ID Cichlidae

ZOOTAXA
1603
A systematic revision of the genus Archocentrus (Perciformes: Cichlidae),

with the description of two new genera and six new species
JUAN J. SCHMITTER-SOTO
Magnolia Press
Auckland, New Zealand
A systematic revision of the genus Archocentrus (Perciformes: Cichlidae), with the description
of two new genera and six new species
(Zootaxa 1603)
78 pp.; 30 cm.
28 Sept. 2007
ISBN 978-1-86977-159-1 (paperback)
ISBN 978-1-86977-160-7 (Online edition)
FIRST PUBLISHED IN 2007 BY

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ISSN 1175-5326 (Print edition)

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Zootaxa 1603: 1–78 (2007) ISSN 1175-5326 (print edition)
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Copyright © 2007 · Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
A systematic revision of the genus Archocentrus (Perciformes: Cichlidae),

with the description of two new genera and six new species
El Colegio de la Frontera Sur (ECOSUR), A.P. 424, MX-77000 Chetumal, QR, Mexico
jschmitt@ecosur.mx
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Key to Archocentrus and allied genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Systematic accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Genus Archocentrus Gill . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Archocentrus centrarchus (Gill, 1877) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Archocentrus multispinosus (Günther, 1867) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Archocentrus spinosissimus (Vaillant & Pellegrin, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Genus Cryptoheros Allgayer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Panamius, n. subgen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Cryptoheros panamensis (Meek & Hildebrand, 1913), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Subgenus Cryptoheros Allgayer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Cryptoheros spilurus (Günther, 1862). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Cryptoheros chetumalensis, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Cryptoheros cutteri (Fowler, 1932), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Bussingius, n. subgen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Cryptoheros septemfasciatus (Regan, 1908) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Cryptoheros altoflavus Allgayer, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Cryptoheros myrnae (Loiselle, 1997) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Cryptoheros nanoluteus (Allgayer, 1994) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Cryptoheros sajica (Bussing, 1974) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Amatitlania, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Amatitlania nigrofasciata (Günther, 1867), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Amatitlania coatepeque, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Amatitlania kanna, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Amatitlania siquia, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Rocio, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Rocio octofasciata (Regan, 1903), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Rocio ocotal, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Rocio gemmata Contreras-Balderas & Schmitter-Soto, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Genus Hypsophrys Agassiz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Hypsophrys nicaraguensis (Günther, 1859) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Hypsophrys nematopus (Günther, 1867), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Accepted by L. Page: 22 Jun. 2007; published: 28 Sept. 2007 3

Abstract
The cichlid genus Archocentrus has been considered one of the most promising (i.e., possibly natural) genera resurrected
to receive some of the species formerly included in Cichlasoma. Evidence is presented to justify generic recognition of
Archocentrus, as well as eight other closely related genera (Caquetaia, Hypsophrys, Parachromis, Amphilophus, Archo-
centrus, Cryptoheros, Amatitlania, and Rocio). Of these, Amatitlania (type species, A. nigrofasciata) and Rocio (type
species, R. octofasciata) are described as new. The present revision treats all nominal species ever assigned to Archocen-
trus, as well as species that have been included in or near the same clade as Archocentrus centrarchus (type species of the
genus) in available phylogenetic analyses. Geographical variation in morphology of the more widespread species was
examined, which has resulted in the description of six new species (Cryptoheros chetumalensis, Amatitlania coatepeque,
A. kanna, A. siquia, Rocio gemmata, and R. ocotal) with a seventh resurrected from synonymy (Cryptoheros cutteri).
Archocentrus includes the type species (Ar. centrarchus), plus Ar. spinosissimus and Ar. multispinosus. Cryptoheros is
restricted to the species complexes of Cr. spilurus (= subgenus Cryptoheros, including also Cr. chetumalensis and Cr.
cutteri) and Cr. septemfasciatus (= Bussingius n. subgen., including also Cryptoheros altoflavus, Cr. nanoluteus, Cr. myr-
nae, and Cr. sajica); Cryptoheros panamensis is placed in Panamius n. subgen. Herotilapia is synonymized with Archo-
centrus, and Neetroplus is synonymized with Hypsophrys, which now includes the type species H. nicaraguensis and H.
nematopus. Lectotypes are designated for Amatitlania nigrofasciata, Archocentrus spinosissimus, Cryptoheros septem-
fasciatus, Cr. spilurus, and Rocio octofasciata. Cichlasoma immaculatum is considered to be a synonym of Archocentrus
spilurus, not of Ar. spinosissimus.
Key words: taxonomy, cichlids, Middle America
Resumen
Archocentrus (Cichlidae) se ha considerado uno de los géneros más prometedores (es decir, posiblemente natural) resuci-
tado para recibir algunas de las especies antes asignadas a Cichlasoma. Se presenta evidencia que justifica el recono-
cimiento a nivel de género de Archocentrus, así como otros ocho géneros cercanamente relacionados (Caquetaia,
Hypsophrys, Parachromis, Amphilophus, Archocentrus, Cryptoheros, Amatitlania y Rocio). De éstos, Amatitlania (espe-
cie tipo, A. nigrofasciata) y Rocio (especie tipo, R. octofasciata) se describen como nuevos. La presente revisión trata
con todas las especies nominales alguna vez asignadas al género, así como las especies que han sido incluidas dentro o
cerca del mismo clado que Archocentrus centrarchus (especie tipo del género) en los análisis filogenéticos disponibles.
Se examinó la variación geográfica en morfología de las especies de distribución más amplia, lo cual resultó en la
descripción de seis especies nuevas (Cryptoheros chetumalensis, Amatitlania coatepeque, A. kanna, A. siquia, Rocio
gemmata y R. ocotal) y una séptima rescatada de la sinonimia (Cryptoheros cutteri). Archocentrus incluye a la especie
tipo (Ar. centrarchus), junto con Ar. spinosissimus y Ar. multispinosus. Cryptoheros se restringe a los complejos de Cr.
spilurus (= subgénero Cryptoheros, incluyendo también a Cr. chetumalensis y Cr. cutteri) y de Cr. septemfasciatus (=
Bussingius n. subgen., incluyendo también a Cr. altoflavus, Cr. nanoluteus, Cr. myrnae y Cr. sajica); Cr. panamensis se
coloca en Panamius n. subgen. Herotilapia se sinonimiza con Archocentrus, y Neetroplus se sinonimiza con Hypsophrys,
el cual ahora incluye a la especie tipo H. nicaraguensis y a H. nematopus. Se designan lectotipos para Amatitlania nigro-
fasciata, Archocentrus spinosissimus, Cryptoheros septemfasciatus, Cr. spilurus y Rocio octofasciata. Cichlasoma
immaculatum se considera sinónimo de Ar. spilurus, no de Ar. spinosissimus.
The arrangement of the American cichlids into genera is a puzzle of great difficulty.
—Hubbs, 1936: 254
Introduction
The cichlid fish genus Archocentrus was originally proposed as a subgenus of Heros Heckel by Gill (in Gill &
Bransford 1877), and later treated as an informal “section” of Cichlasoma Swainson in Regan’s (1905) revi-
sion of the genus. The generic status of Cichlasoma sensu Regan (1905), encompassing more than 100 spe-
cies, has been in a state of uncertainty since Kullander’s (1983) restriction of Cichlasoma sensu stricto to 12
South American species. Of the many remaining groups of species formerly referred to Cichlasoma, those

referred to Archocentrus have been among the strongest candidates for recognition as a monophyletic group
(e.g. Miller 1993, Miller et al. 2005). Species assigned to Archocentrus are major components of the neotropi-
cal Central American ichthyofauna from southern Mexico to Panama, occurring across all the ichthyolimno-
logical provinces of Bussing (1976), but the monophyly of the genus has not been explicitly demonstrated and
its species composition has been far from clear.
Since a comprehensive analysis of Cichlasoma sensu lato appeared unlikely, Kullander (1996) advocated
the provisional use of available generic names, a procedure that some workers had already adopted. Kullander
himself (1983), Stiassny (1991), and Miller (1993) had earlier recommended that the use of ‘Cichlasoma’—
with the quotation marks indicating its informality— would prevent further confusion. Because the systemat-
ics of all species previously referred to Cichlasoma had not yet been completely resolved, Nelson et al. (2004:
151–153) chose the conservative approach and tentatively referred all species to Cichlasoma in the most
recent checklist of fishes for the United States, Canada, and Mexico.
Several species have been assigned to Archocentrus more or less consistently in the literature (Table 1).
Allgayer (2001) restricted Archocentrus to include the two largest species (Ar. centrarchus and Ar. spinosissi-
mus), and referred six others to a new genus, Cryptoheros (type species, Cr. spilurus), which also included the
new species Cr. altoflavus. Other species that have sometimes been assigned to Archocentrus (Table 1)
include: Heros octofasciatus, though more frequently placed in incertae sedis; H. citrinellus, albeit demon-
strated by Roe et al. (1997) to be an Amphilophus; H. multispinosus, generally considered in the monotypic
genus Herotilapia; and Neetroplus panamensis, variously included in Archocentrus (in Kullander 2003),
Theraps (in Eschmeyer 2001), or Hypsophrys (in Bussing 1998).
TABLE 1. Valid and nominal species once or currently included in Archocentrus. Authorship, year of description,
orthography, and present status or name, after Eschmeyer 2005, except for changes proposed or supported in this work.
Basonym Author and year Included in Archo- Present status or name

centrus by:
Heros spilurus Günther, 1862 Regan 1908 Cryptoheros spilurus
Heros citrinellus Günther, 1864 Farias et al. 2000 Amphilophus citrinellus
Heros basilaris Gill, 1877 Synonym of A. citrinellus
Heros multispinosus Günther, 1867 Regan 1908 Archocentrus multispinosus
Heros nigrofasciatus Günther, 1867 Regan 1908 Amatitlania nigrofasciata
Cichlasoma (Archocentrus) centrarchum Gill, 1877 Original author Archocentrus centrarchus
Heros spinosissimus Vaillant & Pellegrin, 1902 Regan 1908 Archocentrus spinosissimus
Heros octofasciatus Regan, 1903 Regan 1908 Rocio octofasciata
Cichlasoma hedricki Meek, 1904 Synonym of R. octofasciata
Cichlasoma spinosissimum Pellegrin, 1904 Regan 1908 Synonym of Cr. spilurus
var. immaculatum
Cichlasoma granadense Meek, 1907 Synonym of Am. citrinellus
Cichlasoma septemfasciatum Regan, 1908 Loiselle 1997 Cryptoheros septemfasciatus
Cichlosoma biocellatum Regan, 1909 Synonym of R. octofasciata
Neetroplus panamensis Meek & Hildebrand, 1913 Kullander 2003 Cryptoheros panamensis
Cichlasoma cutteri Fowler, 1932 Cryptoheros cutteri
Cichlasoma sajica Bussing, 1974 Bussing 1998 Cryptoheros sajica
Archocentrus nanoluteus Allgayer, 1994 Original author Cryptoheros nanoluteus
Archocentrus myrnae Loiselle, 1997 Original author Cryptoheros myrnae
Cryptoheros altoflavus Allgayer, 2001 Kullander 2003 Valid
A SYSTEMATIC REVISION OF ARCHOCENTRUS Zootaxa 1603 © 2007 Magnolia Press · 5

To complicate things further, variation within some Archocentrus species has not been evaluated before.
Hubbs (1936) observed tantalizing meristic and morphometric differences among populations of Cichlasoma
octofasciatum, but he chose not to describe any subspecies because, in his material, geographic differences
were confounded with differences in body size.
Considering the potential for undetected biodiversity within presently recognized species, especially those
more widely distributed, lectotypes are designated in the present work: Amatitlania nigrofasciata, Archocen-
trus spinosissimus, Cryptoheros septemfasciatus, Cr. spilurus, and Rocio octofasciata. Cichlasoma immacula-
tum is shown to be a synonym of Cryptoheros spilurus and not of Archocentrus spinosissimus, although it was
first described as a “variety” of the latter.
The present revision treats all nominal species ever assigned to the genus Archocentrus, as well as species
that have appeared in the same clade as (or as sister group to) Archocentrus centrarchus in relevant phyloge-
netic analyses (Roe et al. 1997, Martin & Bermingham 1998, Farias et al. 2000). Six new species are
described and a seventh is resurrected from synonymy. Archocentrus, Cryptoheros, and Hypsophrys are rede-
fined, and two new genera (Amatitlania and Rocio) are diagnosed.
Genus–level decisions in this work are supported by a phylogeny (Schmitter-Soto, in press), published
separately, which includes most character illustrations and analysis.
Material and methods
The specimens studied include representatives of all nominal species that have at various times been assigned
to the genus Archocentrus (Table 1), as well as comparative material of species that appeared in the same
clade(s) as species assigned to Archocentrus in Roe et al. (1997), Martin & Bermingham (1998), and Farias et
al. (2000) (Table 2). Material of the more widely ranging species was selected in order to include the species’
entire geographic range (see Material examined in Appendix 1).
TABLE 2. Comparative material; the references listed are those that found the species in a clade with putative Archocen-
trus spp. Authorship, year of description, orthography, generic assignation, and present status or name, after Eschmeyer
2005.
Species Author and year Original genus Reference

Petenia splendida Günther, 1862 Petenia Farias et al. 2000
Tomocichla sieboldii Kner, 1863 Heros Martin & Bermingham 1998
Hypsophrys nicaraguensis Günther, 1864 Heros Martin & Bermingham 1998
Parachromis dovii Günther, 1864 Heros Roe et al. 1997
Caquetaia spectabilis Steindachner, 1865 Acara Farias et al. 2000
Hypsophrys nematopus Günther, 1867 Neetroplus Martin & Bermingham 1998
Parachromis loisellei Bussing, 1989 Cichlasoma Martin & Bermingham 1998
Pigmentation patterns were assessed primarily from preserved specimens. Osteological methodology (i.e.,
clearing and staining) follows Taylor and van Dyke (1985), as modified by W.L. Fink (pers. comm.). Radio-
graphs and skeletonized specimens were also used. Bone measurements were taken with an ocular microme-
ter. Some body measurements (e.g. interorbital width) were taken with vernier calipers, but most were taken
from digital photographs using tpsDig (Rohlf 1999). Measurements and counts are standard (Hubbs & Lagler
1958), except that lateral-line scale-count methodology requires discussion. The lateral line in cichlids is sep-
arated into two disconnected segments, which overlap on the posterior part of the body (see Nelson 1994:

381). The procedure used here was to count those scales only in the upper of the two overlapping segments,
then include the remaining scales in the lower segment posterior to the zone of overlap in order to obtain a
total count. The last dorsal and anal fin rays were counted as one only when joined at the base, a condition
usually checked in cleared and stained specimens. The number of scales between the vent and the origin of the
pelvic fin does not include the interpelvic scale. The interpelvic scale is an enlarged scale (sometimes a group
of 2–3 scales) between the bases of the innermost pelvic-fin rays, overlapping the rays; the scales included in
the count do not overlap any ray. Anal creases refer to the intestinal folds, as counted on the anus, viewed
from the outside.
Gut-coiling patterns were illustrated from the right side of the fish, in order to keep stomach and liver,
which are on the left side, out of view. Most widespread within the studied species is the pattern here termed
“simple,” or “S-shaped,” which is retained from juveniles to adults in most cichlids: the gut depicts an S,
forming first (from the vent to the mouth) an “anal loop” rostrally, then turning caudad to form a “medial
loop,” then again rostrad to a “rostral esophageal loop,” then once again caudad to a “caudal esophageal
loop,” and finally to the esophagus, which it joins at approximately the same point as the stomach. The rostral
esophageal loop and the caudal esophageal loop are less abruptly folded than the anal and median loops,
which are rather evenly rounded. The basic simple pattern shows variations of taxonomic interest. The Inter-
secting Point (IP) is a complexity index defined by Yamaoka (1982) as the number of intersections of the gut
with an imaginary left–right line (here modified as a dorsoventral line).
Results
The identification key provided here includes all species in the genera Archocentrus, Cryptoheros, Amatitla-
nia, Rocio, and Hypsophrys, all of which include species once assigned to Archocentrus. The key has been
modified from various sources (Villa 1982; Greenfield & Thomerson 1997; Bussing 1998; Schmitter-Soto
1998; Miller et al. 2005). The genera Amphilophus, Caquetaia, and Parachromis, which showed up within the
ingroup in the phylogenetic hypothesis of Schmitter-Soto (in press) are included for comparison. Abbrevia-
tions: SL, standard length; D., dorsal fin; A., anal fin; Roman numerals after D. or A. indicate spines; Arabic
numerals indicate rays.
Key to Archocentrus and allied genera
1a Maxilla very protractile; ascending process of premaxilla extending beyond orbit, contained ininter-
orbital groove; caudal fin profile strongly rounded .............................................................. Caquetaia
1b Maxilla and premaxilla not as above; caudal fin profile, if rounded, not as strongly as above .........2
2a(1b) Caudal fin emarginate ...................................................................................................Hypsophrys (8)
2b Caudal fin rounded or (sub)truncate ................................................................................................. ..3
3a(2b) Genital papilla long, contours mostly parallel, as seen from rostral face, terminal notch rounded;
upper symphysial teeth abruptly larger than adjacent teeth; lower symphysial teeth smaller than
adjacent teeth, which are enlarged; frenum on lower lip absent.......................................Parachromis
3b Genital papilla not as above; outer teeth on upper jaw gradually increasing in size anteriorly
(symphysial teeth abruptly larger in Rocio, Amatitlania, and some Cryptoheros); frenum on lower
lip present or absent .............................................................................................................................4
4a(3b) Frenum on lower lip absent; total gill-rakers on first arch 14–20; pharyngeal plate longer than wide .
......................................................................................................................................... Amphilophus
4b Frenum present in lower lip or absent; total gill-rakers on first arch 7–15; pharyngeal plate about as

wide as long, or wider......................................................................................................................... 5
5a(4b) Lateral stripe extending from snout to a blotch on mid-body; eight bars on sides; ocellated spot on
caudal fin, dorsal to lateral line; lower jaw usually extending beyond upper jaw..................Rocio (9)
5b Lateral stripe absent; usually 6–7 bars on side of body; caudal blotch, if present, not ocellated, not
entirely dorsal to lateral line, or not entirely on fin; mouth usually terminal ......................................6
6a(5b) First bar on side of body Y-shaped, caudal arm discontinuous; caudal blotch on fin (sometimes obso-
lete) .................................................................................................................... ……Amatitlania (11)
6b First bar on side of body I- or V-shaped (may be Y-shaped in some Cryptoheros, but caudal arm
continuous); caudal blotch usually on peduncle (may lie on fin in some Cryptoheros)......................7
7a(6b) A. IX–XII; genital papilla oval, in females opening strongly crenulated; two interorbital bands.........
..................................................................................................................................Archocentrus (14)
7b A. VI–IX; genital papilla not as above; no interorbital bands ................................... Cryptoheros (16)
Key to species of genus Hypsophrys
8a(2a) Teeth pointed; profile strongly convex (Fig. 28) ........................................ Hypsophrys nicaraguensis
8b Teeth incisor-like; profile with a pronounced slope (Fig. 29) ......................... Hypsophrys nematopus
Key to species of genus Rocio
9a(5a) Pelvic fins usually falling short of anal fin origin; distally two scale rows between longest dorsal-fin
rays; dentary pores usually 4, sometimes 5; total gill-rakers on first arch 8–10; lateral blotch rounded
(Fig. 25); isolated secondary pored scales on caudal fin in addition to pores on extended lateral line;
abdomen reddish in life .................................................................................................... Rocio ocotal
9b Pelvic fins almost always reaching anal fin origin; distally one scale row between longest dorsal-fin
rays; dentary pores always 4; total gill-rakers on first arch 9–12; lateral blotch squarish; no pored
scales on caudal fin, other than one or two on extended lateral line; abdomen not reddish in life ..10
10a(9b) Maxilla reaching both an imaginary vertical line from anteriormost rim of orbit, and a horizontal
line from inferiormost rim of orbit; orbital diameter 25–31% of head length and greater than 85% of
snout length; cheek-scale rows 7; scales from vent to interpelvic scale 9–10; dorsal and anal fins not
bearing filaments; spots on side of body larger than scales and not clearly aligned ....Rocio gemmata
10b. Maxilla reaching only a horizontal line from inferiormost rim of orbit, not a vertical line from
anteriormost rim of orbit; orbital diameter 21–25% of head length and less than 80% of snout length;
cheek-scale rows modally 6 (4-7); scales from vent to interpelvic scale 10–13; dorsal and anal fins
usually bearing filaments; spots on side of body smaller than scales and aligned in ca. 15 regular
series (Fig. 24) .......................................................................................………….Rocio octofasciata
Key to species of genus Amatitlania
11a(6a) Circumpeduncular scales modally 15–16; body depth usually greater than 50% of SL, always
greater than 48% of SL; stripe from snout to eye usually well defined............................................ 12
11b Circumpeduncular scales modally 17–18; body depth usually less than 48% of SL, always less than
50% of SL; stripe from snout to eye usually diffuse......................................................................... 13
12a(11a) Eye blue, gold-rimmed; modally 2–2.5 scales from lateral line to first dorsal fin ray (Fig. 20);
caudal vertebrae modally 15 .................................................................... …………Amatitlania kanna
12b Eye bluish or greenish; modally 1.5 scales from lateral line to first dorsal fin ray (Fig. 21); caudal
vertebrae modally 14................................................................................ …………Amatitlania siquia

13a(11b) Fourth bar on side of body Y-shaped (Fig. 19); body depth usually less than 47% of SL, always
less than 48% of SL; lower lip often with a caudad projection at dorsal angle; abdominal vertebrae
13 ........................................................................................................... …….Amatitlania coatepeque
13b Fourth bar on side of body I-shaped (Fig. 18); body depth usually greater than 47% of SL, always
greater than 46% of SL; lower lip, normal at angle; abdominal vertebrae 12 .......... …………………
................................................................................................................…..Amatitlania nigrofasciata
Key to species of genus Archocentrus
14a(7a) Teeth tricuspid (except symphysial, sometimes truncate) ........................ Archocentrus multispinosus
14b All teeth pointed ................................................................................................................................ 15
15a(14b) Two opercular spots (Fig. 2); D. XVI; A. IX–XI ........................................ Archocentrus centrarchus
15b One opercular spot (part of longitudinal stripe) (Fig. 4); D. XVIII–XIX; A. XI–XII ...........................
.............................................................................................................. ….Archocentrus spinosissimus
Key to subgenera of genus Cryptoheros
16a(7b) Teeth truncate and labiolingually compressed, incisor-like...................................................................

………………………………………..Subgenus Panamius (one species: Cryptoheros panamensis)
16b All teeth pointed ................................................................................................................................ 17
17a(16b) Lateral spot usually distinct (not distinct from lateral bar in C. sajica); 1.5–2.5 scales between lateral
line and first dorsal-fin ray; dorsal-fin interradial scale rows 2–7 scales long, distally in one row
(sometimes with sporadic supplementary scales, but not distally); in females opening of genital
papilla V-shaped at its rostral end ................................................…...……...Subgenus Bussingius, 18
17b No well-defined lateral spot; scales between lateral line and first dorsal fin ray 2–3.5; dorsal-fin
interradial scale rows 4–11 scales long, distally in two rows; opening of genital papilla not as above
.................................................................................................................... Subgenus Cryptoheros, 22
Key to species of genus Cryptoheros (subgenus Bussingius)
18a(17a) Bars on side of body of uniform width (Fig. 16); no lateral spot discernible from third bar; no
blotch on dorsal fin; caudal blotch tenuous or absent; A. VI–VIII......................... Cryptoheros sajica
18b Bars on side of body, not uniform in width; lateral spot present; a black, ocellated blotch on
spinous dorsal fin of mature females; caudal blotch usually present (at least tenuous); A. VIII–IX 19
19a(18b) Lateral spot circular (Fig. 14); caudal blotch tenuous; iris metallic blue in life; breast and throat
orange in life; axil of pectoral fin somewhat dark to black; predorsal scales modally 11.....................
.............................................................................................................………….Cryptoheros myrnae
19b Lateral spot, if present, oval or squarish; caudal blotch tenuous or marked; iris not metallic blue in
life; breast and throat not orange in life, axil of pectoral fin with same coloration as breast or
somewhat darker; predorsal scales modally 12 or more................................................................... 20
20a(19b) Breast, fins and throat greyish; no longitudinal stripe on side of body; dorsal-fin interradial scale
rows not imbricated (i.e. with no supplementary scales); jaw teeth not oval in section; opercular
spot indistinct (Fig. 12); anal creases 12–16 .......................................... Cryptoheros septemfasciatus
20b Breast, fins and throat yellow or yellowish; a longitudinal stripe from opercle to pectoral-fin origin;
dorsal-fin interradial scale rows imbricated (i.e. with supplementary scales); jaw teeth oval in
section, widest medially; opercular spot usually distinct; anal creases 10–14 ................................. 21
21a(20b) Breast, fins and throat only yellowish in life; bars on side of body indistinct (Fig. 13); body depth

45–55% of SL ...................................................................................... ……….Cryptoheros altoflavus
21b Breast, fins and throat intensely yellow in life; bars on side of body intensified medially and
dorsally, as series of spots (Fig. 15); body depth 44–50% of SL ........ ...........Cryptoheros nanoluteus
Key to species of genus Cryptoheros (subgenus Cryptoheros)
22a(17b) First bar on side of body divided, or at least dorsally expanded (Fig. 7); body deep, always greater
than 45% of SL; bars on side of body of uniform intensity.................................Cryptoheros spilurus
22b First bar on side of body not divided, nor distinctly expanded dorsally; body deep or not; bars on
sides of body alternating in intensity or not.......................................................................................23
23a(22b) Bars on side of body of alternating intensity, the second much lighter than the first and third (Fig.
11); body depth usually greater than 49% of SL, always greater than 44% of SL; spot on tail usually
saddled (i.e., extending over top, but not under bottom, of caudal peduncle); lower gill-rakers on
first arch modally 7; circumpeduncular scales modally 19; scales from lateral line to first dorsal fin
ray modally 2.5–3.5; distance between origin of pectoral and first lateral line scale, usually 14–15%
of SL ...................................................................................................................... Cryptoheros cutteri
23b Bars on side of body of uniform intensity (Fig. 9); body depth usually less than 47% of SL; always
less than 48% of SL; spot on tail usually not saddled (i.e., extending neither over top nor under
bottom of caudal peduncle, or else extending both over top and under bottom of peduncle); lower
gill-rakers on first arch modally 6; circumpeduncular scales modally 17; scales from lateral line to
first dorsal fin ray modally 2–2.5; distance between origin of pectoral and first lateral line scale
usually 12–13% of SL ............................................................................... Cryptoheros chetumalensis
Systematic accounts
This section deals only with Archocentrus, Cryptoheros, the two new genera described herein, and Hyp-
sophrys. Where data on gut morphology are omitted, no specimens were available for dissection and examina-
tion. Material examined is listed in Appendix 1.
Genus Archocentrus Gill
Heros (Archocentrus) Gill in Gill & Bransford, 1877: 185 (original description as subgenus).
Herotilapia Pellegrin, 1904: 211 (junior synonym).
Cichlosoma, “section” Archocentrus, Regan 1905: 74 (new combination).
Archocentrus, Allgayer 1994: 13 (new status).
Type species. Heros (Archocentrus) centrarchus Gill, 1877, by monotypy.
Diagnosis. Genus distinguished by three strict synapomorphies (Schmitter-Soto, in press). Anal pterygio-
phores on first haemal spine usually five, anal-fin spines modally 11–12, genital papilla oval, the opening
strongly crenulated or deeply notched; notch not rounded. Further diagnosed from similar genera by a con-
cave posterior end of maxilla, except in Archocentrus centrarchus, and peritoneum moderately pigmented
dorsolaterally (also in Cryptoheros, Petenia, and Amatitlania kanna). Two interorbital bands (also in Rocio).
Description. D. XVI–XIX,7–10; A. X–XIII,7–9; scale rows on cheek 4–6; pored lateral-line scales 25–
29; predorsal scales 12–17; scales from vent to interpelvic scale 6–9; circumpeduncular scales 15–21; anal
creases 10–15. Species small, less than 110 mm SL. Body rather oval, deep, depth 47–61% of SL. Head length

33–35% of SL. Snout short, length about 25% of head length. Orbital diameter 24–33% of head length. Cau-
dal peduncle twice as deep as long; least depth 16–19% of SL. Head profile straight or concave to above
orbits, convex at nape. Lips not medially narrow; pectoral and pelvic fins long, always reaching posteriorly to,
or beyond, 3rd anal-fin spine. Origin of pelvic fin posterior to a vertical from origin of dorsal fin. Gut equal or
longer than body. Seven vertical bars on side of body; dorsal and anal fins immaculate or with dots in a check-
ered or diagonally-aligned pattern. A blotch on scaly base of caudal fin, which crosses lateral line, ocellated or
not.
Distribution. Pacific slope of Central America from Costa Rica (Río Tempisque) to Nicaragua (Río Gua-
saule); Atlantic slope from Costa Rica (Río Matina) through the Great Lakes of Nicaragua to Guatemala (trib-
utaries of Lago Izabal) (Fig. 1).
FIGURE 1. Distribution of the species in the genus Archocentrus. Squares, Ar. centrarchus; circles, Ar. multispinosus;
triangles, Ar. spinosissimus. A symbol may represent more than one collecting site.
Species composition. Archocentrus as restricted here includes only three species: Ar. centrarchus, Ar.
multispinosus, and Ar. spinosissimus.
Remarks. The synonymization of Herotilapia is discussed and justified below.

