species of Bramocharax, including A.
“Ocotal”, considered a Bramocharax by Valdez-Moreno (2005).
Zootaxa 4109 (2) © 2016 Magnolia Press
·
111
PHYLOGENY OF ASTYANAX
23. Metapterygoid, dorsorostral projections
[0] none (Fig. 13a); [1] one (Fig. 13b); [2] two (Fig. 13c)
Thirteen steps, ci=0.23, ri=0.23. State 2 is synapomorphic for the studied species of Astyanax +
Hyphessobrycon; state 1, homoplastically synapomorphic for A. cf. fasciatus “Ceará” + A. panamensis and for the
Nicaraguan clade that includes A. cf. fasciatus “das Velhas”; also in Roeboides and other Astyanax. Several
polymorphisms.
This is Valdez-Moreno’s (1997) character 15, with three states, although she includes not only the number of
dorsorostral projections, but also the absence of a foramen in state 0; however, the foramen can also be absent in
state 2 (see Fig. 12c). I agree with her that A. aeneus, A. “Veracruz”, and A. “Texas” are polymorphic for the trait.
Valdez-Moreno (2005) used five characters from this bone. Her character 54, state 0, dorsoposterior margin
with two processes, was a synapomorphy uniting Bramocharax, Astyanax, Deuterodon, and Knodus.
FIGURE 13. Metapterygoid: (a) no dorsorostral projections, rostral arm somewhat longer than ventral (Brycon guatemalensis,
UMMZ 190656); (b) one dorsorostral projection, rostral arm much longer (Astyanax fasciatus, UMMZ 216281); (c) two such
projections, arms subequal (B. dorioni, UMMZ 193918). Bars are 1 mm long.
24. Arms of metapterygoid
[0] rostral arm somewhat longer than ventral (Fig. 13a); [1] rostral much longer than ventral (Fig. 13b); [2] arms
subequal (Fig. 13c)
Five steps, ci=0.50, ri=0.75. State 1, a strict synapomorphy for the subgenus Astyanax; state 2, strictly
autapomorphic for A dorioni. A reversion to state 0, synapomorphic for B. bransfordii + A. nasutus. Polymorphism
in A. “Costa Rica”.
25. Quadrate dorsal process
[0] distally expanded (Fig. 14a); [1] not expanded (Fig. 14b)
Thirteen steps, ci=0.15, ri=0.27. State 1, synapomorphic for the studied species of Astyanax +
Hyphessobrycon. A reversion to state 0, synapomorphic for the clade including A. mexicanus and for the Atlantic
southern Mexican-Guatemalan clade; convergently autapomorphic in other species, including some Bramocharax.
Polymorphisms.
Valdez-Moreno (2005) coded this same character (her character 49) differently, with state 0 as “semicircular”
vs. state 1 as “straight”, the latter a synapomorphy for all characid genera studied by her.
FIGURE 14. Dorsal process of quadrate: (a) distally expanded, process narrower (Astyanax mexicanus, UMMZ 169835); (b)
not expanded, process wider (A. aeneus, UMMZ 178568). Bars are 0.5 mm long.
SCHMITTER-SOTO
112
·
Zootaxa 4109 (2) © 2016 Magnolia Press
26. Arms of premaxilla
[0] dentigerous longer (Fig. 15a); [1] subequal (Fig. 15b)
Fifteen steps, ci=0.13, ri=0.00. State 1 could be interpreted as a synapomorphy for the Atlantic southern
Mexican-Guatemalan clade followed by a reversion to state 0 in most of its species, parallel in A. aeneus and
others.
FIGURE 15. Premaxilla: (a) dentigerous arm longer (Astyanax “Tamazulapan”, UMMZ 234194); (b) arms subequal (A.
aeneus, UMMZ 178568). Bars are 1 mm long.
27. First row of premaxillary teeth
[0] with nine teeth or more; [1] with three to five teeth (Fig. 15); [2] with two teeth
Four steps, ci=0.50, ri=0.33. State 1 is a synapomorphy for the studied Astyanax + Hyphessobrycon, some
species having 3–4, others 4–5, but most of them constantly with four teeth. State 2 is an homoplastic
synapomorphy for A. cf. fasciatus “das Velhas” + A. cocibolca, the latter being the sole Central American species
with this trait, convergent in A. orthodus, in A. cf. fasciatus “Ceará”, and in Roeboides.
Mirande (2010) coded this trait differently, as two binary characters: 129, with states 0, four or fewer teeth, and
1, five or more; and character 130, states 0, seven or fewer teeth, and 1, eight or more. He coincides with the classic
distinction by Eigenmann (1921) of genera with four teeth, such as Bryconamericus, and genera “with five teeth”,
such as Astyanax. I do not find such constancy in the latter genus.
28. Premaxillary teeth rows
[0] three; [1] two (Fig. 15)
One step, ci=1.0, ri=0.0. State 1 is a strict synapomorphy for the studied species of Astyanax +
Hyphessobrycon (also in Roeboides).
The character was used also by Valdez-Moreno (1997), same polarity found; nevertheless, in a subsequent
multigeneric cladogram (Valdez-Moreno 2005), having three rows of teeth on the premaxilla turned out to be a
synapomorphy for genus Brycon, the condition of having only two rows being plesiomorphic, which is also the
view of Zanata & Vari (2005) for the Characiformes.
This corresponds to Mirande’s (2010) binary character 123, with one or two rows as state 0, typical of most of
the Characidae, and three rows as state 1, a homoplastic synapomorphy for various clades, including the one
containing Brycon. He discusses potential problems in recognizing homology in this character.
Opercular series, characters 29–35
29. Dorsal half of opercle
[0] sides about parallel at mid-length (Fig. 16a); [1] sides divergent (Fig. 16b)
One step, ci=1.00, ri=0.00. State 1 is a strict autapomorphy for A. “Tehuacán”.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
113
PHYLOGENY OF ASTYANAX
FIGURE 16. Sides of upper half of opercle: (a) about parallel at mid-length (Astyanax “Macal”, UMMZ 178599); (b)
divergent at mid-length (A. “Tehuacán”, UMMZ 198853). Bars are 1 mm long.
30. Interopercle
[0] longer; [1] shorter
One step, ci=1.00, ri=0.00. State 1 is a strict synapomorphy for the studied species of Astyanax +
Hyphessobrycon, also in Roeboides.
Valdez-Moreno (1997) first proposed this character, with the same interpretation.
31. Posterior edge of interopercle
[0] with an angle (Fig. 17a); [1] convex, without any angle (Fig. 17b)
FIGURE 17. Interopercle, caudal edge: (a) with an angle (arrow) (Astyanax mexicanus, UMMZ 169835); (b) no such angle (A.
“Texas”, UMMZ 170107). Bars are 1 mm long.
SCHMITTER-SOTO
114
·
Zootaxa 4109 (2) © 2016 Magnolia Press
Fifteen steps, ci=0.13, ri=0.24. State 1, homoplastically synapomorphic for A. bimaculatus + A. orthodus, for
the clade including A. nicaraguensis, and for A. “Texas” + A. “Rioverde”; also in Hyphessobrycon and others.
Several polymorphisms.
32. Subopercular dorsorostral projection
[0] absent; [1] present
One step, ci=1.00, ri=0.00. State 1 is a strict synapomorphy for the studied species of Astyanax.
This is Valdez-Moreno’s (1997) character 14, same interpretation.
33. Preopercular ventral rim
[0] straight, at least anteriorly (Fig. 18a); [1] convex (Fig. 18b)
Five steps, ci=0.40, ri=0.00. State 1 appears autapomorphically in parallel in A. “Acatlán”, A. “Tamazulapan”,
and A. “Rioverde”; polymorphisms in A. “Costa Rica” and A. “Veracruz”.
Valdez-Moreno (1997) combined this trait and the following one into her character 12, with three states: 0,
ventral rim straight and anterodorsal edge concave; 1, ventral rim curved and anterodorsal edge convex; 2, ventral
rim curved and anterodorsal edge concave. Her state 1 apparently occurred only in one population of A. “Texas”.
FIGURE 18. Preopercle: (a) ventral rim nearly straight, anterodorsal edge straight-concave (Astyanax cf. fasciatus “Alto São
Francisco”, UMMZ 216281); (b) ventral rim convex (A. “Rioverde”, UMMZ 192510); (c) anterodorsal edge with a median
convexity, arrow (A. “Campeche”, UMMZ 143428). Bars are 1 mm long.
34. Preopercular anterodorsal edge
[0] straight-concave (Fig. 18a); [1] with a median convexity (Fig. 18c)
Thirteen steps, ci=0.15, ri=0.39. State 1, synapomorphic for the studied Astyanax; synapomorphic reversions
to state 0 at the larger clades for Atlantic southern Mexico-Guatemala and central-northern Mexico, with several
polymorphisms.
35. Preopercular canals
[0] two (Fig. 18); [1] one
One step, ci=1.0, ri=0.0. State 1 is a strict autapomorphy for A. cocibolca.
Axial skeleton, characters 36–37
36. Edge of epuric plate on last neural spine
[0] concave; [1] straight, not indented (Fig. 19a); [2] convex, not indented (Fig. 19b); [3] straight, indented (Fig.
19c); [4] roundish, indented (Fig. 19d)
Eighteen steps, ci=0.28, ri=0.13. State 1, synapomorphic for the studied Astyanax. State 2, synapomorphic for
A. nasutus + B. bransfordii and for the northern Mexican-Texan clade. State 3, convergently autapomorphic for A.
nicaraguensis, A. “Cubilhuitz”, and B. caballeroi. State 4, a strict autapomorphy for B. dorioni. Polymorphisms in
the Atlantic southern Mexican-Guatemalan clade.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
115
PHYLOGENY OF ASTYANAX
FIGURE 19. Edge of epuric plate on last neural spine (arrows): (a) straight, no indentation (Astyanax orthodus, UMMZ
160225); (b) convex, no indentation (A. “Texas”, UMMZ 170107); (c) straight, indented (A. “Rioverde”, UMMZ 192510); (d)
roundish, indented (A. “Quiché”, UMMZ 186378)
37. Total vertebrae
[0] 42; [1] 34; [2] 32 or 33; [3] 31
Eleven steps, ci=0.36, ri=0.13. State 1 is strictly autapomorphic for A. atratoensis; state 2, a synapomorphy for
genus Astyanax; state 3, a synapomorphy for subgenus Astyanax. Autapomorphic reversions to state 2 in A. cf.
fasciatus “Ceará”, A. “Quiché”, B. caballeroi, and A. nasutus. Astyanax “Costa Rica and A. nicaraguensis are
polymorphic for this trait.
Mirande (2010) coded the character with just two states: 0, 40 vertebrae or fewer, which includes my states 1-
3; and state 1, 41 vertebrae or more. He acknowledged that this coding is “rather subjective”; most of the characids
that he examined had 35-38 vertebrae.
Squamation, characters 38–42
38. Scales on lateral line
[0] 48 or more; [1] 43 or fewer
One step, ci=1.00, ri=1.00. State 1 is a strict synapomorphy for the studied species of Astyanax.
39. Scale rows between lateral line and pelvic fin origin
[0] mean 5 or 6; [1] mean 7 or 8; [2] mean 4
Six steps, ci=0.50, ri=0.00. State 1, homoplastically autapomorphic for A. atratoensis, A. “Macal”, and A.
“Costa Rica”; state 2, for A. cf. fasciatus “das Velhas” and A. orthodus. Astyanax panamensis is polymorphic for
the trait.
40. Predorsal scales
[0] mean 16; [1] mean 11–13; [2] mean 10
Six steps, ci=0.50, ri=0.40. State 1 is synapomorphic for the studied species of Astyanax. State 2 is a strict
autapomorphy for A. “Tamiahua”. Polymorphism in several South American species and also in A. “Macal”.
SCHMITTER-SOTO
116
·
Zootaxa 4109 (2) © 2016 Magnolia Press
41. Predorsal scale series
[0] incomplete, an unscaled space behind tip of supraoccipital process (Fig. 20a); [1] complete to caudal tip of
supraoccipital process (Fig. 20b)
The character was originally proposed by Eigenmann (1921) to separate his subgenera Poecilurichthys and
Astyanax. Two steps, ci=0.50, ri=0.67; state 1 is a synapomorphy for the subgenus Astyanax (parallel in
Hyphessobrycon), to the exclusion of species in Poecilurichthys (A. atratoensis, A. bimaculatus, A. orthodus); the
latter subgenus is not recovered as monophyletic.
Mirande (2010) found the opposite polarity for this character.
FIGURE 20. Predorsal scale series: (a) incomplete at nape, arrow (Astyanax bimaculatus, UMMZ 206863); (b) complete to
supraoccipital (A. aeneus, UMMZ 178568).
42. Scaly sheath at anal fin base
[0] with imbricated scales, long; [1] simple, long; [2] simple, short; [3] with imbricated scales, short
Eight steps, ci=0.50, ri=0.50. State 1, synapomorphic for genus Astyanax. State 2, synapomorphic for subgenus
Astyanax sensu Eigenmann (1921). State 3, a strict autapomorphy for B. dorioni. Autapomorphic reversions to
state 0 in A. panamensis, A. cocibolca, B. bransfordii, and A. altior. Astyanax “Costa Rica” is polymorphic for the
trait.
Fins, pectoral and pelvic girdles, characters 43–51
43. Anal-fin rays
[0] mean 33–36; [1] mean 28–30; [2] mean 25–27; [3] mean 21–24; [4] mean 19
Eighteen steps, ci=0.28, ri=0.24. State 1 is a strict synapomorphy for A. orthodus + A. bimaculatus. State 2 is
homoplastically synapomorphic for A. nasutus + B. bransfordii, autapomorphic for several species formerly in
Bramocharax and for A. aeneus, A. fasciatus “Alto São Francisco”, A. “Belize”, and Hyphessobrycon. State 3 is
synapomorphic for the subgenus Astyanax sensu Eigenmann (1921); state 4, a strict autapomorphy for A.
“Rioverde”. Several polymorphisms.
Mirande (2010) split this multistate character into four binary characters. He observes that “its phylogenetic
utility may be mostly restricted to the resolution of rather small clades.” In his view, a low rather than a high
number of rays is the plesiomorphic state, because most non-characid Characiformes have few anal-fin rays; his
character 286, state 1, 11 rays or more, is typical of most Characidae.
44. First branched anal-fin ray
[0] width about normal (Fig. 21a); [1] noticeably wider than the rest (Fig. 21b)
Eight steps, ci=0.25, ri=0.40. State 1 is homoplastically synapomorphic for A. bimaculatus + A. orthodus and
the two Nicaraguan clades; autapomorphic in most former Bramocharax species and A. “Tamazulapan”.
Polymorphism in A. “Costa Rica” and A. panamensis.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
117
PHYLOGENY OF ASTYANAX
FIGURE 21. First unbranched anal-fin ray: (a) about same width as the rest, arrow (Astyanax “Texas”, UMMZ 170107); (b)
noticeably wider than the rest (B. dorioni, UMMZ 193918).
45. Post-anal element
[0] shorter (Fig. 22a); [1] longer (Fig. 22b)
Four steps, ci=0.50, ri=0.00. State 1 is parallel for A. “Petén” and B. bransfordii. Polymorphic in A. “Belize”
and A. “Quiché”.
FIGURE 22. Post-anal element (arrow): (a) shorter (Astyanax “Texas”, UMMZ 170107); (b) longer (B. bransfordii, FMNH
5919).