Archocentrus centrarchus (Gill, 1877)
Figures 1–2
Heros (Archocentrus) centrarchus Gill in Gill & Bransford 1877: 185 (original description).
Cichlasoma (Archocentrus) centrarchus, Jordan & Evermann 1898: 1526 (new combination).
Archocentrus centrarchus, Allgayer 1994: 15 (new combination).
Holotype. USNM 16878, J. F. Bransford, Mar. 1, 1876. Lake Nicaragua, Nicaragua. No paratypes (see
Remarks).
Diagnosis. Autapomorphies (Schmitter-Soto, in press): total gill rakers on first arch 16–19; gill rakers
elongate, slender; two opercular spots, vertically aligned. Further diagnosed from other species of Archocen-
trus by absence of a concavity at posterior end of maxilla; circumpeduncular scales modally 18 or fewer (vs.
19 or more); presence (vs. absence) of rows of secondary pored scales on caudal fin; bars on side of body
extending partially onto dorsal and anal fins (vs. bars confined to sides of body), absence (vs. presence) of
medial intensification in bars on side of body, presence (vs. absence) of ocellus on dorsal fin of mature
females; abdomen predominantly yellow-green in life (vs. abdomen predominantly bluish, whitish, yellow or
black); and caudal blotch ocellated (vs. not ocellated).
Description. D. XV–XVII,7–9; A. IX–XI,7–9. Gill rakers on lower limb of first arch 12–17; gill rakers
long, especially on ventral side, and serrated. Scale rows on cheek 4–6; pored lateral-line scales (not counting
scales overlapping between the two segments of the lateral line) 25–29; scales from lateral line to base of first
dorsal-fin ray 1.5–3; circumpeduncular scales 15–17. Additional meristic data appear in Table 3.
Largest examined specimen 102 mm SL. Body rather oval, deep (50–57% of SL), horizontally almost
symmetrical in profile. Head profile straight or concave above orbits, convex at nape. Head length 33–34% of
SL; orbital diameter 24–27% of head length (further morphometric data appear in Table 4). Maxilla extending
to ventral rim of orbit; premaxilla extending to anterior rim of orbit. Teeth embedded, conical, slightly ret-
rorse, upper symphysial teeth not abruptly larger, lower symphysial teeth subequal to adjacent teeth. Lower
jaw slightly protruding. Lower lip at corner of mouth not tapering, rounded or squarish. Frenum of lower lip
usually absent (not “present”, contra to the original description).
Pectoral fins reaching posteriorly to 5th–6th anal-fin spine; pelvic fins reaching to 6th–9th anal-fin spine; dis-
tal pelvic ray not bearing filaments. Filamentous rays of dorsal fin extending to about middle of caudal fin.
Caudal fin definitely truncate to slightly emarginate. Scales weakly ctenoid. Subsidiary pored scales on caudal
fin forming 3-scale–long rows; scales between anal fin rays in one or two rows, 6–11 scales long.
Gut simple, about as long as body; intestinal anal and esophageal loops adjacent. Genital papilla tongue-
shaped, in adults deeply notched or strongly crenulate; longer than wide in females, may be wider than long in
males; pigmented on tip, but with little pigment on posterior side.
A vertical bar on head; two interorbital bands; a suborbital streak; diffuse stripe from snout to eye; no
speckles on cheek; two spots characteristically on opercle. Eyes golden, yellow or reddish. Seven vertical bars
on sides; 1st bar curved on head, all bars well marked medially and dorsally, 4th and 6th bars extending onto
dorsal fin, no lateral spot discernible on bars. Blotch on dorsal fin over 6th bar sometimes ocellated; soft dorsal
fin occasionally with two rows of dots forming a checkered pattern, but elsewhere unpaired fins mostly
immaculate,. Rows of spots on sides 11–14, smaller than scales; breast region golden in juveniles, bronze-
green in adults. Axil of pectoral fin and base of pectoral fin with same coloration as breast. A black caudal
blotch, on fin, across lateral line (not “chiefly above,” contra the original description), ocellated.

TABLE 3. Meristics and other quantitative discrete characters of species in the genera Archocentrus, Cryptoheros, Ama-
titlania, Rocio, and Hypsophrys. Numbers are absolute frequencies; sample size varied among traits and species.
Pectoral fin reaching caudad on anal-fin spines (in parentheses: pelvic fin reaching posteriorly on anal-fin spines)
not to 1st to 2nd to 3rd to 4th to 5th to 6th to 7th to 8th to 9th to 10th
reaching spine
Archocentrus
centrarchus 3 5(3) 2(5) (2) (1)
spinosissimus 1 1(2) 1(1) 7(4) 1(2) (1) (1)
multispinosus 5 4(1) 1(7) (1)
Cryptoheros
panamensis 3(1) (1) (1)
spilurus (1) 5(5) 4(6) (10) 1(1) (1)
chetumalensis 1(1) 1(1) 1(1) 1(1) 1(1)
cutteri 2 3(3) 3(5) 4(2) 1(4) 1 (1)
septemfasciatus 3 5 3(6) 1(5) (2) (1)
altoflavus 1 1 1(1) (1) (1)
myrnae 6 3(2) 1(4) (1)
nanoluteus 3(1) (1) (1)
sajica 1 5(1) 3(4) 1(5)
Amatitlania
nigrofasciata 2(1) 4(2) 8(3) 5(11) 3(2) 1(4)
coatepeque 1 5(1) (2) 4(5) 1(4) 1(2)
kanna 2 2(3) 1 4(2) (3) (5) (1)
siquia (1) 5 7(1) 6(9) 3(1) 1(6) (5) (2) (1)
Rocio
octofasciata 1(2) 2(1) 13(14) 13(36) 5(54) (13) (7) (1)
ocotal 3(2) (2) 1 1(1) (1)
gemmata 1 1(1) (1) 1(1)
Hypsophrys
nicaraguensis 2 1 2 4(4) 2(3) 1(3)
nematopus 3 (1) (1) (1)

TABLE 3 (cont.)
Number of scales in longest interradial row on soft dorsal fin
2 3 4 5 6 7 8 9 10 11 12 >13
Archocentrus
centrarchus 3 5 1
spinosissimus 1 4 5 1 1
multispinosus 2 2 5 1
Cryptoheros
panamensis 1 1 1
spilurus 1 6 1 3 2 2 1 2 1 4
chetumalensis 1 1 1 1 1 1 1 1 1
cutteri 2 4 7 9 3 4 3 4 2 4
septemfasciatus 1 7 7 2 2
altoflavus 2 1 1
myrnae 2 5 4 1
nanoluteus 2 1 1 1
sajica 2 4 3 1
Amatitlania
nigrofasciata 1 6 3 1 1 1
coatepeque 1 3 4 1 1 1
kanna 1 1 1 1 1 1 1 1
siquia 6 7 2 1 1
Rocio
octofasciata 1 15 15 4 1 1
ocotal 2 3
gemmata 1 1
Hypsophrys
nicaraguensis 6 2 1
nematopus 2 1 7 1

TABLE 3 (cont.)
Number of scales in longest interradial row on soft anal fin
1 2 3 4 5 6 7 8 9 10 11 12 >13
Archocentrus
centrarchus 6 2 1 1 1 1
spinosissimus 1 1 1 4 1 2
multispinosus 2 1 3 1 2 1 2
Cryptoheros
panamensis 1 1 1 1
spilurus 1 1 1 4 1 2 1 3
chetumalensis 1 1 1 1 1 1
cutteri 4 3 1 2 3 4 1 1
septemfasciatus 3 7 5 1
altoflavus 1 1 1 1
myrnae 2 4 1
nanoluteus 1 1 1
sajica 2 7 1 1
Amatitlania
nigrofasciata 2 5 3 1
coatepeque 1 5 2 2 1
kanna 1 1 1 1
siquia 3 1 4 5 1
Rocio
octofasciata 2 12 6 3
ocotal 2 1 1
gemmata 1 1
Hypsophrys
nicaraguensis 2 3 4
nematopus 1 1 1 2 1 1 2 2

TABLE 3 (cont.)
Dorsal-fin spines Dorsal-fin rays
XV XVI XVII XVIII XIX XX 7 8 9 10 11 12
Archocentrus
centrarchus 10 2 1 9 2
Cryptoheros
panamensis 1 2 9 1 2 6 3 1
spilurus 12 40 13 1 21 34 9
chetumalensis 2 20 3 1 1 5 7
cutteri 10 45 6 10 41 13
altoflavus 2 4 2 2 1
myrnae 8 6 1 11 1
nanoluteus 1 3 3 1
sajica 11 1 1 11
Amatitlania
nigrofasciata 9 55 4 2 45 23 3
kanna 1 13 4 13 4 1 1
siquia 23 64 3 1 26 62 12
Rocio
octofasciata 26 122 16 13 48 106 7
ocotal 5 1 2 4
gemmata 3 1 2
Hypsophrys
nicaraguensis 3 8 1 1 7 4
nematopus 3 10 5 7 1

TABLE 3 (cont.)
Anal-fin spines Anal-fin rays
VI VII VIII IX X XI XII XIII 6 7 8 9
Archocentrus
centrarchus 4 8 2 10
Cryptoheros
panamensis 8 3 3 6
spilurus 1 15 43 7 7 52 8
cutteri 7 42 10 1 10 36 8
septemfasciatus 8 12 3 3 14 3
altoflavus 1 4 1 2 2
myrnae 4 9 1 9 4
nanoluteus 2 1 2 1
sajica 1 10 1 5 7
Amatitlania
kanna 3 14 1 2 11 5
siquia 16 52 24 1 6 50 51 1
Rocio
octofasciata 1 1 103 71 1 62 141 18
ocotal 2 2 1 1 1 1
gemmata 2 1 1 2
Hypsophrys
nicaraguensis 4 9 6 6 1
nematopus 11 1 1 3 10

TABLE 3 (cont.)
Pectoral-fin rays
13 14 15 16 17
Archocentrus
centrarchus 5 5
spinosissimus 3 4 6 1
multispinosus 1 1
Cryptoheros
panamensis 3 1
spilurus 1 26 6
chetumalensis 2 1 1
cutteri 1 8 12
septemfasciatus 1 1
altoflavus 1 1
myrnae 1 1
nanoluteus 1 1
sajica 1 1 1
Amatitlania
nigrofasciata 10 5 9
coatepeque 8 6 1
kanna 1 6
siquia 15 6
Rocio
octofasciata 8 13 7
ocotal 1 1
gemmata 1 1 1
Hypsophrys
nicaraguensis 1 1
nematopus 1 1

TABLE 3 (cont.)
Gill-rakers on lower limb of first arch (in parentheses: total), not counting rudiments
5 6 7 8 9 10 11 12 13 14 15 16 17 18 >19
Archocentrus
centrarchus 1 1 8 1 1(1) (1) (1) (10)
spinosissimus 1 8 5(1) (8) (4) (10)
multispinosus 8 2 (8) (2)
Cryptoheros
panamensis 6 4 (4) (6)
spilurus 4 62 10(3) (43) (8) (4)
chetumalensis 2 10 4(2) (10) (3) (1)
cutteri 1 21 38(1) (14) (33) (9)
septemfasciatus 14 1 2(5) (11) (1)
altoflavus 3 1 (2) (2)
myrnae 11 3 (12) (2)
nanoluteus 3 1(1) (2) (1) (1)
sajica 10 1 (10) (1)
Amatitlania
nigrofasciata 13 23 7(13) 21(25) (8) (1)
coatepeque 15 10 (11) (13) (1) (1)
kanna 2 9 5(2) (7) (4)
siquia 25 38 8(18) (61) (9) (2)
Rocio
octofasciata 5 57 42(3) 3(36) (51) (16) (2)
ocotal 1 5 (1) (4) (1)
gemmata 2 1 (2) (1)
Hypsophrys
nicaraguensis 1 8 4(1) (8) (2) (1) (1)
nematopus 8 1 (9)

TABLE 3 (cont.)
Scale rows on cheek (in parentheses: scales between interpelvic scale and vent)
2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
Archocentrus
centrarchus 2 8 1(3) (7)
spinosissimus 2 8 1(5) (3) (3) (1)
multispinosus 4 6 (3) (4) (3)
Cryptoheros
panamensis 1 9 1 (3) (3) (1) (1) (1)
spilurus 18 22 3(1) (2) (6) (6) (2) (3) (3) (1)
chetumalensis 7 2 (1) (1) (5) (1) (1) (1) (1)
cutteri 12 15 3 1(2) (6) (5) (6) (4) (1) (1) (1)
septemfasciatus 3 5 (10) (7) (3)
altoflavus 3 (2) (1) (1)
myrnae 4 8 (2) (1) (4) (3)
nanoluteus 3 1 (2)
sajica 1 1 (1) (1) (5) (1)
Amatitlania
nigrofasciata 5 14 1 (3) (1) (4) (1)
coatepeque 10 2 (5) (4) (4) (3)
kanna 6 18 2(1) (1) (3) (1)
siquia 20 21 2(1) (4) (9) (4) (1)
Rocio
octofasciata 2 35 39 12(1) 2(5) (14) (21) (26) (19) (8) (1) (4)
ocotal 2 3 (1) (1) (2) (2)
gemmata 3 (1) (2)
Hypsophrys
nicaraguensis 1 1 (1) (1) (1) (1)
nematopus 7 2 (3) (6)

TABLE 3 (cont.)
Number of predorsal scales
9 10 11 12 13 14 15 16 17 18 19 20
Archocentrus
centrarchus 2 2 4 1 1 2
Cryptoheros
panamensis 4 4 1 1
spilurus 2 8 14 8 2 6 2 1
cutteri 2 4 10 3 8 4 1
altoflavus 2 2 1 1
myrnae 3 7 3 1 2
nanoluteus 1 3
sajica 2 2 3 2 2
Amatitlania
nigrofasciata 1 1 2 2 3 3 1
coatepeque 1 1 2 1 2 5 1 3
kanna 2 3 1 3 1
siquia 2 2 1 5 3 2 1
Rocio
octofasciata 1 1 4 10 6 4 1 1
ocotal 1 1 1 1
gemmata 1 1
Hypsophrys
nicaraguensis 1 2 4 2 2
nematopus 2 3 1 1 1 1

TABLE 3 (cont.)
Scale rows from lateral line to base of first dorsal-fin ray

(in parentheses: scale rows from lateral line to base of first dorsal-fin spine)
1 1.5 2 2.5 3 3.5 4 4.5 5 5.5 6 6.5 7
Archocentrus
centrarchus 3 6 1 1 (3) (6) (1) (1)
spinosissimus 4 4 2 (2) (5) (5)
multispinosus 10 (5) (5)
Cryptoheros
panamensis 7 1 1 (1) (8) (1)
spilurus 27 49 2 (16) (45) (14)
chetumalensis 5 9 2 (3) (6) (2) (1)
cutteri 11 42 5 1 (1) (9) (22) (24) (1)
septemfasciatus 7 12 1 (1) (10) (7) (2)
altoflavus 3 (2) (1) (1)
myrnae 1 4 10 (1) (6) (4) (4) (1)
nanoluteus 2 (2) (1)
sajica 1 1 (1) (1) (1)
Amatitlania
nigrofasciata 17 35 2 (8) (30) (15) (4)
coatepeque 2 14 10 (4) (8) (8) (4)
kanna 12 6 (5) (9) (3) (2)
siquia 1 64 14 7 1 (1) (18) (34) (9) (3)
Rocio
octofasciata 5 15 1(5) 1(11) (5) (1)
ocotal 2 1 3(2) (3) (1)
gemmata 2 1(1) (2)
Hypsophrys
nicaraguensis 10 (1) (9)
nematopus 10 (9) (1)