46. Predorsal elements
[0] 11; [1] 4–6
One step, ci=1.00, ri=1.00. State 1 is a strict synapomorphy for the studied Astyanax. This is Mirande’s (2010)
character 281 (as “number of supraneurals”), opposite polarity found, with states 0 and 1 defined respectively by
him as seven or fewer vs. eight or more.
47. First dorsal pterygiophore, rostral edge
[0] curved (Fig. 23a); [1] with a spine (Fig. 23b)
Nine steps, ci=0.22, ri=0.30. State 1, synapomorphic for subgenus Astyanax sensu Eigenmann (1921), parallel
in A. bimaculatus. Synapomorphic reversal to state 0 for A. nicaraguensis + (A. cocibolca + A. cf. fasciatus “das
Velhas”). Polymorphism in A. mexicanus, A. “Campeche”, and A. “Bacalar”.
SCHMITTER-SOTO
118
·
Zootaxa 4109 (2) © 2016 Magnolia Press
FIGURE 23. First dorsal pterygiophore, rostral edge: (a) curved (Astyanax “Costa Rica”, UMMZ 159153); (b) angled (A.
“Rioverde”, UMMZ 192510).
48. Cleithrum, suture to coracoid
[0] single, deep, with two convexities (Fig. 24a); [1] 2–3 interdigitations (Fig. 24b); [2] 4–5 interdigitations (Fig.
24c); [3] single, triangular, broad (Fig. 24d); [4] 6 interdigitations (Fig. 24e)
Eleven steps, ci=0.46, ri=0.14. State 1, synapomorphic for Astyanax. State 2, homoplastically autapomorphic
for A. “Ocotal” and A. altior. State 3, strict autapomorphy for B. caballeroi; state 4, for A. “Rioverde”. A reversion
to state 0 is synapomorphic for A. cocibolca + A. cf. fasciatus “das Velhas”, autapomorphic for A. “Cubilhuitz”.
Polymorphisms in A. aeneus and other species.
FIGURE 24. Suture cleithrum-coracoid: (a) single, deep, with two convexities or angles (Astyanax “Quiché”, UMMZ
173731); (b) with 2–3 interdigitations (A. “Texas”, UMMZ 186469); (c) with 4–5 interdigitations (Oaxacan A. aeneus, UMMZ
184801); (d) single, triangular, broal (Honduran A. aeneus, UMMZ 144617); (e) six interdigitations (A. “Rioverde”, UMMZ
192510). Bars are 0.5 mm long, except in (c) and (e), 1 mm.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
119
PHYLOGENY OF ASTYANAX
49. Postcleithrum, caudal process
[0] concave-sided dorsally and ventrally (Fig. 25a); [1] concave-sided dorsally, almost straight ventrally (Fig. 25b);
[2] squarish (Fig. 25c); [3] concave-sided, with a dorsal angle (Fig. 25d); [4] straight-sided, with a ventral proximal
indentation (Fig. 25e)
Nineteen steps, ci=0.26, ri=0.18. State 1 is homoplastically synapomorphic for A. nicaraguensis + (A. cf.
fasciatus “das Velhas” + A. cocibolca); state 2, autapomorphic for A. cf. fasciatus “Ceará”, A. aeneus, A.
“Tamiahua”, B. dorioni, and B. bransfordii; state 3, synapomorphic for the Atlantic southern Mexican-Guatemalan
clade, with several polymorphisms. State 4 is a strict autapomorphy for A. mexicanus.
FIGURE 25. Postcleithrum, caudal process: (a) concave-sided (Brycon guatemalensis, UMMZ 190656); (b) concave-sided
dorsally, almost straight ventrally (Astyanax “Veracruz”, UMMZ 184761); (c) squarish (A. aeneus, UMMZ 178490); (d)
concave-sided, with a dorsal angle (A. “Quiché”, UMMZ 161739); (e) straight-sided, with a ventral proximal indentation (A.
mexicanus, UMMZ 169835).
50. Pelvic bone, proximal edge
[0] curved to irregular (Fig. 26a); [1] straight (Fig. 26b)
Three steps, ci=0.33, ri=0.33. State 1, synapomorphic for the A. nicaraguensis clade, autapomorphic for A.
“Quiché” and B. dorioni.
FIGURE 26. Pelvic bone, proximal edge: (a) curved (Astyanax aeneus, UMMZ 178568); (b) straight (A. “Quiché”, UMMZ
193886).
51. Caudal fin lobes
[0] subequal; [1] inferior longer
Three steps, ci=0.33, ri=0.0. State 1, parallel in A. “Macal”, A. panamensis, and A. cocibolca.
SCHMITTER-SOTO
120
·
Zootaxa 4109 (2) © 2016 Magnolia Press
Coloration, character 52
52. Caudal spot
[0] on peduncle and on rays; [1] not on rays, only on peduncle
Two steps, ci=0.50, ri=0.00. State 1 is a convergent autapomorphy for A. atratoensis and A. panamensis.
Phylogeny
Forty-three equally most parsimonious trees were recovered. The strict consensus tree is 454 steps long (CI=0.289,
RI=0.264, Fig. 27; matrix shown in Table 1). Synapomorphies are mentioned above for every character concerned.
The genus Astyanax is recovered as monophyletic, but, to test this further, many more genera need to be
included. Astyanax atratoensis is the sister group to the larger clade of Central American and Mexican Astyanax,
i.e., the subgenus Astyanax, monophyletic, including four South American species. On the contrary, the subgenus
Poecilurichthys is not recovered as monophyletic.
Five clades stand resolved out of a large polytomy, three of them including one Brazilian species. A purely
Nicaraguan (Great Lakes) clade is A. nasutus + B. bransfordii. A second, partly Nicaraguan, clade features the
widespread A. nicaraguensis sister to A. cocibolca plus the Brazilian form A. cf. fasciatus “das Velhas”. The
northern Brazilian form A. cf. fasciatus “Ceará” is in a same clade with A. panamensis.
The largest resolved clade, with nine species, summarized as “Atlantic southern Mexican-Guatemalan” (but
again including a South American species, in a derived position), shows the widespread Gulf of Mexico form A.
“Veracruz” sister to the rest, which includes an inland central Mexican microendemic, A. “ Acatlán”, in turn sister
to a clade with species from northeastern Guatemala, Belize, and southeastern Mexico.
Finally, a central-northern Mexican-Texan clade shows the northernmost species in the genus, A. “Texas”, in a
derived position.
Bramocharax species were not recovered together. Two of them occur in separate resolved clades; the other
three Bramocharax occur in the large polytomy corresponding to subgenus Astyanax, same as A. aeneus s. str.
(restricted to Pacific southern Mexico and northern Central America) and A. fasciatus s. str. (lower Rio São
Francisco, Brazil).
Discussion
Eleven out of the 30 cranial characters used by Valdez-Moreno (1997) were also included in the present study,
albeit many of them with different decisions on character states. Since hers was a population tree, the comparison
to the present hypothesis is not clear, except that northern forms (what I call Astyanax “Texas”) occur in a derived
position.
Twelve out of the 360 morphological characters used by Mirande (2010) were also used in this analysis; he
coded all additive characters as binary, which means that often one of my multistate character corresponds to two
or more of his characters. He found B. bransfordii to lie in its own clade (with species of Oligosarcus, one
undescribed), outside the Tetragonopterinae. No other authors have found Bramocharax so far away
phylogenetically from Astyanax, hypotheses ranging from finding the former genus polyphyletic, every one of its
component species being sister to a sympatric species of Astyanax (Ornelas-García et al. 2008), to recovering it as
monophyletic if some species are excluded, but making Astyanax paraphyletic if recognized (Valdez-Moreno
1997).
Mirande (2010) found his “A. mexicanus” (which corresponds to the form here called A. “Texas”) to be the
sister taxon to a clade including many South American Astyanax. The “back to South America” pattern, a recurrent
presence of South American species within primarily Middle American clades, has been observed in several
groups. For example, Schmitter-Soto (2007) found the South American Caquetaia nested within Middle American
cichlids, and Bermingham and Martin (1998) found Roeboides meeki from Colombia within Lower Central
American species of the genus. Chakrabarty and Albert (2011) reviewed several Neotropical fish phylogenies
suggesting that much of the Central American species diversity can be explained by older biogeographic events
Zootaxa 4109 (2) © 2016 Magnolia Press
·
121
PHYLOGENY OF ASTYANAX
FIGURE 27. Strict-consensus species cladogram for Middle American Astyanax and comparative material (361 steps, CI
0.363, RI 0.431). Undescribed species of Astyanax are designated by their region or locality. Numbers above lines are
apomorphies, with the superindex indicating the character state; for character numbering, see Character description and
analysis.
SCHMITTER-SOTO
122
·
Zootaxa 4109 (2) © 2016 Magnolia Press
TABLE 1. Matrix of characters and species used for parsimony analysis of character distribution in Central American
and Mexican Astyanax and Bramocharax. Question marks designate not only unknown states, but also polymorphisms
and inapplicable characters. See cladogram in Figure 27.
......continued on the next page
12345678910 11 12 13 14 15 16 17 18
Brycon 000000000000000000
Hyphessobrycon 00?002000000?01002
Roeboides 00?003010? 0 0 ? 0 0 ? 1 0
“Acatlán” 00010 202?1100?1112
aeneus ?000? ?0???000?0? 12
altior 0001? 102?1110?0111
“Campeche” 0 0 0 1 0 10211000?1111
“Texas” ?0??? 2024? 11111105
atratoensis 00000 1011000011111
“Bacalar” ? 0 0 1 0 10200100?11? 1
baileyi ????? 1????000?0011
“Belize” 10010 1020000??10??
bimaculatus 00000 10200 ???00111
bransfordii 101?? 00241001400?4
“Quiché” ? 0 0 1 ? 2 ? 2 ? 1 ? 0 ? ? 1 ? ? 1
caballeroi 00010 1020100110001
cf. fasciatus “Alto
São Francisco”
000?? 10201011111?1
cf. fasciatus “Ceará” 1 0 0 1 0 1010000011011
cocibolca 21100 30121002110?1
“Cubilhuitz” 0 ? ? 1 0 10010001110?1
cf. fasciatus “das
Velhas”
00100 1013111101003
dorioni 20010 1113010011011
fasciatus ????? 1????0001101?
“Veracruz” ?00?1 1010110031?11
“Macal” 0 ? ? 1 0 20241110011?1
mexicanus 20001 2010100011105
nasutus 201?? 2013100130???
nicaraguensis 10000 10?0100131012
“Ocotal” 00010 10??100001111
“Costa Rica” 0 1 1 1 ? 1 0 ? 0 ?????11?1
orthodus ?0000 10200 ???21112
panamensis 100?? 101?? 101?1?12
“Petén” 10000 10?01 ? 0 ? 1 1 1 ? 1
“Rioverde” 2 0 0 1 1 20201111?1101
“Tamazulapan” 00010 2000100141005
“Tamiahua” ? 00?0 201?011?101?1
“Tehuacán” 2??00 2220100130001
Zootaxa 4109 (2) © 2016 Magnolia Press
·
123
PHYLOGENY OF ASTYANAX
TABLE 1. (cont.)
......continued on the next page
19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35
Brycon 00000000000000000
Hyphessobrycon 1001201011011?000
Roeboides 0?0010002101?000?
“Acatlán” ?1?121011101 ?1100
aeneus ? ?11? 1111101 ?10?0
altior 1401211?1101 11000
“Campeche” 1 ? 0121??1101 ?10?0
“Texas” 1401?10?1101 11000
atratoensis 010120101101 0?010
“Bacalar” 1 ? ? 121??1101 11010
baileyi 1104???01101 0?0?0
“Belize” 1101?1? ?1101 ?1010
bimaculatus 120120101101 11010
bransfordii 040420001101 01010
“Quiché” 1 4 0121??1101 ?10?0
caballeroi 110421111101 01000
cf. fasciatus “Alto
São Francisco”
110111101101 01000
cf. fasciatus “Ceará” 1 1 0111102101 01010
cocibolca 110211112101 11011
“Cubilhuitz” 1 1 0121001101 01010
cf. fasciatus “das
Velhas”
030111102101 11010
dorioni 140422001101 01010
fasciatus 1?01?????1?? ??0??
“Veracruz” ?101?1111101 ? 1?00
“Macal” 1 4 0221101101 01010
mexicanus 1101210?1101 01000
nasutus ???3201??101 0?010
nicaraguensis 1?1311101101 11010
“Ocotal” 140421001101 0? ?00
“Costa Rica” 0 1 ? 1 2 ? 1 ? 1 1 0 1 ? 1 ? ? 0
orthodus 110120102101 11010
panamensis ??? 1101?1101 110?0
“Petén” 140121001101 010?0
“Rioverde” 0 4 0111001101 11100
“Tamazulapan” 111111001101 01110
“Tamiahua” 1??1?1111101 ?10?0
“Tehuacán” 01?111001111 01000
SCHMITTER-SOTO
124
·
Zootaxa 4109 (2) © 2016 Magnolia Press
TABLE 1 (cont.)
36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52
Brycon 00000000000000000
Hyphessobrycon ??0001020?0????0?
Roeboides ??000?000? 0????00
“Acatlán” ?3101123001113000
aeneus ? 31011220011?2000
altior 1310110300112?000
“Campeche” ? 310112?001? 1?000
“Texas” 231011230010? ?000
atratoensis 11111010001010001
“Bacalar” 1 310112?001???000
baileyi ??1011221?? ????00
“Belize” ?31011220? 111?000
bimaculatus 1210?011101110?00
bransfordii 23101102111??2?00
“Quiché” 1 21011230?11? ?100
caballeroi 32101122001133000
cf. fasciatus “Alto São
Francisco”
1310?122001113000
cf. fasciatus “Ceará” 3 3101123001112000
cocibolca 23101102101003?10
“Cubilhuitz” 4 3101123001103000
cf. fasciatus “das
Velhas”
1212?123101001100
dorioni 12101133101012100
fasciatus ??10?1?2001?????0
“Veracruz” ?310112?00111? 000
“Macal” 1 211?123001113010
mexicanus 23101123001?15000
nasutus 22101??2101??30?0
nicaraguensis 3?101122101011100
“Ocotal” 13101?2300? 123?00
“Costa Rica” ? ? 1111???010??000
orthodus 1212?011101011000
panamensis 131?110??0111? 011
“Petén” ? 310112301111? 000
“Rioverde” ? 3101124001143000
“Tamazulapan” 23101123101113000
“Tamiahua” 23102123001112000
“Tehuacán” 1310112300111?000
Zootaxa 4109 (2) © 2016 Magnolia Press
·
125
PHYLOGENY OF ASTYANAX
between Central and South America, and that the faunal interchange made possible by the rise of the isthmus led to
several Plio-Pleistocene reinvasions of Central American taxa back into northwestern South America. A Paleocene
first closure of the Isthmus was already proposed by Myers (1966), and geological evidence for this view exists
(e.g. Iturralde-Vinent 2006).
The occurrence of A. “Acatlán” in a relatively basal position within its clade is interesting, The new species
(Schmitter-Soto, unpubl. data) is endemic to central Mexico; its sister-group relationship to Guatemalan-
Southeastern Mexican species, in turn sister to a clade including a Brazilian species, bespeaks again a probable
“return to South America”.
In contrast, other highland, microendemic, undescribed species occur in derived positions within their clades.
These are A. “Macal”, from the Maya mountains of Belize, and A. “Ocotal”, from an endorheic lake in Chiapas
which harbors at least one other endemic, Rocio ocotal (Schmitter-Soto 2007).