TABLE 3 (cont.)
Pored lateral-line scales, caudal section

(in parentheses: total count, not counting overlap between the two sections of lateral line)
<7 8 9 10 11 12 >13 <25 26 27 28 29 30 31 >32
Archocentrus
centrarchus 2 5 3 1 (1) (5) (4) (1) (1)
spinosissimus 5 6 3 (2) (4) (7)
multispinosus 1 5 3 1 (4) (5) (2)
Cryptoheros
panamensis 3 2 3 3 1 (1) (3) (1) (4) (3)
spilurus 2 14 27 23 7 2 (3) (22) (31) (17) (1)
chetumalensis 1 1 6 5 2 1 (1) (2) (6) (6) (1)
cutteri 2 12 23 18 1 (1) (6) (19) (26) (6)
septemfasciatus 3 4 8 6 (2) (11) (2) (1)
altoflavus 1 2 1 1 (1) (3) (1)
myrnae 2 5 4 1 (2) (5) (6)
nanoluteus 1 1 (1) (1)
sajica 1 8 1 1 (9) (3)
Amatitlania
nigrofasciata 3 13 24 12 (2) (12) (24) (10) (3)
coatepeque 1 6 11 7 (1) (6) (8) (8) (1)
kanna 1 5 6 4 1 (7) (6) (4) (1)
siquia 6 26 27 12 1 (7) (37) (27) (4)
Rocio
octofasciata 2 13 57 52 22 4 (7) (61) (60) (21) (4)
ocotal 3 1 2 (5) (1)
gemmata 2 1 (1) (1) (1)
Hypsophrys
nicaraguensis 1 4 4 2 (1) (3) (8)
nematopus 1 6 3 1 (2) (9) (2)

TABLE 3 (cont.)
Number of circumpeduncular scales
14 15 16 17 18 19 20 21 22
Archocentrus
centrarchus 4 6 1
spinosissimus 3 7 3
multispinosus 3 7 1
Cryptoheros
panamensis 1 1 9
spilurus 10 46 14 5 1
chetumalensis 2 11 3
cutteri 1 22 28 9
septemfasciatus 9 8 1
altoflavus 2
myrnae 4 7 1
nanoluteus 1 2
sajica 2 8 2
Amatitlania
nigrofasciata 1 2 24 32 8
coatepeque 1 12 12 2
kanna 1 11 5
siquia 2 36 62 15 1
Rocio
ocotal 4 1 1
gemmata 1 1
Hypsophrys
nicaraguensis 1 8 1 1
nematopus 1 9 1

TABLE 3 (cont.)
Number of anal creases
10 11 12 13 14 15 16 17 >18
Archocentrus
centrarchus 3 1 1 2
spinosissimus 9
Cryptoheros
panamensis 1 1 3
spilurus 1 1 3 2 7 5 1
chetumalensis 1 1 1 1 1
cutteri 1 2 3 3 1 2 5
altoflavus 1 1 1
myrnae 1 2 2 5 1 1
nanoluteus 1 1 1
sajica 1 2 2 2 1 1
Amatitlania
coatepeque 3 3 1 1 2
kanna 1 1 1
siquia 3 1 7 1 1 1 2
Rocio
octofasciata 3 12 6 8 7 2 1
ocotal 3 3
gemmata 1 1 1
Hypsophrys
nicaraguensis 6 2
nematopus 3 1 6 1

TABLE 3 (cont.)
Caudal vertebrae Total vertebrae
<14 15 16 17 26 27 28 29 30
Archocentrus
centrarchus 1 2 1 1
spinosissimus 4 1 4 1
Cryptoheros
panamensis 1 1 1 1 1
spilurus 1 1 1 1
chetumalensis 2 1 1 1
cutteri 1 1 1 1
altoflavus 1 1
myrnae 1 6 1 1
nanoluteus 1 1
sajica 1 1 1 1
Amatitlania
nigrofasciata 2 2 2 2
coatepeque 2 2
kanna 2 2
siquia 5 4 4 5
Rocio
ocotal 6 5 1
gemmata 1 1 1 1
Hypsophrys
nicaraguensis 2 1 2 1
nematopus 3 1 3

TABLE 4. Selected morphometric proportions of cichlid species in the genera Archocentrus, Cryptoheros, Amatlania,
Rocio, and Hypsophrys. Specimens measured are those marked “dig.” in Appendix 1. HL, head length.
Body Head Inter- Orbit Post- Pre-dor- Least Gut length,

depth, length, orbital, diameter orbital, sal, %SL depth, %SL
%SL %SL %HL %HL %HL %SL
Archocentrus
centrarchus 50-57 33-34 38-40 24-27 49-50 48-49 17-18 101
spinosissimus 57-61 34-35 35-40 28-29 51-52 49-50 18-19 109
multispinosus 47-54 33-35 37-40 26-33 45-46 45-47 16-17 264-271
Cryptoheros
panamensis 45-51 30-34 29-40 27-32 47-52 41-45 16-18 nd
spilurus 52-56 32-37 27-38 28-32 40-42 43-50 15-17 97-186
chetumalensis 42-49 30-37 24-33 23-35 37-44 39-42 15-16 94-161
cutteri 44-54 32-43 23-38 20-40 42-46 40-49 14-17 82-230
septemfasciatus 42-56 30-36 28-37 23-29 49-53 38-44 16-17 92-141
altoflavus 45-56 31-36 24-31 22-35 45-49 42-44 16-17 151
myrnae 42-50 33-37 16-25 25-30 35-48 41-44 15-16 117-129
nanoluteus 44-52 29-35 23-29 27-32 42-54 44-46 16-18 150
sajica 46-49 34-37 27-32 28-31 46-49 43-45 16-18 107-116
Amatitlania
nigrofasciata 46-50 32-38 26-35 21-34 44-51 42-44 15-17 58-124
coatepeque 43-48 32-38 16-32 21-34 49-57 42-44 16-18 108-113
kanna 48-58 33-37 27-33 27-33 51-58 44-49 15-17 94
siquia 48-55 31-38 26-36 27-41 51-53 42-45 16-17 111-188
Rocio
octofasciata 39-51 33-42 24-38 21-31 48-50 39-47 15-17 67-98
ocotal 41-46 35-41 24-31 21-23 48-52 41-46 14-17 76
gemmata 41-46 35-41 24-31 25-30 49-53 43-45 15-17 102
Hypsophrys
nicaraguensis 44-46 33-34 26-36 26-30 45-46 44-45 14-15 80
nematopus 38-46 31-39 26-35 26-33 42-43 41-12 13-14 136-231
Distribution. Pacific slope, in tributaries of the Golfo de Fonseca, Honduras and Nicaragua; Atlantic
slope, from the Río Chirripó (Río Matina) of Costa Rica to the Río San Juan and associated drainages of Nic-
aragua, including the Great Lakes (Fig. 1).

Remarks. Although the specimen BMNH 1905.3.27.2 (Fig. 2) is labeled “cotype”, Gill and Bransford
(1877: 186) left no doubt that they had only one specimen (i.e., the holotype) at hand. The BMNH specimen
therefore has no formal type status.
FIGURE 2. Archocentrus centrarchus, BMNH 1905.3.27.2. Photo, BMNH, London.
Archocentrus multispinosus (Günther, 1867)

Figures 1, 3
Heros multispinosus Günther, 1867: 601 (original description).

Cichlasoma (Archocentrus) multispinosum, Jordan & Evermann 1898: 1525 (new combination).
Herotilapia multispinosa, Pellegrin 1904: 211 (new combination).
Archocentrus multispinosus, Burgess 2000: 54 (new combination).
Holotype. BMNH 1865.7.20.34, 68 mm SL (Fig. 3) , J. M. Dow. Lake Managua, Nicaragua. No paratypes.

Diagnosis. Autapomorphies (Schmitter-Soto, in press): both upper and lower symphysial teeth tricuspid;
basally two rows of interradial scales on anal fin; rostral arm of articular bone only a little longer than dorsal
process (ratio ca. 0.87, vs. 0.77 or less in other Archocentrus); gut with IP = 6. Also diagnosable from all other
Archocentrus by a sigmoid (vs. irregular) posterior edge of urohyal; coronal pore triple (vs. single); narrowest
point of dentigerous arm of premaxilla at caudal tip (vs. before tip). Truncate caudal-fin profile, synapomor-
phic with A. centrarchus (vs. rounded-truncate in A. spinosissimus).
Description. D. XVIII–XIX,8–9; A. XI–XII,7–8; gill rakers on lower limb of first arch 8–9; gill rakers
blunt, their ends trapezoidal, serrated. Scale rows on cheek 4–5 (contra Stawikowski & Werner 1998, who
counted 3); pored lateral-line scales (not counting scales overlapping between the two segments of the lateral
line) 27–29; 2.5 scales from lateral line to base of first dorsal-fin ray; circumpeduncular scales 19–21 (further
meristic data appear in Table 3).
Largest examined specimen 87 mm SL. Body rather oval, usually not so deep (47–54% of SL) and not so
symmetrical horizontally, the curve of the nape somewhat more pronounced than the curve of the thoracic
region. Head profile straight above orbits, convex at nape. Head length 33–35% of SL; orbital diameter 26–

33% of head length (other morphometric data appear in Table 4). Maxilla extending to ventral rim of orbit;
premaxillary dorsal process not reaching anterior rim of orbit. Teeth not embedded; incisors tricuspid (Pelle-
grin 1904: fig. 20, 4), symphysial teeth aligned with adjacent teeth. Lower jaw not protruding. Lower lip at
corner of mouth square, not rounded, the angle varying from right-angled to acute. Frenum present on lower
lip.
FIGURE 3. Archocentrus multispinosus, holotype, BMNH 1865.7.20.34. Photo, BMNH, London.
Pectoral fins reaching posteriorly to 3rd–5th anal-fin spine, and pelvic fins extending to 4th–6th anal-fin
spine; distal pelvic ray may bear filaments. Filamentous rays of dorsal and anal fins extending to about proxi-
mal third of caudal fin. Caudal fin truncate (contra Günther 1869 and Pellegrin 1904, who considered it
rounded). Scales strongly ctenoid. A few, sporadic subsidiary pored scales on caudal fin, not forming rows;
scales between dorsal fin rays, proximally in two rows, the rows up to 14 scales long.
Gut strongly coiled, IP = 6. Genital papilla longer than wide (may be wider than long in females), widest
at base, deeply notched, sunken; no pigmentation, except for base and sides.
A vertical bar on head; two interorbital bands; no suborbital streak; diffuse stripe from snout to eye; no
speckles on cheek; a spot on opercle, which represents an anterior expansion of longitudinal stripe on side of
body. Eyes orange. Seven sharp vertical bars on sides, broader than interspaces, and usually double; 1st bar dif-
fusely Y-shaped; a longitudinal stripe from orbit to a circular, jet-black blotch on 4th bar; bars on side of body
not extending onto dorsal fin. No ocellus on dorsal fin; dorsal and anal fins immaculate or with 3–4 rows of
dots. About 12 rows of spots on sides, smaller than scales; breast region olive-yellowish. Axil of pectoral fin
with same coloration as breast or dusky; base of pectoral fin with same coloration as breast or slightly paler. A
black caudal blotch, on fin, across lateral line, the blotch not ocellated.
Distribution. Atlantic slope, from Costa Rica (Río Matina) through the Great Lakes of Nicaragua to Hon-
duras (Río Patuca); Pacific slope, from Costa Rica (Río Tempisque) to Nicaragua (Río Guasaule) (Fig. 1).
Panamanian records (e.g. BMNH 1925.3.6.165) are based on misidentifications.
Remarks. Pellegrin (1904) put the species in a monotypic genus because of the “dentition tout à fait car-
actéristique” (surprisingly not observed by Günther 1867 or 1869); on the other hand, he considered it
“[v]oisin de Neetroplus.”

A. multispinosus exhibits striking and abundant autapomorphies, but its relationships to the other two
Archocentrus are not resolved (Schmitter-Soto, in press). Considering this, it seems best to apply only one
name to the monophyletic group formed by the three species.
Archocentrus spinosissimus (Vaillant & Pellegrin, 1902)

Figures 1, 4
Heros (Cichlasoma) spinosissimus Vaillant & Pellegrin 1902: 87 (original description).

Cichlasoma (Archocentrus) spinosissimum, Pellegrin 1904: 188 (new combination).
Archocentrus spinosissimus, Allgayer 1994: 15 (new combination).
Lectotype. MNHN A-0352, 69 mm SL (Fig. 4), F. Bocourt. Río Polochic, Guatemala. The lectotype, herein
designated, is the only one with an intact caudal fin.
FIGURE 4. Archocentrus spinosissimus, lectotype, MNHN A-0352. Photo, C. Ferrara.
Paralectotypes. MNHN 2005-0780 (3; 58, 59, and 74 mm SL).

Diagnosis. Autapomorphies (Schmitter-Soto, in press): vertical bar on head, extending just across nape,
not present on opercle; distally three rows of anal interradial scales. In addition, caudal fin rounded-truncate
(vs. definitely truncate) in the other species of Archocentrus.
Description. D. XVII–XIX,7–10; A. XI–XIII,7–9; gill rakers on lower limb of first arch 6–8; gill rakers
bifid or at least distally expanded, trapezoidal, serrated. Scale rows on cheek 4–6; pored lateral-line scales (not
counting scales overlapping between the two segments of the lateral line) 26–28; scales from lateral line to
base of first dorsal-fin ray 3.5–4.5; circumpeduncular scales 19–21 (additional meristic data appear in Table
3).
Largest specimen observed, 74 mm SL, the species often reaching 110 mm SL according to Conkel
(1993). Body rather oval, deeper than other species of Archocentrus (57–61% of SL), dorsal and ventral pro-
files almost symmetrical, or upper profile more strongly curved. Head profile straight above orbits, convex at

nape. Head length 34–35% of SL, head deeper than long; orbital diameter 28–29% of head length (further
morphometric data appear in Table 4). Maxilla not reaching a vertical line from anteriormost rim of orbit, but
extending to a horizontal line from inferiormost rim of orbit; premaxilla reaching anterior rim of orbit. Teeth
embedded, canine, pointed, cylindrical, rather long, slightly retrorse; symphysial teeth subequal to adjacent
teeth. Lower jaw slightly protruding. Lower lip tapering, the ventral corner acute. Frenum present on lower
lip.
Pectoral fins reaching posteriorly to 4th–8th anal-fin spine; pelvic fins reaching to 5th–10th anal-fin spine.
Filamentous rays of dorsal fin extending posteriorly to distal fourth of caudal fin. Caudal fin rounded-trun-
cate. Scales strongly ctenoid. Pored scales on caudal fin vestigial or absent; scales between dorsal and anal fin
rays distally in two or three rows, up to 8 scales long.
Gut simple, anal and esophageal loops not adjacent. Genital papilla rounded or vase-shaped, wider than
long; large feminine pore, tip crenulated to deeply notched; pigmented on margins (males) or just around
opening (females).
A bar extending across nape from preopercle to preopercle; two interorbital bands; suborbital streak usu-
ally pointed, slightly concave dorsally; no stripe from snout to eye; speckles on cheek aligned from chin to
angle of preopercle and along anteroventral edge of opercle; only one opercular spot, which is part of a longi-
tudinal stripe running from orbit to lateral blotch. Eyes coppery, bluish. Seven vertical bars on sides, rather
diffuse; 1st bar diffusely Y-shaped; 1st to 5th sometimes double, none extending onto dorsal fin; a squarish or
rounded spot on 3rd bar. No ocellus on dorsal; dots on soft dorsal and anal fins in 4–6 rows, concentric on base
of third or fourth ray, a basal row all along dorsal fin. About 10–14 rows of well-marked spots on sides,
smaller than scales; breast region olive. Base of pectoral fin paler than breast, sometimes yellowish. Caudal
blotch on scaly base of fin, extending across lateral line, never forming a saddle over top of caudal peduncle,
connected to posterior speckles between rays of caudal fin.
Distribution. Ríos Polochic, Dulce, and other rivers in Lago Izabal drainage, Caribbean versant of Guate-
mala (Fig. 1).
Remarks. Pellegrin (1904) considered that the species “présente tout à fait l’aspect d’Herotilapia multis-
pinosa […] elle n’est pas éloignée de C. nigrofasciatum […] et de C. centrarchus…”
Cichlasoma spinosissimum var. immaculata Pellegrin (1904), which occurs sympatrically with A. spino-
sissimus (and with Cryptoheros spilurus) in the Río Polochic, was considered a valid species by Regan (1905)
and Miller (1966), mainly because it lacks speckles on the fins and has up to 19 dorsal fin spines, as few as 9
anal-fin spines, and as many as 29 pored lateral-line scales. Pellegrin himself (1904) was of the opinion that,
in spite of the differences, it was not possible to categorically separate this form to species (he even called it a
“variety,” not a subspecies). Allgayer (2001) noted that it might be the female of Archocentrus spinosissimus,
the reduction or absence of speckles being a “fait aussi observé en captivité.” After examining the types, I
conclude that they are somewhat decolored (although not totally descaled, contra Allgayer 2001) specimens
of Cryptoheros spilurus. Thus, Cichlasoma immaculatum (the nomen used for this taxon by Regan [1908] and
Miller [1966]) is hereby synonymized with Cryptoheros spilurus, as discussed in the account of that species.
Genus Cryptoheros Allgayer
Cryptoheros Allgayer 2001: 14 (original description).
Type species. Heros spilurus Günther, 1862, by original designation.
Diagnosis. Genus distinguished by one strict synapomorphy (fig. 3c in Schmitter-Soto, in press): one to five
short, acute interdigitations in sutural connection between halves of lower pharyngeal jaw. Also characterized