Including cytogenetic data, available only for A. “Texas”, A. “Veracruz” (both 2n=50: Klinkhardt et al. 1995,
d’Artola-Barceló2009), and A. fasciatus (2n=46 or 48: Pazza et al. 2007), does not alter the topology of the tree.
Mirande (2010) coded lower chromosome numbers as plesiomorphic, but the prevailing view (e.g. Portela et al.
1988) is that this decrease in chromosome number is a derived trend.
Esquivel-Bobadilla’s (2011) unrooted mtDNA tree shows one clade for northern Mexico (= A. “Texas”),
another for cave forms (also in northern Mexico), and a third clade for central-southern forms, in the pattern B.
caballeroi + (A. “Veracruz” + A. “Campeche”), which is congruent with the present hypothesis. More extended in
material examined and choice of genetic markers is the phylogeny by Ornelas-García et al. (2008), based on
mtDNA (CO1, cyt b, 16 S) and nuclear DNA (RAG1). These authors found A. fasciatus s. str. to be the sister taxon
to all studied Middle American Astyanax, not part of the ingroup. Astyanax panamensis (as per my identification)
is in their cladogram the following sister species to the rest, which consists of two large clades: excluding new
species and localities not considered here (e.g., hypogean populations), the first group unites A. nasutus, A.
nicaraguensis + A.bransfordii, A. “Macal”, and A. “Belize”; a second, more northern, group includes A. “Quiché”,
A. aeneus, B. dorioni + A.”Petén”, A. altior, B. caballeroi, A. “Veracruz”, A. mexicanus, and A. “Texas”.
As Degnan and Rosenberg (2006) observed, “because of the stochastic way in which lineages sort out during
speciation, gene trees may differ in topology from each other and from species trees”. Nevertheless, the present
morphological cladogram does show interesting congruence with the molecular proposal by Ornelas-García et al.
(2008). Both cladograms find “Nicaraguan clades”, with A. nasutus and B. bransfordii as sister taxa, as well as a
derived position for the northernmost species. However, the placing of many species, such as A. “Macal” and A.
aeneus, is at variance between our hypotheses, even given the uncertainty caused by the large polytomy in the
present strict-consensus proposal.
As mentioned above, Mirande (2010) found A. “Texas” in the basal position of a South American Astyanax
clade; nevertheless, it is difficult to compare his hypothesis to the one here proposed, given that there are no other
taxa in common (except for B. bransfordii), since his aim was to recover the phylogenetic relationships of the
Characidae, not to test any lower-level relationships. Mirande (2010) found species of Hyphessobrycon,
Psellogramus, and Markiana in Astyanax-dominated clades; a broader sampling of Astyanax species and related
genera is the still-missing bridge between his revision and the present contribution.
Most species in this study are polymorphic for several characters; among the few taxa with no polymorphisms
are B. caballeroi, B. dorioni, and A. cf. fasciatus “Ceará”. However, invoking the pervading phenomenon to
preserve an “A. aeneus s. lat.”, at any level, does not seem useful: despite polymorphisms, all species in this study
are diagnosable under a phylogenetic species concept (unpubl. data.).
The fact that contact zones with genetic exchange are ubiquitous between species of Astyanax has been used
by some authors (Strecker et al. 2004, Hausdorf et al. 2011) to sustain the view that all of these forms belong in just
one species, A. fasciatus, under a strict biological species concept. However, these authors themselves (e.g.
Strecker 2005) have endorsed a different view concerning the species of Cyprinodon in Lake Chichancanab, many
of which not only share haplotypes, but are indistinguishable by most molecular markers and display a certain
percentage of hybrids in nature. In the laboratory, the females of C. beltrani prefer the males of the larger species
C. maya (Strecker 1996). Hybrid zones are also common among species of Cyprinidae in Europe (Machordom et
al. 1990).
The nominal A. emperador Eigenmann & Ogle 1907 and A. robustus Meek 1912 have been considered species
of Bryconamericus (Meek 1914, Grey 1947, Román-Valencia 2002). Hence they were not included in this study.
SCHMITTER-SOTO
126
·
Zootaxa 4109 (2) © 2016 Magnolia Press
Also omitted were García-Ornelas et al.’s (2008) “species 2” (Río Máquinas, Veracruz) and “species 3”
(Montebello, Pacific Chiapas), because of unavailability of material for osteological study. The equivalence of
Ornelas-García et al.’s (2008) “species 5” (Ciruelas to Chires, Panama), “species 6” (Ciruelas, Panama to
Tempisque, Costa Rica), and “species 8” (Chagres, Panama) is not clear, since material from their exact localities
was not available for study. The Panamanian area deserves further analysis, since several Astyanax species coexist
in the region, as is true also for the complex Guatemalan Petén.
My hypothesis does not recover a Bramocharax clade. Ornelas-García et al. (2008) also found Bramocharax
to be polyphyletic. Rosen (1972) proposed that this genus, understood as composed by B. baileyi, B. bransfordii,
and B. dorioni (the latter two, considered subspecies of B. bransfordii—found quite apart in the present phylogeny),
originated in the Usumacinta system, with B. baileyi as the most primitive species. Valdez-Moreno (2005) found
the same pattern, adding a new species from Chiapas (= B. “Ocotal”) as the most basal taxon. She claimed
Bramocharax to be monophyletic, but the inclusion of the other Middle American Astyanax in the analysis proves
otherwise.
Moreover, the clustering together of former Bramocharax in Valdez-Moreno’s (2005) cladogram may be due
to the large number of “trophic” characters used. These species did not appear together in one clade in the
molecular hypothesis of Ornelas-García et al. (2008), their phenotypic resemblance being attributed to adaptive
convergence, as every examined Bramocharax species was in a same clade with its respective sympatric Astyanax
species.
As for the grouping of B. bransfordii and Oligosarcus in Mirande’s (2010) hypothesis, the author himself
acknowledged that his material was not sufficient to solve the problem, and he suggested that “[t]he inclusion of
some Mesoamerican species of Astyanax with relatively high number of maxillary teeth, such as A. nasutus Meek
and some morphologically conservative species of Bramocharax, such as B. baileyi Rosen, would be useful to test
the monophyly and position of this clade” (interestingly, A. nasutus was found to be sister to B. bransfordii in the
present hypothesis). The sole synapomorphy for his Bramocharax clade is the form of the epioccipital bridge,
medially depressed. Valdez-Moreno (2005) included O. hepsetus in her study, and found it to lie in a group with
species of Acestrorhynchus, far outside her Bramocharax-Astyanax clade, in fact in a position basal respective to
Roeboides and Hyphessobrycon.
An ontogenetic character state shift for A. mexicanus in “trophic” traits has been documented, namely a
transition from unicuspid to multicuspid teeth (Trapani et al. 2005). Valdez-Moreno (1997) observed not just
variation in the number of cusps, but also a confounding effect of erosion in older individuals. I have made little
use of dental characters in this review, and none of cuspidization.
The use of quantitative characters in cladistics can be controversial (Pimentel & Riggins 1987). Nevertheless,
overlapping variability exists in “qualitative” characters as well (Kitching et al. 1998), and statistical coding
procedures help formulate objective decisions (Rae 1998). On the other hand, Mirande (2010) and other authors
have made substantial use of quantitative characters without any explicit criterion for defining states, aside from
coding as “polymorphic” any species with overlapping variability.
Ayache and Near (2009) state that morphological data are valuable not just for the discovery and description of
new species, but also for building phylogenetic hypotheses, notwithstanding the fact that resolution and robustness
of cladograms based on anatomical traits tend to be much lower than those using DNA-sequence data. In spite of
the shortcomings, morphological datasets may underscore problematic areas of phylogenies deserving further
study. Such is the present case.
Acknowledgments
This work was done while on sabbatical stay at the University of Michigan Museum of Zoology, Division of
Fishes, under a partial grant from the Mexican Consejo Nacional de Ciencia y Tecnología. I thank most warmly my
hosts, Bill Fink and Jerry Smith, as well as the collection manager, Doug Nelson, who handled loans from ANSP,
BMNH, ECOCH, FLMNH FMNH, GCRL, LACM, MNCN, MNHN, UANL, USM, USNM, ZMB, and ZMUC.
Humberto Bahena processed the photographs. Janneth Padilla prepared the maps and all figures.
Zootaxa 4109 (2) © 2016 Magnolia Press
·
127
PHYLOGENY OF ASTYANAX
References
Archie, J. (1989) Homoplasy excess ratios: new indices for measuring levels of homoplasy in phylogenetic systematics and a
critique of the ocnsistency index. Systematic Zoology, 38, 253–269.
http://dx.doi.org/10.2307/2992286
Ayache, N.C. & Near, T.J. (2009) The utility of morphological data in resolving phylogenetic relationships of darters as
exemplified with Etheostoma (Teleostei: Percidae). Bulletin of the Peabody Museum of Natural History, 50 (2), 327–346.
http://dx.doi.org/10.3374/014.050.0203
Bussing, W.A. (1998) Peces de las aguas continentales de Costa Rica. Revista de Biología Tropical, 46, 1–468.
Bermingham, E., & Martin, A.P. (1998) Comparative mtDNA phylogeography of neotropical freshwater fishes: testing shared
history to infer the evolutionary landscape of lower Central America. Molecular Ecology, 7 (4), 499–517.
http://dx.doi.org/10.1046/j.1365-294x.1998.00358.x
Chakrabarty, P. & Albert, J. (2011) Not so fast. A new take on the Great American Biotic Interchange. In: Albert, J. & Reis,
R.E. (Eds.), Historical Biogeography of Neotropical Freshwater Fishes. University of California Press, Berkeley, pp.
293–305.
http://dx.doi.org/10.1525/california/9780520268685.003.0018
Crisp, M.D. & Chandler, G.T. (1996) Paraphyletic species. Telopea, 6 (4), 813–844.
http://dx.doi.org/10.7751/telopea19963037
d’Artola Barceló, A.L. (2009) Aspectos citogenéticos de dos poblaciones de la sardina de agua dulce Astyanax aeneus (Pisces:
Characidae). Unpubl. B.Sc. Diss., Universidad Juárez Autónoma de Tabasco, Villahermosa, Mexico.
Degnan, J.H. & Rosenberg, N.A. (2006) Discordance of species trees with their most likely gene trees. PLoS Genetics, 2 (5),
762–768.
http://dx.doi.org/10.1371/journal.pgen.0020068
Eigenmann, C.H. (1921) The American Characidae. Part 3. Memoirs of the Museum of Comparative Zoology, 43, 209–310.
Esquivel-Bobadilla, S. (2011) Análisis genético de Astyanax mexicanus (Characidae, Teleostei, Pisces) de la vertiente
atlántica de México usando microsatélites. Unpubl. M.Sc. Diss., Centro de Investigaciones Biológicas del Noroeste, La
Paz, Mexico, 83 pp.
Farris, J.S. (1989) The retention index and the rescaled consistency index. Cladistics, 5, 417–419.
http://dx.doi.org/10.1111/j.1096-0031.1989.tb00573.x
Felsenstein, J. (2009) PHYLIP (Phylogeny Inference Package) version 3.69. Department of Genome Sciences, University of
Washington, Seattle.
Greenfield, D.W. & Thomerson, J.E. (1997) Fishes of the Continental Waters of Belize. University Press of Florida,
Gainesville, 311 pp.
Grey, M. (1947) Catalogue of type specimens of species in Chicago Natural History Museum. Fieldiana: Zoology, 32 (3),
109–205.
Hausdorf, B., Wilkens, H., & Strecker, U. (2011) Population genetic patterns revealed by microsatellite data challenge the
mitochondrial DNA based taxonomy of Astyanax in Mexico (Characidae, Teleostei). Molecular Phylogenetics and
Evolution, 60 (1), 89–97.
http://dx.doi.org/10.1016/j.ympev.2011.03.009
Iturralde-Vinent, M.A. (2006) Meso-Cenozoic Caribbean paleogeography: implications for the historical biogeography of the
region. International Geology Review, 48 (9), 791–827.
http://dx.doi.org/10.2747/0020-6814.48.9.791
Kitching, I.J., Forey, P.L., Humphries, C.J. & Williams, D.M. (1998) Cladistics. The Theory and Practice of Parsimony
Analysis. 2nd Ed. Oxford University Press, Oxford, 228 pp.
Klinkhardt, M., Tesche, M. & Greven, H. (1995) Database of fish chromosomes. Westarb Wissenschaften, Heidelberg, 237 pp.
Machordom, A., Berrebi, P., & Doadrio, I. (1990) Spanish barbel hybridization detected using enzymatic markers: Barbus
meridionalis Risso × Barbus haasi Mertens (Osteichthyes, Cyprinidae). Aquatic Living Resources, 3 (04), 295–303.
http://dx.doi.org/10.1051/alr:1990030
Maddison, W.P. & Maddison, D.R. (2011) Mesquite: A modular system for evolutionary analysis, version 2.75. Available from:
http://mesquiteproject.org (accessed 27 November 2012)
Meek, S.E. (1914) An annotated list of fishes known to occur in the fresh-waters of Costa Rica. Field Museum of Natural
History Publications, Zoölogical Series, 10 (10), 101–134.
http://dx.doi.org/10.5962/bhl.title.5576
Melo, F.A.G. de (2005) Revisão taxonômica do complexo de espécies Astyanax fasciatus (Cuvier, 1819) (Teleostei:
Characiformes: Characidae). Ph.D. Diss., Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil.
Miller, R.R. (1986) Composition and derivation of the freshwater fish fauna of Mexico. Anales de la Escuela Nacional de
Ciencias Biológicas, Instituto Politécnico Nacional, 30, 121–153.
Miller, R.R., Minckley, W.L. & Norris, S.M. (2009) Peces Dulceacuícolas de México. Comisiòn Nacional para el
Conocimiento y Uso de la Biodiversidad, ECOSUR, Sociedad Ictiológica Mexicana, and Desert Fishes Council, Mexico
City, 559 pp.
Mirande, J.M. (2010) Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy.
SCHMITTER-SOTO
128
·
Zootaxa 4109 (2) © 2016 Magnolia Press
Neotropical Ichthyology, 8, 385–568.
http://dx.doi.org/10.1590/S1679-62252010000300001
Myers, G.S. (1966) Derivation of the freshwater fish fauna of Central America. Copeia, 1966, 766–773.
http://dx.doi.org/10.2307/1441405
Ornelas-García, C.P., Domínguez-Domínguez, O. & Doadrio, I. (2008) Evolutionary history of the fish genus Astyanax Baird
& Girard (1854) (Actinopterygii, Characidae) in Mesoamerica reveals multiple morphological homoplasies. BMC
Evolutionary Biology, 8, 340.
http://dx.doi.org/10.1186/1471-2148-8-340
Pazza, R., Kavalco, K.F. & Bertollo, L.A.C. (2007) Chromosome polymorphism in Astyanax fasciatus (Teleostei, Characidae).
1. Karyotype analysis, Ag-NORs and mapping of the 18S and 5S ribosomal genes in sympatric karyotypes and their
possible hybrid forms. Cytogenetic and Genomic Research, 112, 313–319.
Pimentel, R.A. & Riggins, R. (1987) The nature of cladistic data. Cladistics, 3, 201–209.
http://dx.doi.org/10.1111/j.1096-0031.1987.tb00508.x
Portela, A.L.B.S., Galetti Jr., P.M. & Bertollo, L.A.C. (1988) Considerations on the chromosome evolution of
Tetragonopterinae (Pisces: Characidae). Revista Brasileira de Genética, 11 (2), 307–316.