by a twisted median loop in adult gut (except in Cr. myrnae and Cr. sajica), and peritoneum moderately pig-
mented dorsolaterally (also found in Archocentrus, Petenia, and Amatitlania kanna).
Description. D. XVI–XIX,9–11; A. VII–X (VI in Cr. panamensis),7–9. Gill rakers on lower limb of first
arch 6–7, occasionally 5. Pored lateral-line scales 26–30; circumpeduncular scales 16–18, rarely 19 or 20. A
genus of small cichlids, less than 130 mm SL. Body rather oval, deep, depth 42–56% of SL. Maxilla not
reaching orbit, premaxillary dorsal process not reaching anterior rim of orbit (except in young). Frenum
present in lower lip. Filamentous rays of dorsal and anal fins extending to mid-caudal fin or beyond. Origin of
pelvic fin located behind a vertical from origin of dorsal fin. Caudal fin truncate or rounded. Scales usually
strongly ctenoid (moderately ctenoid in Cr. sajica). Up to two pored scales on caudal fin, as an extension of
the lateral line. Gut simple, with variations (secondary loops) in some species; anal and anterior esophageal
loops adjacent. Genital papilla usually tongue-shaped or rounded. No spots on cheek; vertical bar on head,
absent or faint, as a darkening of opercle and nape; interorbital bands faint or absent. Seven vertical bars on
sides, diffuse or well marked. Breast region often yellowish or olive. Caudal blotch usually two-thirds or more
on peduncle, never ocellated.
Distribution. Panama to Mexico, mostly on Atlantic versant (Fig. 5).
FIGURE 5. Distribution of the species in the genus Cryptoheros. Closed squares, Cr. panamensis; open squares, Cr. spi-
lurus; closed circles, Cr. chetumalensis; open circles, Cr. cutteri; closed triangles, Cr. septemfasciatus; open triangles, Cr.
altoflavus; “×” sign, Cr. myrnae; “+” sign, Cr. nanoluteus; stars, Cr. sajica. A symbol may represent more than one col-
lecting site.

Species composition. Nine species: Cr. spilurus, Cr. chetumalensis, Cr. cutteri, Cr. septemfasciatus, Cr.
altoflavus, Cr. myrnae, Cr. nanoluteus, Cr. sajica, and Cr. panamensis. Three clades diagnosable, here
described as subgenera, one of which is monotypic.
Remarks. The generic name was introduced by Allgayer (2001) for the “smaller Archocentrus” species,
i.e., those species included here in the subgenera Cryptoheros and Bussingius, and the genus Amatitlania, but
excluding Cr. panamensis. Allgayer (2001) also united Archocentrus, Cryptoheros, and Herotilapia in a new
subtribe, Archocentrina. However, this decision was not supported by cladistic analyses, and Allgayer did not
consider conflicting evidence, such as that presented by Roe et al. (1997) (see below).
Panamius, n. subgen.
Type species. Neetroplus panamensis Meek & Hildebrand, 1913, by monotypy.
Diagnosis, description, distribution. As for the species. See below.

Species composition. Monotypic.
Etymology. From Panama, the country to which the type species is endemic. Gender masculine.
Cryptoheros panamensis (Meek & Hildebrand, 1913), n. comb.

Figures 5–6
Neetroplus panamensis Meek & Hildebrand, 1913: 90 (original description).

Cichlasoma panamense, Conkel 1993 (new combination).
Theraps panamense, Eschmeyer 2001 (new combination).
Archocentrus panamensis, Kullander 2003: 617 (new combination).
Holotype. FMNH 7601, 79 mm SL, S. E. Meek and S. F.Hildebrand, Feb. 2, 1911. Río “Mandingo” [Mand-
inga] at “Bas Obispo”, Canal Zone, Panama.
Paratypes. FMNH 8105–8112 (20), from several localities.
FIGURE 6. Cryptoheros panamensis, holotype, FMNH 7601. Photo, P. Willink.

Diagnosis. Autapomorphy (Schmitter-Soto, in press): teeth truncate and labiolingually compressed, inci-
sor-like, but tips rounded (pointed in juveniles), their biting edges not forming a line. Further distinguished
from other species in the genus by having modally three rows of scale rows on cheek (vs. four or more); pres-
ence of single spot at posteroventral angle of opercle (vs. opercular spots absent, or single spot present but not
located at angle in other species except Cr. altoflavus); a medial intensification only of 4th lateral bar on side of
body (vs. either only on 3rd bar or none at all); dorsal edge of the articular bone straight (vs. convex); first neu-
ral spine retrorsely directed (vs. antrorse in all other Cryptoheros, except Cr. spilurus and Cr. cutteri), and two
dorsal elements between first two epineural spines (vs. three). Cr. panamensis is the only member of its genus
in which the paired fins seldom or never extend posteriorly to the first anal-fin spine, and also the only one
with six anal-fin spines (vs. seven or more) and as few as 16 dorsal-fin spines (vs. always 17 or more). Sum of
dorsal- and anal-fin spines 23 (vs. 24 or more).
Description. D. XVI–XVIII,9–10; A. VI,7–8. Gill rakers on lower limb of first arch 6–7; gill rakers trap-
ezoidal, serrated, no basal process. Scale rows on cheek 2–4; predorsal scales 9–12; pored lateral-line scales
(not counting scales overlapping between the two segments of the lateral line) 25–29; scales between lateral
line and origin of dorsal fin 3.5–4.5; scales from lateral line to base of first dorsal-fin ray 1.5–2.5; circumpe-
duncular scales 14–16 (further meristic data appear in Table 3).
Maximum size 130 mm SL (Conkel 1993), although largest examined specimen during this study only 79
mm SL. Body relatively deep (45–51% of SL). Head profile convex to straight. Head length 30–34% of SL;
orbital diameter 27–32% of head length (further morphometric data appear in Table 4). Teeth not embedded;
the only incisor teeth are those near symphyses, which are rounded, its edges not forming a line; other teeth
are conical (in young, even symphysial teeth conical). Upper and lower symphysial teeth lower than adjacent
teeth in adults. Lower jaw slightly receding (contra Stawikowski & Werner 1998, who considered it terminal).
Upper lip medially narrowed; lower lip square, not rounded at corner, its lower angle acute.
Pectoral fins not extending caudad to 1st anal-fin spine; pelvic fins sometimes reaching 3rd anal-fin spine.
Filamentous rays of dorsal fin reaching posteriorly to distal third of caudal fin. Scales strongly ctenoid. Two
lateral-line pored scales on caudal fin, subsidiary pores few and sporadic; scales between dorsal and anal fin
rays in one row, up to 4 scales long.
Genital papilla tongue-shaped, rounded, longer than broad, sunken; some melanophores on basal margins.
No interorbital bands; no suborbital streak; a faint stripe from snout to eye; only one opercular spot, asso-
ciated to the medial intensification of the 1st bar. Eyes greyish, reddish. No longitudinal stripe. Bars on sides
diffuse, 1st bar not divided, somewhat curved on head; 1st, 4th and posterior ones more intense medially, as
series of blotches. No bars on dorsal and anal fins, but an ocellus present on dorsal fin of mature females,
between spines X or XII to XVI or XVIII. Dots on soft dorsal and anal fins in 4–6 undulated rows, concentric
on base of third or fourth ray. About 11 rows of light spots on sides, smaller than scales; breast region olive-
yellowish. Axil of pectoral fin dark, sometimes with a spot. Base of pectoral fin either whitish or same colora-
tion as breast. Caudal blotch on both caudal fin and peduncle, two-thirds above lateral line.
Distribution. Atlantic drainages of Panama; Lake Gatún, rivers Mandinga, Chagres, Ipetí and others,
including areas formerly within the Canal Zone (Fig. 5).
Remarks. The list of objective synonyms underscores the obscurity in phylogenetic position of this spe-
cies. It appears clear, nevertheless, that its resemblance to Neetroplus (=Hypsophrys) nematopus is a conver-
gence (Rogers 1981, Schmitter-Soto, in press). The affinity with Cryptoheros was earlier recognized by
Kullander (2003), who considered it a species of Archocentrus. This relationship was overlooked by all previ-
ous workers, perhaps because of the low anal-fin spine count of Cr. panamensis, which is atypical for Archo-
centrus and related taxa (Table 3) and is approached only by Cr. sajica.

Subgenus Cryptoheros Allgayer
Cryptoheros Allgayer 2001: 14 (original description).
Type species. Same as for the genus.
Diagnosis. No strict synapomorphies discovered. Distinguished from the subgenera Panamius and Bussingius
by having the interradial scales on dorsal fin distally in two rows, 4–11 or more scales long; the absence of a
well-defined lateral spot (vs. usually distinct); scales between lateral line and origin of dorsal fin 4.5–5.5 (vs.
4.5 or fewer); scales between lateral line and first dorsal fin ray 2–3.5 (vs.1.5–2.5); opening of genital papilla
not V-shaped at its rostral end (also not so in Panamius, vs. V-shaped).
Description. D. XVII–XIX (rarely XX), 9–11 (occasionally 8); A. VIII–X (rarely VII or XI), 7–9. Gill
rakers on lower limb of first arch 5–7; gill rakers elongated but blunt, ventrally curved, and distally expanded
(club-shaped). Scale rows on cheek 4–6; predorsal scales 11–18; pored lateral-line scales (not counting scales
overlapping between the two segments of the lateral line) 26–30; scales between lateral line and origin of dor-
sal fin 4–6; scales between lateral line and first dorsal fin ray 2–3; circumpeduncular scales 16–20. Medium-
sized group of Cryptoheros, less than 112 mm SL. Head about as deep as long. Teeth bicuspid and usually
pointed, always slightly labiolingually compressed and retrorse. Symphysial teeth subequal to adjacent teeth,
not abruptly larger. Lower jaw not protruding. Pectoral and pelvic fins always reaching caudad beyond 2nd
anal-fin spine. Gut variable in length, 80–230% of SL, longer and more complex with growth. Genital papilla
rounded or oval. No interorbital bands, but postorbital–nape region may be somewhat dark; a stripe extending
from snout to eye, though sometimes diffuse; no opercular spots, but opercle dark; no longitudinal stripe; 3rd
bar on side of body and sometimes also posterior bars may extend slightly onto the base of the dorsal fin.
Body mostly olive.
Distribution. Atlantic versant, from Honduras to Mexico (Yucatan Peninsula) (Fig. 5).
Species composition. Three species: Cr. spilurus, Cr. chetumalensis, and Cr. cutteri.
Cryptoheros spilurus (Günther, 1862)

Figures 5, 7–8
Heros spilurus Günther, 1862: 289 (original description).

Cichlasoma spilurum, Jordan & Evermann 1898: 1520 (new combination).
Cichlasoma (Archocentrus) spilurum, Pellegrin 1904: 167 (new combination).
Cichlosoma spilurum, Regan 1905: 75 (unjustified emendation).
Archocentrus spilurus, Allgayer 1994: 15 (new combination).
Cryptoheros spilurus, Allgayer 2001: 14 (new combination).
Cichlasoma spinosissimum var. immaculata Pellegrin, 1904: 189 (junior synonym).
Cichlosoma immaculatum, Regan 1905: 77 (new status).
Lectotype. Herein designated as BMNH 1864.1.26.52, O. Salvin. Specimen 63 mm SL from the syntypic
series (Fig. 7). Lake “Isabel” (=Izabal), Guatemala.
Paralectotypes. BMNH 1864.1.26.53–55 (2), collected with lectotype.
Diagnosis. Autapomorphy (Schmitter-Soto, in press): first bar on side of body, Y-shaped or at least dor-
sally expanded, usually well-marked, arms continuous, rostral arm not strongly curved forward (see Günther
1867, plate 73, fig. 1). (Not to be confused with Y-shaped first bar of Amatitlania spp., in which rostral arm
not curved and caudal arm usually discontinuous.)
Description. D. XVII–XIX,9–11 (one specimen of 66 with 20 spines); A. VIII–X,7–9 (one specimen of
66 with 7 spines). Gill rakers on lower limb of first arch modally 6. Scale rows on cheek 4–6; scales from lat-

eral line to origin of dorsal fin 4.5–5.5; scales from lateral line to base of first dorsal-fin ray 2–2.5, sporadi-
cally 3; circumpeduncular scales modally 17 (additional meristic data appear in Table 3).
FIGURE 7. Cryptoheros spilurus, lectotype, BMNH 1864.1.26.52. Photo, BMNH, London.
FIGURE 8. Cichlasoma spinosissimum var. immaculata, syntype, MNHN 9846. Photo, C. Ferrara.
Largest examined specimen 80 mm SL. Body deeper than other Cryptoheros (52–56% of SL); head length
32–37% of SL; orbital diameter 28–32% of head length (further morphometric data appear in Table 4). Head
profile convex, straight, or concave. Teeth not embedded; pointed or bluntish, slightly labiolingually com-
pressed and retrorse, bicuspid (i.e. with a strong lingual cusp), occasionally incisor-like. Upper and lower
symphysial teeth subequal to adjacent teeth, not abruptly larger. Lips not medially narrow; lower lip squarish
or rounded at corner, its lower angle acute.