Rae, T.C. (1998) The logical basis for the use of continuous characters in phylogenetic systematics. Cladistics, 14, 221–228.
http://dx.doi.org/10.1111/j.1096-0031.1998.tb00335.x
Román-Valencia, C. (2002) A systematic review of the species of the genus Bryconamericus (Teleostei: Characidae) from
Central America. Revista de Biología Tropical, 50 (1), 173–192.
Rosen, D.E. (1972) Origin of the characid fish genus Bramocharax and a description of a second, more primitive, species in
Guatemala. American Museum Novitates, 2500, 1–21.
Sabaj Pérez, M.H. (Ed.) (2012) Standard symbolic codes for institutional resource collections in herpetology and ichthyology:
an online reference. Vers. 3.0. Available from: http://www.asih.org/ (accessed 20 April 2014)
Schmitter-Soto, J.J. (2007) Phylogeny of species formerly assigned to the genus Archocentrus (Perciformes: Cichlidae).
Zootaxa 1618, 1–50.
Schmitter-Soto, J.J., Valdez-Moreno, M.E. Rodiles-Hernández, R.& González-Díaz, A.A. (2008) Astyanax armandoi, a junior
synony m of Astyanax aeneus (Teleostei: Characidae). Copeia, 2008 (2), 409–413.
http://dx.doi.org/10.1643/CI-07-012
Simon, C. (1983) A new coding procedure for morphometric data with an example from periodical cicada wing veins. In:
Felsenstein, J. (Ed.), Numerical Taxonomy. Springer, Berlin, 378–383.
http://dx.doi.org/10.1007/978-3-642-69024-2_40
Strecker, U. (1996) Molekulargenetische und ethologische Untersuchungen zur Speziation eines Artenschwarmes der Gattung
Cyprinodon (Cyprinodontidae, Teleostei). Ph.D. Diss., Univ. Hamburg. Shaker Verlag, Hamburg.
Strecker, U. (2005) Description of a new species from Laguna Chichancanab, Yucatan, Mexico: Cyprinodon suavium (Pisces:
Cyprinodontidae). Hydrobiologia 541, 107–115.
http://dx.doi.org/10.1007/s10750-004-4821-3
Strecker, U., Faúndez, V.H. & Wilkens, H. (2004) Phylogeography of surface and cave Astyanax (Teleostei) from Central and
North America based on cytochrome b sequence data. Molecular Phylogenetics and Evolution, 33 (2), 469–481.
http://dx.doi.org/10.1016/j.ympev.2004.07.001
Taylor, W.R. & van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone
and cartilage study. Cybium, 9, 107–119.
Trapani, J., Yamamoto, Y. & Stock, D.W. (2005) Ontogenetic transition from unicuspid to multicuspid oral dentition in a teleost
fish: Astyanax mexicanus, the Mexican tetra (Ostariophysi: Characidae). Zoological Journal of the Linnean Society, 145
(4), 523–538.
http://dx.doi.org/10.1111/j.1096-3642.2005.00193.x
Valdez-Moreno, M.E. (1997) Estudio comparativo osteológico del género Astyanax en diversas cuencas de México. Unpubl.
M.Sc. Diss., Universidad Nacional Autónoma de México, Mexico City.
Valdez-Moreno, M.E. (2005) Osteología craneal del subgénero Catemaco género Bramocharax (Teleostei: Characidae):
relaciones filogenéticas y biogeografía. Unpubl. Ph.D. Diss., Universidad Autónoma de Nuevo León, Monterrey, Mexico.
Zanata, A.M. & Vari, R.P. (2005) The family Alestidae (Ostariophysi, Characiformes): a phylogenetic hypothesis of a trans-
Atlantic clade. Zoological Journal of the Linnean Society, 145 (1), 1–144.
http://dx.doi.org/10.1111/j.1096-3642.2005.00183.x
Zootaxa 4109 (2) © 2016 Magnolia Press
·
129
PHYLOGENY OF ASTYANAX
APPENDIX 1. Material examined (in bold, with cleared-and-stained or skeletonised specimens). Order by country and
locality, then collection codon and catalog number. Institutional codons follow Sabaj-Pérez (2012). Further data
available from the author on request.
Astyanax “Acatlán”: UMMZ 203317, Mexico, Río Acatlán; UMMZ 191698, Mexico, Río Jía.
Astyanax “Bacalar”: ECOCH 1764, UMMZ 196478, 246454, Mexico, Laguna Bacalar; UMMZ 202874, Belize, Gabourel
Creek; UMMZ 246840, Belize, River Sittie; ECOCH 1976, Mexico, Laguna Encantada; UMMZ 196540, Mexico, pool
near Tulum; UMMZ 210875, Mexico, Río Hondo; UMMZ 194103, Guatemala, Río Mopán; FLMNH 101171, Mexico,
Tulum.
Astyanax “Belize”: MNHN 5224-5225 (ST of Tetragonopterus belizianus), Belize, Mullins River; UMMZ 246831, Belize,
Moho River; BMNH 1861.8.12.20-21 (ST of T. brevimanus), MRAC 7057 (ST of T. brevimanus), ZMB 6801 (ST of A.
panamensis), Guatemala, Lake Izabal; UMMZ 194018, Guatemala, Motagua; UMMZ 194002, Guatemala, Río Agua
Fría; UF 115561, Guatemala, Río Dulce; UMMZ 197303, Guatemala, Río Trincheras; USM 34095, Honduras, Aguán;
USM 35490, Honduras, Cangrejal; USM 36032, 36056, 36066, Honduras, Chamelecón; UF, Honduras, Colón; USM
31071, Honduras, La Ceiba; USM 35611, Honduras, Lancetilla; UMMZ 155869, Honduras, Río Celán; LACM 32458-1,
USM 36370, 36408, Honduras, Río Coco; UMMZ 228667, Honduras, Río Jutiapa; UMMZ 199524, Honduras, Río
Patuca; USM 34131, 36057, Honduras, Río Sico; UMMZ 155868, Honduras, Río Ulúa; USM 31910, Honduras, Tulián.
Astyanax “Campeche”: BMNH ex 1857.7.31.9 (syntypes of T. angustifrons), “Mexico”; UMMZ 143326, Guatemala, Arroyo
Subín; UMMZ 143428, Guatemala, Río San Pedro; UMMZ 102209, UMMZ 196571, Mexico, Río Champotón; UMMZ
196625, Mexico, Río Mamantel; UMMZ 246448, Mexico, Río Ulumán; UMMZ 64475, Mexico, aguada at Tuxpeña;
UMMZ 187429, Mexico, Zohlaguna.
Astyanax “Costa Rica”: ZMUC 948, ZMB 9197, ZMUC 947, ZMUC 955, ZMUC 956 (ST of T. orstedii), Costa Rica/
Nicaragua, Río San Juan; UMMZ 243884, Costa Rica, Corobicí; LACM 8311, Costa Rica, Divia; FMNH 6257 (HT of A.
regani), GCRL 5095, Costa Rica, Las Cañas; LACM 4831, UMMZ 194212, Costa Rica, Puntarenas; UMMZ 245906,
Costa Rica, Quebrada Blanca; UMMZ 245887, Costa Rica, Río Blanco; UMMZ 138248, Costa Rica, Siquirres; UMMZ
194221, Costa Rica, Tárraba; UMMZ 159153, 243890, Costa Rica, Tempisque; UMMZ 145677, Panama, Chiriquí.
Astyanax “Cubilhuitz”: UMMZ 188007, Guatemala, Río Dolores.
Astyanax “Macal”: UMMZ 178599, Belize, River Macal.
Astyanax “Ocotal”: UMMZ 171139, Mexico, Laguna Ocotal.
Astyanax “Petén”: BMNH 1864.1.26.374 (ST), Guatemala, Lake Petén Itzá ; UMMZ 190975, Guatemala, Arroyo El Chorro;
UMMZ 143332, Guatemala, Arroyo Jolomax; UMMZ 143441, Guatemala, Eckibix; UMMZ 143433, Guatemala, Lago
Petén Itzá; UMMZ 143424, Guatemala, Laguna Perdida; UMMZ 97877, Guatemala, Uaxactún; UMMZ 186378, Mexico,
Río Chiapa.