Pectoral and pelvic fins always reaching caudad beyond 3rd anal-fin spine. Filamentous rays of dorsal fin
extending to distal third of caudal fin or beyond. One or two lateral-line pored scales on caudal fin; subsidiary
pores always present, occasionally forming two-scale rows. Scales between dorsal and anal fin rays distally in
two rows, up to 16 scales long.
Genital papilla oval, with pigment on margins and basal half, rarely also on tip.
A faint vertical bar on head, starting behind orbits; no interorbital bands; a stripe from snout to eye; no
opercular spots. Eyes golden-green. No longitudinal stripe, but said to have a blue streak in life. Second bar on
side of body weaker dorsally; bar intensities otherwise not varying; 1st bar Y-shaped (see Diagnosis); 3rd to 7th
bars may extend onto base of dorsal fin. An ocellus sporadically present on spinous dorsal fin of females. Soft
dorsal and anal fins usually immaculate; sometimes with light dots discernible at bases. About 12–13 rows of
light spots on sides, smaller than scales; breast region coppery, yellow-orange, or pink-red. Axil of pectoral fin
somewhat darker, base of pectoral fin either white or same coloration as breast. Caudal blotch two-thirds or
more on peduncle, one third or less on caudal fin; the blotch crosses the lateral line, and usually forms a sad-
dle, i.e., blotch continuous across the dorsal part of the caudal peduncle but not across the ventral part.
Distribution. Endemic to rivers flowing into Lake Izabal, Guatemala (Fig. 5).
Remarks. Although earlier references (e.g., Schmitter-Soto 1998) have included this species in the Mexi-
can fauna, Kullander (2003) correctly considered the species not to be present in Mexico. Alleged records
from that country are based on Cr. chetumalensis (see below). Costa Rican and Panamanian specimens origi-
nally identified as “Cichlasoma spilurum” at UMMZ (190367 and 147222) are Cr. septemfasciatus and Cr.
altoflavus, respectively. The Nicaraguan records mentioned by Allgayer (2001) and Kullander (2003) might
be Cr. cutteri (see below), but I was unable to locate any specimen of the subgenus Cryptoheros from Nicara-
gua.
Cichlasoma immaculatum, originally described as a “variety” of C. spinosissimum by Pellegrin (1904; see
above) is actually a junior synonym of Cr. spilurus. The two syntypes (MNHN 9846, Fig. 8), although decol-
ored and partly descaled, are readily identifiable as Cr. spilurus based on meristic (e.g., A. IX, not A. XI;
scales between lateral line and first dorsal fin spine 4.5, not 5–6) and morphological grounds (e.g., the triple
coronal pore characteristic of Cryptoheros).
Cr. cutteri is removed from the synonymy of Cr. spilurus (see below).
Cryptoheros chetumalensis, new species

Figures 5, 9–10
Archocentrus spilurus (part. et non Günther), Schmitter-Soto 1998; Valtierra-Vega & Schmitter-Soto 2000; Miller et al.
2005 (misidentifications).
Holotype. ECOCH 5467, 63 mm SL (Fig. 9), M. Navarro-Mendoza, Jan. 15, 1988. Arroyo Aguadulce, a trib-
utary of the Río Hondo, at Sabidos, near Chetumal, Quintana Roo, Mexico.
Paratypes. ECOCH 1005 (12), 1465 (2), 1536 (1), 1559 (4), 1593 (3), 1693 (8), 1900 (13), 2328 (7),
UMMZ 210888 (9).
Diagnosis. No unique autapomorphy, but Cr. chetumalensis differs from the other two species in the sub-
genus by having the secondary pored scales on caudal fin not forming rows (vs. forming rows); rostral end of
maxilla convex (vs. notched or concave); first neural spine slanting rostrad instead of caudad; dorsal elements
between first two epineural spines three (vs. two); and a spinous anterodorsal process on first dorsal pterygio-
phore present (vs. absent).
Description. D. XVII–XIX,8–10 (one specimen of 26, with XX spines); A. VIII–X,7–9. Gill rakers on
lower limb of first arch modally 6. Scale rows on cheek 4–5; scales from lateral line to origin of dorsal fin

4.5–6; scales from lateral line to base of first dorsal-fin ray modally 2.5; circumpeduncular scales modally 17
(additional meristic data appear in Table 3).
FIGURE 9. Cryptoheros chetumalensis, holotype, ECOCH 5467. Photo, H. Bahena.
FIGURE 10. Cryptoheros chetumalensis live, from the outlet of Laguna Encantada into Río Hondo, ca. 10 km upstream
from Chetumal, Quintana Roo, Mexico. Photo, H. Bahena.
Maximum size observed, 97 mm SL. Body much less deep than in the other species of Cryptoheros, 42–
49% of SL; orbital diameter 28–32% of head length (further morphometric data appear in Table 4). Head pro-
file convex or straight, concave above orbits. Teeth moderately embedded; canine, pointed, slightly labiolin-
gually compressed, slightly retrorse, bicuspid. Upper and lower symphysial teeth subequal to adjacent teeth,
not abruptly larger. Lips may be medially narrow; lower lip squarish at corner, its lower angle acute.
Pectoral and pelvic fins always reaching caudad beyond 3rd anal-fin spine. Longest rays of dorsal fin
extending to mid-caudal fin or beyond. Subsidiary pored scales on caudal fin always present, but never form-

ing rows; scales between dorsal and anal fin rays distally in two rows, up to 11 scales long; in juveniles, no
scales between anal fin rays.
In specimens above ca. 45 mm SL, gut of simple type but with two intermediate loops; anal and medial
loops twist and directed dorsad; anal and anterior esophageal loops adjacent. Peritoneum only dorsally pig-
mented, mainly on posterior half; black melanophores on tan background. Genital papilla oval or tongue-
shaped; some melanophores on its base, margins and posterior side, or none.
A faint vertical bar on head; no interorbital bands, but this area dark in young; a stripe running from snout
to eye (sometimes diffuse); suborbital streak slightly curved, sharp-ended; no opercular spots. Eyes green,
grey, or bluish; sometimes golden. No longitudinal stripe. Bars on side of body, medially more intense, espe-
cially at scale edges; no conspicuous alternation of intensity, except that 3rd bar most intense and 2nd bar least
intense; first bar I-shaped, somewhat inclined on head; 1st and 2nd bars may coalesce medially as a humeral
blotch, and 3rd bar may display an oval spot. Ocellus absent from dorsal fin. Bases of soft dorsal and anal fins
usually darkened; sometimes with rows of light dots discernible. Abdomen whitish or greyish in life, not
wine-colored (Fig. 10). Axil of pectoral fin somewhat darker than breast; base of pectoral fin usually white.
Caudal blotch either more on peduncle than on fin, or entirely on peduncle, oval in shape, extending from dor-
sum to abdomen; edge of blotch diffuse, usually not saddled, coalescent or not with last bar.
Distribution. Belize (Belize River) and the Guatemalan Petén (Río Sarstún) north to Quintana Roo, Mex-
ico (as far north as Laguna Kaná, 19°30’N —ECOCH 1559) (Fig. 5).
Etymology. The type locality is about 10 km upstream from the river mouth at Chetumal, the city after
which the species is named. An adjective.
Cryptoheros cutteri (Fowler, 1932), n. comb.

Figures 5, 11
Cichlasoma cutteri Fowler 1932: 380 (original description).

Archocentrus cutteri, Allgayer 1994: 15 (new combination).
Cryptoheros spilurus (part. et non Günther), Allgayer 2001: 14 (misidentification).
Holotype. ANSP 53930, 112 mm SL (Fig. 11), C. B. Worth, Aug. 24, 1930. Río Lancetilla, Atlántida Dept.,
Honduras.
Paratypes. ANSP 53931–53933 (3, 38–108 mm SL), paratopotypes.
FIGURE 11. Cryptoheros cutteri, holotype, ANSP 53930. Photo, K. Luckenbill.

Diagnosis. Autapomorphy (Schmitter-Soto, in press): a wine-colored abdomen in life (in adults). Bars on
sides of body with alternating intensity, the second one much lighter than first and third. Further distinguished
from other species in the subgenus by arms of first epibranchial bone, which are divergent (vs. parallel);
supraoccipital crest undulating (vs. straight); ventral angle of articular obtuse (vs. right-angled); absence (vs.
presence) of an anteriorly directed pronounced convexity on ventral process of articular; procurrent rays of
caudal fin 2 (vs. 3–4).
Description. D. XVII–XIX,9–11; A. VIII–X,7–9 (one specimen of 60 with 11 spines). Gill rakers on
lower limb of first arch modally 7. Scale rows on cheek 4–6; scales from lateral line to origin of dorsal fin 4–
6 (contra Fowler 1932, who counted 7); scales from lateral line to base of first dorsal-fin ray 2.5–3.5; cir-
cumpeduncular scales modally 18 (further meristic data appear in Table 3).
Largest examined specimen, 112 mm SL. Body depth (44–54%, usually greater than 49% of SL). Head
length 32–43% of SL (further morphometric data appear in Table 4). Head profile convex or straight, concave
above orbits. Teeth embedded; pointed or bluntish, slightly labiolingually compressed and retrorse, bicuspid.
Upper symphysial teeth subequal to adjacent teeth, not abruptly larger; lower symphysial teeth may be
slightly shorter than adjacent teeth. Upper lip medially narrow; lower lip squarish or rounded at corner, its
lower angle acute.
Pectoral and pelvic fins always reaching caudad beyond 2nd anal-fin spine. Filamentous rays of dorsal fin
at least to mid-caudal fin. Subsidiary pored scales on caudal fin always present, occasionally forming two-
scale rows; scales between dorsal fin rays distally in two rows, up to 11 scales long.
In specimens from ca. 35 mm SL and larger, gut simple with two intermediate loops; medial and anal
loops twisted and directed dorsad. Peritoneum pigmented only dorsally (to half sides), mainly anteriorly; with
isolated blotches posteriorly. Genital papilla rounded, as long as broad, or oval, longer than broad or broader
than long; in some, the tip crenulated or notched; sunken; no pigment, except on margins, basal half, and pos-
terior (caudal) side.
No vertical bar on head; no interorbital bands, but postorbital–nape region somewhat dark; a stripe from
snout to eye, sometimes diffuse; suborbital streak straight, sharp-ended; no opercular spots, but opercle dark,
except on postorbital region. Eyes blue-greenish. No longitudinal stripe. Bars on sides of body, medially much
more intense; 1st bar I-shaped, somewhat inclined on head; intensity alternating: 1st, 3rd, 5th, 7th bars, especially
3rd, much more intense than 2nd, 4th, 6th; 2nd, 4th bars may be dorsally evanescent; 3rd bar and sometimes also 5th
and 7th may extend slightly onto base of dorsal fin. An ocellus may be present on spinous dorsal fin of mature
females. Soft dorsal and anal fins usually immaculate; sometimes light dots discernible at bases. About 13
rows of light spots on sides, smaller than scales; breast region yellowish-greenish or olive (or “wood brown”
according to Fowler [1932]). Axil of pectoral fin somewhat dark (part of 1st bar); base of pectoral fin yellow-
ish or white. Caudal blotch two-thirds or more on peduncle, across lateral line, usually forming a saddle.
Distribution. Atlantic Honduras (Ríos Celán, Lancetilla, Aguán, Jutiapa, and others) north to Guatemala
(Ríos Achuelo, Matasano, and others) (Fig. 5).
Remarks. Synonymized with Cichlasoma spilurum by Miller (1966) with no explicit justification. This
move was followed by Allgayer (2001), who added that the Honduran so-called “«Archocentrus» sp.
«Yojoá»” also belonged in Cr. spilurus, but acknowledged that “[p]lusieurs formes locales, notamment du
Honduras[,] sont connues.”
Fowler’s (1932: 381) figure is not accurate: it depicts the holotype as showing all the bars of approxi-
mately the same intensity, which is not the case (Fig. 11). The figure contains other errors, e.g. the size of the
scales on the breast, which are much smaller and numerous than shown. Fowler (1932) thought his new spe-
cies was “[r]elated to Heros octofasciatus” and felt that he needed to compare Cr. cutteri with R. octofasciata,
not with Cr. spilurus.
I was unable to examine specimens of the “Río Papaloteca” Archocentrus (Velasco 2001) from northern
Honduras, but nearby specimens (from Río Jutiapa, UMMZ 228665) are identifiable as Cr. cutteri.

Bussingius, n. subgen.
Type species. Cichlosoma septemfasciatum Regan, 1908.
Diagnosis. Synapomorphies (Schmitter-Soto, in press): genital papilla oval, in females opening V-shaped at
its anterior (rostral) end; upper symphysial teeth oval in section, labiolingually compressed (incisor-like), wid-
est medially (not so in Cr. septemfasciatus); an abdominal black blotch in mature females (not so in Cr.
sajica). Further distinguished from other two subgenera of Cryptoheros by dorsal-fin interradial scales distally
in one row (vs. two) and 2–7 scales long (vs. 11 or more).
Description. D. XVII–XVIII,9–11; A. VIII–IX (usually VII in Cr. sajica),7–9. Gill rakers on lower limb
of first arch 6–8. Scale rows on cheek 4–5; predorsal scales 10–16; pored lateral-line scales (not counting
scales overlapping between the two segments of the lateral line) 26–30; scales between lateral line and origin
of dorsal fin 3–4.5 (occasionally up to 5.5); scales between lateral line and first dorsal fin ray 1.5–2.5; cir-
cumpeduncular scales 16–18. A group of relatively small species of Cryptoheros, not exceeding 100 mm SL.
Lower jaw not protruding. Pectoral and pelvic fins always reaching caudad beyond 2nd anal-fin spine. No ver-
tical bar on head; no interorbital bands, snout usually somewhat dark; a suborbital streak usually present. Eyes
greenish-blue, body usually yellowish, orange or definitely yellow. Bars on sides of body, often diffuse, usu-
ally medially and dorsally more intense; first bar V-shaped or like an inverted triangle, its apex behind the
base of pectoral fin; mature females with a black ocellated blotch on dorsal fin (except Cr. sajica); lateral spot
oval or circular (not discernible from bar in Cr. sajica); caudal blotch often diffuse, ventrally triangular,
mainly on peduncle (except in Cr. sajica).
Distribution. Atlantic Panama and Atlantic and Pacific Costa Rica (Fig. 5).
Species composition. Five species: Cr. septemfasciatus, Cr. altoflavus, Cr. myrnae, Cr. nanoluteus, and
Cr. sajica.
Etymology. Gender masculine. Named in honor of William Bussing, arguably the most influential recent
ichthyologist in Costa Rica.
Cryptoheros septemfasciatus (Regan, 1908)

Figures 5, 12
Cichlosoma septemfasciatum Regan, 1908: 461 (original description).

Archocentrus septemfasciatus, Allgayer 1994: 15 (new combination).
Cryptoheros septemfasciatus, Allgayer 2001: 16 (new combination).
Lectotype. Herein designated as BMNH 1909.3.13.82 (Fig. 12), C. F. Underwood. Río Iroquois, Costa Rica.
The specimen, 69 mm SL, is the second largest individual in the type series, inasmuch as the largest syntype is
rather deformed.
Paralectotypes. BMNH 1909.3.13.83–91 (17, 60–100 mm TL), collected with lectotype.
Diagnosis. No unique autapomorphies, but the only species of Bussingius with symphysial teeth conical
rather than labiolingually compressed, and a caudal blotch both on fin and peduncle (vs. either exclusively on
the fin or on the peduncle). Further distinguished by bars on sides of body only medially (not dorsally) more
intense; anal creases 12–16, modally 14 (vs. 10–15, modally 11–13); and lower symphysial teeth bicuspidate,
with a small lingual cusp (also in Cr. altoflavus).
Description. D. XVII–XVIII,9–10 (one specimen of 20 with 8 rays); A. VIII–X,7–9. Gill rakers on lower
part of first arch modally 6; gill rakers trapezoidal, sometimes bifid. Scales strongly ctenoid. Predorsal scales
modally 13; pored lateral-line scales (not counting overlapping scales between the two segments of the lateral
line) modally 27; scales from lateral line to base of first dorsal-fin ray 2 or fewer (one specimen of 20 with
2.5); circumpeduncular scales 16–17, occasionally 18 (additional meristic data appear in Table 3).