Astyanax “Quiché”: BMNH 1864.1.26.388, 1864.1.24.177 (ST of T. brevimanus), Guatemala, Río San Gerónimo; MNHN
5219 (ST of T. cobanensis, 6 dig), Guatemala, Cobán; LACM 37765-1, Guatemala, Ixcán; UMMZ 131142, Guatemala,
Río Copón; UMMZ 193886, Guatemala, Río Sachichá; UMMZ 173731, Mexico, Cintalapa; UMMZ 167714, Mexico, Río
Grande de Comitán; UMMZ 161769, Mexico, Río Santa Cruz.
Astyanax “Rioverde”: UMMZ 192510, 172194, Mexico, “Rioverde”; UMMZ 193447, Mexico, Río Santa María.
Astyanax “Tamazulapan”: TNHC 25027, TU 185676, UANL 14306, 14327, 22296, UMMZ 234194, USNM 357728, Mexico,
Ojo de Agua de Tamazulapan.
Astyanax “Tamiahua”: UMMZ 97362, Mexico, Río Cucharas; UMMZ 167489, Mexico, Río Nautla.
Astyanax “Tehuacán”: UMMZ 198853, Mexico, Río Salado.
Astyanax “Texas”: BMNH 1883.12.14.107 (ST of A. argentatus), USA, Río Nueces; UMMZ 182081, Mexico, Cuatro
Ciénegas; UMMZ 186469, Mexico, Río Apodaca; UMMZ 192476 (161 spms., 8 dig), Mexico, Río Limón; UMMZ
179171, Mexico, Río Salado; UMMZ 211059, Mexico, Durango; UMMZ 170107, USA, Rio Pecos.
Astyanax “Veracruz”: MNHN 5223 (ST of T. finitimus), Mexico, near Orizaba; UMMZ 183900, Mexico, Cocolapa; UMMZ
215480 (PT of A. armandoi), Mexico, Gregorio Sánchez Magaña; UMMZ 196382, Mexico, laguna near Tierra Blanca ;
UMMZ 191727, Mexico, near Palenque; UMMZ 97335, Mexico, Río Chachalacas; ANSP 15598-608, 32271, 32272 (ST
of T. streetsii), Mexico, Río Coatzacoalcos; UMMZ 184761, Mexico, Río Jaltepec; UMMZ 97336, Mexico, Río Paso San
Juan; UMMZ 184703, Mexico, Río Teapa; BMNH 1905.12.6.20, USNM 127094 (ST of T. macrophthalmus), Mexico,
Río Tonto.
Astyanax aeneus: BMNH 1907.4.10.3, BMNH 1860.6.17.41-42, Mexico, Oaxaca; UMMZ 197102, Guatemala, creek near
Taxisco; UMMZ 190523, Guatemala, ditches near Escuintla; BMNH 1865.4.29.43-44 (syntypes of T. microphthalmus),
BMNH 1865.4.29.45-50 (synty pes of T. humilis), Guatemala, Lake Amatitlán; UF 115560, UF 115933, Guatemala,
Escuintla; USM 36089, Honduras, Goascorán; UMMZ 144614, USM 33957, Honduras, Río Nacaome; UMMZ 144617,
UMMZ 144618, UMMZ 144621, Honduras, Río Choluteca; UMMZ 191719, Mexico, Huixtla; UMMZ 161510, México,
pond in Tehuantepec; UMMZ 168919, Mexico, Río Cacaluta; UMMZ 108596, UMMZ 108597, UMMZ 178490, UMMZ
181829, México, Río Papagayo; UMMZ 184737, Mexico, Río Tapanatepec; UMMZ 178568, Mexico, Río Tequisistlán;
UMMZ 184801, Mexico, Río Arenas.
Astyanax altior: UMMZ 102144 (holotype), UMMZ 102145 (paratypes), Mexico, Progreso; ECOCH 2988, Mexico, cenote
near Celestún; ECOCH 2955, Mexico, cenote near Chunchucmil; UMMZ 196550, Mexico, cenote at Dzibilchaltún;
SCHMITTER-SOTO
130
·
Zootaxa 4109 (2) © 2016 Magnolia Press
UMMZ 196564, Mexico, cenote near Sisal.
Astyanax atratoensis: UMMZ 61965, 160232, Colombia, Truando.
Astyanax bimaculatus: UMMZ 206863, Paraguay, Arroyo Tobatiry.
Astyanax cf. fasciatus “Alto São Francisco”: UMMZ 216281, Brazil, upper Río São Francisco.
Astyanax cf. fasciatus “Ceará”: UMMZ 147331, Brazil, Riacho do Constantino
Astyanax cf. fasciatus “das Velhas”: UMMZ 216372 (117 spms., 1 dig), Brazil, Río das Velhas.
Astyanax cocibolca: LACM 56648-1 (HT), 56648 (PT), UMIM 3601, UMMZ 209838, Nicaragua, Lake Nicaragua.
Astyanax fasciatus: MNHN 8653 (ST), MNHN 9896 (ST), Brazil, Río São Francisco.
Astyanax mexicanus: ZMUC 941, MZUT 149, ZMUC 942 (ST), “lake near Mexico”; UMMZ 169835, Mexico, Barranco
Capire; MNHN 5194 (ST of T. fulgens), MNHN 5191 (ST of T. nitidus), Mexico, near Cuernavaca; UMMZ 178410,
Mexico, Río Aguililla; UMSNH uncat., Mexico, Río Cupatitzio; UMMZ 160764, Mexico, Colima.
Astyanax nasutus: FMNH 5909 (HT), 5908 (PT), Nicaragua, Lake Managua.
Astyanax nicaraguensis: USNM 55653 (HT), USNM 55657 (PT), “Nicaragua”; UMMZ 166469, Costa Rica, Laguna del
Misterio; UMMZ 162475 (PT of A. aeneus var. costaricensis), Costa Rica, Río Parismina; USNM 45381, Nicaragua,
Greytown; USNM 44322, Nicaragua, Río Escondido; UMMZ 165772, Nicaragua, Matagalpa; USNM 44192, Nicaragua,
San Carlos; UMMZ 145686, Panama, Río Sixaola.
Astyanax orthodus: UMMZ 160225, Colombia, Truando.
Astyanax panamensis: BMNH 1864.1.26.415 (ST), “Pacific Panama”; GCRL 13409, Panama, Calobre ; FLMNH 16696,
Panama, Canal Zone; LACM 56197-2, Panama, Cárdenas; UF 12973, Panama, Chepo; UMMZ 61263, Panama, San Blas;
UF 19666, Panama, Tonosí.
Bramocharax baileyi: AMNH 30197 (HT), Guatemala, Río San Simón; UMMZ 190763, Guatemala, near Río Canilla.
Bramocharax bransfordii: USNM 16885 (ST), FMNH 5922 (HT of B. elongatus), UMMZ 180608, 16885 (PT of B.
elongatus), Nicaragua, Lake Nicaragua.
Bramocharax caballeroi: UANL 5681 (HT), UMMZ 184539, Mexico, Lake Catemaco.
Bramocharax dorioni: AMNH 29411 (HT), Guatemala, Río Semococh; UMMZ 187944, Guatemala, Río de la Pasión;
UMMZ 193918, Guatemala, Río San Simón..
Brycon guatemalensis: UMMZ ex 246840, Belize, Sittie; UMMZ 190656, Guatemala, Livingston.
Hyphessobrycon compressus: ECOCH 1449, Mexico, Arroyo Huay Pix; UANL 5784, 5849, after Valdez-Moreno (2005).
Roeboides guatemalensis: UANL 1696, after Valdez-Moreno (2005).