FIGURE 12. Cryptoheros septemfasciatus, lectotype, BMNH 1909.3.13.82. Photo, BMNH, London.
Largest specimen examined, 82 mm SL (maximum size 100 mm SL [Conkel 1993]). Body depth 42–56%
of SL; head length, 30–36% of SL; orbital diameter 23–29% of head length (further morphometric data appear
in Table 4). Head profile convex in young; in adults, profile is concave-straight above orbits, convex on nape.
Teeth moderately embedded; conical, pointed, slightly labiolingually compressed. Upper symphysial teeth,
slightly shorter than adjacent teeth; lower symphysial teeth may also be slightly lower than adjacent teeth.
Upper lip medially narrow; lower lip square or rounded at corner, slightly tapering.
Pectoral fins always reaching caudad beyond 2nd anal-fin spine, and pelvic fins extending beyond 4th anal-
fin spine. Filamentous rays of dorsal fin reaching distal quarter of caudal fin or beyond. Up to two lateral-line
pored scales on caudal fin. Usually no subsidiary scales on caudal fin. Dorsal-fin interradial scale rows, not
imbricated (i.e. with no supplementary scales), up to 5 scales long.
Gut simple in juveniles, acquiring a double medial-loop in adults. Peritoneum only dorsally pigmented,
mainly anteriorly. Genital papilla with contours mostly parallel as seen from rostral face, or divergent, wider
before tip; pigmented overall except on tip.
No vertical bar on head; no interorbital bands; suborbital streak parallel to body axis, diffuse, blunt-ended;
no stripe from snout to eye, but snout somewhat dark; opercular spot indistinct or absent, but lower opercle
darker in some specimens. Eyes dark, greenish or bluish. No longitudinal stripe. Bars on side of body with no
dorsal intensifications; diffuse, except for the medial darkening; 1st bar diffuse, Y- or V-shaped (inversely tri-
angular), inclined on head; on the 3rd and often on the last bar the medial intensification forming a distinct oval
spot. Bars not extending onto dorsal fin. A dorsal blotch in females, starting on 8th or 9th spine, ending on 11th
or 12th. Soft dorsal and anal fins immaculate; their bases darkened. About 12 rows of light spots on sides,
smaller than scales, clearer on breast; breast, fins and throat yellowish-olive. Axil of pectoral fin either dusky
or with same coloration as breast; base of pectoral fin whitish. Caudal blotch more on peduncle than on fin,
occasionally entirely on peduncle; always ventrally pointed, its edge diffuse.
Distribution. Atlantic Costa Rica, Río Banano to Río San Juan, at the Nicaraguan border (Fig. 5).
Remarks. Regan (1908) considered this species to be “very close to C. spilurum.” Misidentifications are
common; many museum specimens of the other species in the subgenus are determined as Cryptoheros sep-

temfasciatus, especially records from Panama (e.g. ANSP 156820, reidentified as Cr. altoflavus) and southern
Costa Rica (e.g. ANSP 168312, reidentified as Cr. myrnae).
Cryptoheros altoflavus Allgayer, 2001

Figures 5, 13
Cryptoheros altoflavus Allgayer 2001: 16 (original description).

Archocentrus altoflavus, Kullander 2003: 616 (new combination).
Holotype. MNHN 2001-1163, 90 mm SL (Fig. 13), P. de Rham and J.-C. Nourissat, Apr. 22, 1998. Río
Cañaveral, Panama.
FIGURE 13. Cryptoheros altoflavus, holotype, MNHN 2001-1163. Photo, C. Ferrara.
Paratypes. MNHN 2001-1164 to 2001-1167 (6, 59–79 mm SL), 4 paratopotypes and 2 paratypes from
Río Caña, an affluent of Río Cañaveral.
Diagnosis. No unique autapomorphies. Breast, fins and throat yellowish (vs. greyish or strongly yellow in
other species of Bussingius); basal process on the largest first-arch gill-rakers; lower symphysial teeth bicusp-
idate, with a small lingual cusp (also in Cryptoheros septemfasciatus); distally two rows of interradial scales
on anal fin (also in Cr. nanoluteus); spots on opercle, part of a rather indistinct longitudinal stripe.
Description. D. XVII–XVIII,9–10; A. VIII–IX,7–9. Gill rakers on lower part of first arch 6; gill rakers
distally expanded, sometimes bifid. Scales strongly ctenoid. Predorsal scales modally 14; pored lateral-line
scales (not counting scales overlapping between the two segments of the lateral line) modally 28; scales from
lateral line to base of first dorsal-fin ray 2.5; circumpeduncular scales 17 (other meristic data appear in Table
3).
Largest specimen examined, 90 mm SL. The deepest-bodied species of subgenus Bussingius, depth 45–
56% of SL, mean 53% of SL; head length 31–36% of SL (this trait not diagnostic vs. Cr. nanoluteus, contra
Allgayer 2001); orbital diameter 22–35% of head length (further morphometric data appear in Table 4). Head
profile concave above orbits, steep, straight. Mouth terminal (contra Allgayer 2001, who found it “légèrement
rétrognathe”). Teeth moderately embedded; strongly labiolingually compressed, sides concave, tip triangular,

retrorse. Upper symphysial teeth, often one larger than the other; the smaller upper symphysial tooth not
abruptly larger than adjacent tooth; lower symphysial teeth subequal to adjacent teeth. Lips not narrow medi-
ally; lower lip slightly tapering or not, squarish at corner.
Pectoral fins always reaching caudad beyond 2nd anal-fin spine; pelvic-fin filamentous rays always reach-
ing beyond 4th and sometimes to 9th anal-fin spine. Filamentous rays of dorsal fin to distal quarter of caudal
fin. Two pored scales continuing lateral line on caudal fin, subsidiary scales present. Interradial scale rows of
dorsal and anal fins imbricated (i.e. with supplementary scales), up to 7 scales long.
Gut simple, with a secondary medial loop. Genital papilla tongue-shaped or rounded, notched, longer than
wide, wider at base, pigmented on base, sides, and posterior (caudal) face.
No vertical bar on head, but lower half of opercle darker; opercular spot usually distinct, forming part of
longitudinal stripe from opercle to pectoral-fin origin. Eyes greenish, greyish, or bluish. Bars on side of body
diffuse; 1st bar V-shaped with coalescent arms, like an inverted triangle, inclined on head; 2nd and 3rd intensi-
fied medially, all intensified dorsally; 5th and 6th more marked; medial intensification of 3rd bar an oval spot.
Bars not extending onto dorsal fin. Ocellus on dorsal fin of mature females starting on 6th or 7th spine, ending
between 10th and 13th spines. Soft dorsal and anal fins immaculate; their bases darkened. About 9 rows of light
spots on sides, smaller than scales; breast, fins and throat yellowish. Axil of pectoral fin somewhat dark, sur-
rounded by a lighter ring; base of pectoral fin whitish. Caudal blotch more on peduncle than on fin, occasion-
ally entirely on peduncle, across lateral line, saddled or diamond-shaped (always pointed ventrally), its edge
diffuse.
Distribution. In addition to the type locality, also known from Río Cricamola at Konkintu, Panama, a
new record that extends the range of the species westward (Fig. 5).
Remarks. Allgayer (2001) counted 15 abdominal + 15 caudal vertebrae, whereas I count 13+16 (also on
the published radiograph, ibidem); the present redescription is also at variance with the original description
with regard to other meristic data.
The collectors believed the species to be absent from the Río Cricamola “d’après les Indiens.” The Cri-
camola specimens of Cr. altoflavus approach Cr. nanoluteus in some characters, notably in body depth and
shape of caudal blotch.
Cryptoheros myrnae (Loiselle, 1997)

Figures 5, 14
Archocentrus myrnae Loiselle, 1997: 3 (original description).

Cichlasoma septemfasciatum (part. et non Regan), Bussing 1987: 219.
Cryptoheros myrnae, Allgayer 2001: 16 (new combination).
Holotype. AMNH 59079, 81 mm SL, W. A. Bussing. Río Cocolis, tributary of Río Sixaola, 3.5 km SE of Shi-
roles, Limón, Atlantic Costa Rica.
Paratypes. AMNH 59080 (8), LACM 44988-2, 44989-2 (30); UCR 144-2 (10); UMMZ 217739 (20)
(Fig. 14).
Diagnosis. No unique autapomorphies, but distinguished from other species of the subgenus Bussingius
as follows: upper symphysial teeth usually abruptly larger than adjacent teeth (vs. not abruptly larger); gill
rakers on first arch digitiform, blunt (vs. trapezoidal or bifid); a diffuse but complete longitudinal stripe, end-
ing in a tenuous blotch on the caudal peduncle (vs. on the fin); lateral spot circular (vs. oval); predorsal scales
modally 11 (vs. modally 12 or more).
Description. D. XVII–XVIII,9–11; A. VIII–IX,8–9. Gill rakers on lower limb of first arch modally 6; gill
rakers digitiform. Scales strongly ctenoid. Predorsal scales modally 11; pored lateral-line scales modally 27

(not counting scales overlapping between the two segments of the lateral line); scales from lateral line to base
of first dorsal-fin ray 1.5–2.5; circumpeduncular scales 16–18 (further meristic data appear in Table 3).
FIGURE 14. Cryptoheros myrnae, holotype, AMNH 59079. Photo, M. Stiassny.
Largest specimen examined, 68 mm SL (maximum size 80 mm SL: Kullander 2003). The most slender
species of Bussingius, depth 42–50% of SL; head length 33–37% of SL; orbital diameter 25–30% of SL (other
morphometric data appear in Table 4). Head profile straight above orbits, convex on nape. Teeth not embed-
ded; labiolingually compressed, the sides not concave. Upper symphysial teeth usually abruptly larger than
adjacent teeth; lower symphysial teeth subequal to adjacent teeth. Lips medially narrow; lower lip slightly
tapering or not, rather square at corner; often with fleshy hair-like protuberances on lower lip.
Pectoral and pelvic fins always reaching caudad beyond 3rd anal-fin spine. Filamentous rays of dorsal fin
extending to mid-caudal fin. One or two pored scales of lateral line continuing onto caudal fin; usually no sub-
sidiary scales present elsewhere between caudal-fin rays. Interradial scale rows on dorsal and anal fin imbri-
cated (i.e. with supplementary scales), up to 6 scales long.
Gut simple, with secondary loops. Peritoneum heavily pigmented dorsoanteriorly. Genital papilla globose
and distally pigmented in females, tip triangular and immaculate in males.
No interorbital bars, but snout darkened; suborbital streak, if visible, posteriorly sharp; opercular spot usu-
ally not distinct. Eyes metallic blue in life. Longitudinal stripe diffuse but complete, extending from orbit to
caudal fin. Bars on side of body diffuse, sometimes absent or incomplete; 1st bar V-shaped with coalescent
arms, like an inverted triangle, inclined on head; medial intensification of 3rd bar a circular spot. Bars not
extending onto dorsal fin. Ocellus on dorsal fin of mature females starting on 8th or 9th spine and ending
between 11th and 13th spine. Three hyaline rows of dots on soft dorsal fin, iridescent blue in life. About 6 rows
of light spots on sides (golden in life in thoracic region), smaller than scales; breast and throat orange in life; a
black triangular region ventral to pectoral fin in females. Axil of pectoral fin somewhat dark to black; base of
pectoral fin whitish. Caudal blotch mainly on peduncle, situated across lateral line; coalescent with last bar
and pointed ventrally.
Distribution. Atlantic Central America, from Río Guarumo, Panama, to Río Estrella, Costa Rica (Fig. 5).
Remarks. Cr. myrnae is replaced by Cr. septemfasciatus north of the Río Estrella (Bussing 1998). Speci-
mens of Cr. myrnae have, in the past, been used to demonstrate the supposed variability of Cr. septemfasciatus
(Bussing 1978).

Cryptoheros nanoluteus (Allgayer, 1994)
Figures 5, 15
Archocentrus nanoluteus Allgayer, 1994: 9 (original description).

Cryptoheros nanoluteus, Allgayer 2001: 16 (new combination).
Holotype. MNHN 1993-0260, 64 mm SL (Fig. 15), J.-C. Nourissat, Feb. 3, 1993. Río Guarumo, “Boca del
Toro” (Bocas del Toro), Chiriquí Grande, Panama.
FIGURE 15. Cryptoheros nanoluteus, holotype, MNHN 1993-0260. Photo, C. Ferrara.
Paratypes. MNHN 1993-0261–0263 (3, 52–62 mm SL).

Diagnosis. Unique autapomorphies (Schmitter-Soto, in press): one spot on opercle, not at angle; first bar
on sides of body, Y- or V-shaped, arms continuous and never coalesced, rostral arm curved forward. Further
distinguished from all other species of subgenus Bussingius by intense yellow, in life, on breast, fins and
throat (vs. yellowish or differently coloured); caudal blotch diffuse and entirely confined to caudal fin (vs.
caudal blotch at least partly on peduncle); sum of dorsal and anal-fin spines 28 (vs. 27 or fewer).
Description. D. XVII–XVIII,10–11; A. VIII–IX,8–9. Gill rakers on lower limb of first arch 6–7; gill rak-
ers digitiform, serrated. Scales strongly ctenoid. Predorsal scales 14; pored lateral-line scales (not counting
scales overlapping between the two segments of the lateral line) 28–29; scales from lateral line to base of first
dorsal-fin ray 2.5; circumpeduncular scales 16–17 (further meristic data appear in Table 3).
Largest specimen examined and maximum known length, 64 mm SL. Body moderately deep, 44–52% of
SL; head length 29–35% of SL; orbital diameter 27–32% of head length (further morphometric data appear in
Table 4). Head profile concave above orbits, convex on nape. Teeth moderately embedded; upper symphysial
teeth spatulate, sides concave, tip triangular, labiolingually compressed. Upper symphysial teeth not abruptly
larger, almost subequal to adjacent teeth; lower symphysial teeth subequal to adjacent teeth. Lips not medially
narrow; lower lip not tapering at corner, even sometimes expanded, squarish.
Pectoral and pelvic fins always reaching caudad beyond 4th anal-fin spine. Filamentous rays of dorsal fin
to distal third of caudal fin. One or two pored scales continuing lateral line on caudal fin, Subsidiary pores on
caudal fin, usually two on each scale. Dorsal- and anal-fin interradial scale rows, imbricated (i.e. with supple-
mentary scales), up to 8 scales long.

Genital papilla rather oval but wider at base, pigmented on base and sides.
Suborbital streak, if visible, anteriorly sharp; one opercular spot. Eyes greenish, bluish, greyish. Longitu-
dinal stripe incomplete, reaching just to base of pectoral fin. Bars on side of body intensified medially, rather
as series of spots, with a second series dorsally, darker posteriorly; 1st bar Y- or V-shaped, arms never coales-
cent, intensified medially and dorsally, rostral arm inclined on head; medial intensification of 3rd bar, discern-
ible as an oval spot; 4th bar weakest. Bars not extending onto dorsal fin. Ocellus on dorsal fin of mature
females starting on on 8th or 9th spine, ending between 10th and 14th spine. Dorsal and anal fins immaculate, just
bases darkened. About nine rows of light spots on sides, smaller than scales; breast, dorsal and anal fins and
isthmus intensely yellow in life. Axil of pectoral fin lighter than breast; base of pectoral fin whitish. Caudal
blotch on fin, two-thirds above lateral line, coalescent with last bar.
Distribution. Ríos Guarumo and Pejebobo, Atlantic Panama (Fig. 5).
Remarks. See Remarks for Cr. altoflavus. Allgayer (1994) considered it “assez proche de l’espèce A.
nigrofasciatus.”
Cryptoheros sajica (Bussing, 1974)

Figures 5, 16
Cichlasoma sajica Bussing, 1974: 30 (original description).

Archocentrus sajica, Allgayer 1994: 15 (new combination).
Cryptoheros sajica, Allgayer 2001: 16 (new combination).
Holotype. LACM 33902–1, 71 mm SL (Fig. 16), W. A. Bussing. A tributary of Río Sierpe, 2 km S of Palmar
Sur, Pacific Costa Rica.
FIGURE 16. Cryptoheros sajica, holotype, LACM 33902–1. Photo, R. Feeney and J. Seigel.
Paratypes. AMNH 58117 (3), LACM 2760 (1), 2928 (3), 4830 (61), 4853 (5), 33903-1–05-1 (50); UCR
69-7 (353), 111-19 (89), 112-16 (60), 114-12 (25), 163-3 (4), 164-7 (57), 165-3 (8), 166-3 (30), 172-3 (3),
173-6 (22), 175-3 (2), 179-1 (4), 250-3 (29), 251-7 (3), 300-5 (17), 309-1 (10), 311-4 (53), 380-8 (1), 393-11
(5), 757-6 (2); USNM 194247 (5), 211617-19 (50).

Diagnosis. No unique autapomorphies, but the only Bussingius with at least the third (main) lateral bar
uniformly wide (its width not uniform in other species); usually no caudal blotch, no lateral spot or medial
intensification of bars on side of body, no ocellus on dorsal fin, no abdominal blackening in mature females;
palatine arms subequal (vs. anterior longer); posteriad projection on ventroposterior angle of retroarticular
absent (vs. present); articular with a right angle ventrally (vs. obtuse); an anteriorly directed pronounced con-
vexity on ventral process of articular present (vs. absent); anal-fin spines modally 7 (vs. 8–9).
Description. D. XVII–XVIII,9–10; A. VI–VIII,7–8. Gill rakers on lower limb of first arch 6; larger gill
rakers bifid or at least distally expanded, trapezoidal, some digitiform. Scales moderately ctenoid. Predorsal
scales modally 14; pored lateral-line scales (not counting scales overlapping between the two segments of the
lateral line) 27–28; scales from lateral line to base of first dorsal-fin ray 2 or fewer; circumpeduncular scales
16–18 (additional meristic data appear in Table 3).
Largest specimen examined 53 mm SL (maximum size 90 mm SL: Kullander 2003). Body depth 46–49%
of SL; head length, 34–37% of SL; orbital diameter 28–31% of head length (further morphometric data appear
in Table 4). Head profile nearly straight to convex. Teeth not embedded; conical, labiolingually compressed.
Upper symphysial teeth not abruptly larger than adjacent teeth, but not subequal either; lower symphysial
teeth subequal to adjacent teeth. Upper lip normal or medially narrow; lower lip square squarish at corner,
often tapering.
Pectoral fins always reaching caudad beyond 2nd, pelvic fins beyond 3rd anal-fin spine. Filamentous rays of
dorsal fin to mid-caudal fin or beyond. Up to two lateral-line pored scales on caudal fin, subsidiary scales usu-
ally present. Dorsal-fin interradial scale rows, not imbricated (i.e. with no supplementary scales), up to 5 or 6
scales long.
Gut simple. Genital papilla tongue-shaped, oval, sunk; sometimes twice as long as broad; pigmented on
margins and tip.
No definite vertical bar on head; no interorbital bands; suborbital streak present; no stripe from snout to
eye; no opercular spot. Eyes blue with a golden rim (more noticeable in life). No longitudinal stripe. First bar
on side of body, rather V-shaped, but diffuse; 3rd bar best defined and uniformly wide; 3rd and 6th bars extend
onto dorsal fin, but there is no dorsal (female) spot. Hints of 2–3 spot rows on soft dorsal. About 11 rows of
spots on sides, smaller than scales and located on the scale rim; breast beige. Axil of pectoral fin with a dark
dorsal spot; base of pectoral fin paler than breast. Caudal blotch tenuous or absent, rather a dark region on
base of fin, not on peduncle.
Distribution. Río Parrita to Río Coloradito, Pacific versant of Costa Rica (Fig. 5).
Remarks. Without a cladistic analysis, Bussing (1974) hypothesized the species to be the Pacific sister-
group of the Atlantic Cr. septemfasciatus; my analysis could not support this assertion. However, several char-
acter states of Cr. sajica are plesiomorphic (Schmitter-Soto, in press), which suggests that the species could be
basal in Bussingius. The rest of the species in the subgenus are distributed in the Atlantic versant.
Amatitlania, new genus
Heros, Günther 1867: 601 (part.).

Cichlasoma, Jordan & Evermann 1898: 1525 (part.).
Archocentrus, Allgayer 1994: 15 (part.).
Cryptoheros, Allgayer 2001: 15 (part.).
Type species. Heros nigrofasciatus Günther, 1867, by original designation.
Diagnosis. Three strict synapomorphies (Schmitter-Soto, in press): first bar on side of body, Y-shaped, well-
marked, caudal arm discontinuous; bars from sides of body extending fully to the edge of dorsal and anal fins;

medial intensifications on 2nd and 3rd bars, sometimes also on 1st. Differs from most other heroine genera also
as follows: mouth terminal (also in Cryptoheros and Ar. multispinosus); spots on opercle part of vertical bar
on head (also in Caquetaia and H. nematopus), which extends fully across opercle (also in Rocio and Archo-
centrus); three procurrent rays on caudal fin (also in most Cryptoheros, Petenia, and Tomocichla); dentary
symphysial teeth lower than adjacent teeth (also in Rocio, Cr. panamensis, and Petenia—not so abruptly lower
as in Parachromis); premaxillary symphysial teeth abruptly larger than adjacent teeth (also in Rocio, Parach-
romis, Cr. myrnae, and Caquetaia); ascending premaxillary arm reaching only to anterior orbit rim (also in
Rocio); maxilla dorsally serrated (also in Cryptoheros and Tomocichla).
Description. D. XVII–XIX,7–10 (up to 11 in Am. kanna); A. VIII–X (to XI in Am. siquia and Am.
kanna),6–8 (to 9 in Am. siquia); first dorsal fin ray divided. Gill rakers on lower limb of first arch 5–8, total
gill rakers 7–11; gill rakers trapezoidal or bifid, with a basal process. Scales strongly ctenoid. Scale rows on
cheek 4–6; pored lateral-line scales (not counting scales overlapping between the two segments of the lateral
line) 25–29; scales from lateral line to first dorsal fin spine 4–5.5, rarely 3.5; scales between vent and interpel-
vic scale 6–12. Lower jaw external teeth 22–24. Anal creases modally 14.
Maximum size, less than 100 mm SL. Body rather oval, depth 43–58% of SL. Maxilla reaching only to
ventral rim of orbit, not to a vertical from anteriormost rim (except in some specimens of Am. nigrofasciata);
premaxillary dorsal process extending only to anterior rim of orbit; mouth terminal or lower jaw protruding
slightly; frenum present in lower lip; teeth moderately embedded, occasionally with a small lingual cusp,
lower symphysial teeth lower than adjacent teeth; pectoral and pelvic always reaching anal-fin origin and usu-
ally to 7th anal-fin spine; in Am. kanna and Am. siquia, pelvic fins may extend caudad to 10th anal-fin spine.
Origin of pelvic fin behind origin of dorsal fin. Caudal fin rounded-truncate. Interradial scale-rows in dorsal
and anal fins, up to 7–8 scales long (to 11 in Am. kanna). In gut, anal loop and rostral esophageal loop adja-
cent. Genital papilla oval, in females opening oval, not much crenulated. Seven bars on sides, first bar clearly
Y-shaped, medial intensifications on 2nd and 3rd bars, and sometimes also on 1st bar; no speckles on cheek; no
interorbital bars, or diffuse; a sharp vertical bar on head, across all of opercle and nape, coalescent with oper-
cular spots; no longitudinal stripe on body; no abdominal black blotch on females; caudal blotch entirely or
mostly on fin (sometimes obsolescent), across lateral line, not ocellated.
Distribution. From Panama (Atlantic) to Guatemala (Atlantic and Pacific) (Fig. 17).
Species composition. Four species: Am. nigrofasciata, Am. coatepeque, Am. kanna, Am. siquia.
Etymology. Gender feminine, derived from the type locality of the type species; “Amatitlán” means “a
place abundant in amate” in Nahuatl, “amate” being a kind of rustic paper made from the bark of Ficus peti-
olaris or F. indica (León-Portilla 1959).
Remarks. Regan (1908) suspected a relationship between Amatitlania and Rocio. The genus does show
putative synapomorphies with Rocio and Cryptoheros, but the cladistic analysis (Schmitter-Soto, in press) did
not find substantial support for such a clade.
Amatitlania nigrofasciata (Günther, 1867), n. comb.

Figures 17–18
Heros nigrofasciatus Günther, 1867: 601 (original description).

Cichlasoma nigrofasciatum, Jordan & Evermann 1898: 1525 (new combination).
Archocentrus nigrofasciatus, Allgayer 1994: 15 (new combination).
Cryptoheros nigrofasciatus, Allgayer 2001: 15 (new combination).
Lectotype. BMNH 1865.4.19.76, 64 mm SL (Fig. 18), O. Salvin. Designated herein from the largest speci-
men in the syntypic series. Lake Amatitlán, Guatemala (apparently not Lake Atitlán—see Remarks).

FIGURE 17. Distribution of the species in the genus Amatitlania. Squares, Am. nigrofasciata; circle, Am. coatepeque;
triangles, Am. kanna; stars, Am. siquia. A symbol may represent more than one collecting site.
Paralectotypes. BMNH 1865.4.19.77–78 (12), ZMB 6882 (1). There were several syntypes, not only one
(see Remarks).
Diagnosis. No unique autapomorphies, but the only species in the genus with two (vs. one) distal rows of
interradial scales on anal fin, and arms in the first epibranchial bone parallel (vs. divergent). In addition, pos-
terior end of dentigerous arm of dentary rounded or squarish (vs. triple-spined or bluntly pointed). Peritoneal
coloration uniformly dark (vs. not uniformly dark). Rostrad directed pronounced convexity on the ventral pro-
cess of the articular absent (vs. present). Also morphometric differences (body less deep) vs. Am. kanna and
Am. siquia, and coloration differences (4th bar not Y-shaped) vs. Am. coatepeque.
Description. D. XVII–XIX,7–9; A. VIII–X,6–7. Larger gill rakers elongated, rounded or pointed, curved
ventrad. Scales from lateral line to base of first dorsal-fin ray modally 2.5; circumpeduncular scales usually
17–19, modally 18; total vertebrae 27–28 (further meristic data appear in Table 3).
Largest specimen examined 88 mm SL (maximum size 100 mm SL, according to Kullander 2003). Body
depth 46–50% of SL, usually less than 48% of SL (further morphometric data appear in Table 4). Head profile
nearly straight on orbits to convex on nape. Teeth conical, pointed. Upper symphysial teeth abruptly larger
than adjacent teeth; lower symphysial teeth lower than adjacent teeth. Lips not medially narrow; lower lip
often tapering, corner dorsally rounded, ventrally angled.

FIGURE 18. Amatitlania nigrofasciata, lectotype, BMNH 1865.4.19.76. Photo, BMNH, London.
Pectoral fins always reaching caudad beyond 2nd anal-fin spine, pelvic fins extending beyond 3rd anal-fin
spine. Filamentous rays of dorsal fin to distal quarter of caudal fin. Up to two lateral-line pored scales on cau-
dal fin, subsidiary scales usually present. Dorsal- and anal-fin interradial scale rows arranged in one or two
rows, up to 8 scales long (contra Günther 1869, who found the soft dorsal and anal fins to have “scarcely any
scales on their base”).
Gut simple, usually shorter than standard length of fish. Peritoneum silvery. Genital papilla tongue-
shaped, somewhat oval-tubular, slightly notched, tip bluntly triangular, not sunken; pigmented on margins,
tip, and base on posterior (caudal) side.
Suborbital streak diffuse; stripe from snout to eye usually diffuse. Eyes bluish-green. Fourth bar on side of
body I-shaped. Ocellus on spinous dorsal fin of females absent (present in 0.3% of the specimens examined).
Breast olive. Axil of pectoral fin dark; base of pectoral fin usually definitely white. Caudal blotch present as a
bar on fin, not on peduncle.
Distribution. The range of this species is restricted as follows: Pacific slope, from Río Sucio, El Salvador
to Río Suchiate, Guatemala; Atlantic slope, from Río Patuca, Honduras to Río Jutiapa, Guatemala; in neither
slope to Panama, Costa Rica or even Nicaragua, as formerly considered. Those southern populations, and the
one isolated at Lake Coatepeque, El Salvador, belong to the three species described below (Fig. 17). Intro-
duced elsewhere, e.g. in the Río Balsas basin, central Mexico (Contreras-Balderas 1999).
Remarks. Günther (1867) mentioned both lakes, Amatitlán and Atitlán, as the type locality; however, the
original labels of the syntypes at BMNH mention only Lake Amatitlán, same as did Günther (1869). On the
other hand, Eschmeyer (2005) mentions just two British syntypes, but there are 13 specimens in the jar (plus
one skeleton), and Günther (1867) wrote: “numerous examples…were collected by Mr. Salvin.”
The extensive experimental (ethological, physiological, etc.) literature mentioning Archocentrus nigrofas-
ciatus or Cichlasoma nigrofasciatum should eventually be revised regarding the origin of the material used
(see Distribution). The same applies to the study of biological invasions by the “convict cichlid,” considered a
potential pest outside its natural range (Welcomme 1988).

Amatitlania coatepeque, new species
Figures 17, 19
Cichlasoma nigrofasciatum (part. et non Günther), Hildebrand 1925: 274 (misidentification).
Holotype. UMMZ 245584, 87 mm SL (Fig. 19), P. L. Clifton, Jan. 19, 1958. North shore of island in Lake
Coatepeque, El Salvador.
FIGURE 19. Amatitlania coatepeque, holotype, UMMZ 245584.
Paratypes. FMNH 12136 (10), island in Lake Coatepeque; UMMZ 181823 (18), 202805 (4), Santa Ana,
eastern coast of Lake Coatepeque at Monterrey.
Diagnosis. Unique autapomorphies (fig. 24h in Schmitter-Soto, in press): fourth bar on side of body Y-
shaped (formed by ventral coalescence of 4th and 5th bars) (vs. I-shaped); posterior end of dentigerous arm of
dentary triple-spined (vs. rounded), squarish or bluntly pointed; and a characteristic double medial-loop in gut
present (vs. absent). Differs from the other Amatitlania species also by posteriad projection at dorsal corner
present in lower lip (vs. absent), peritoneal pigmentation uniformly sparse (vs. uniformly dark or silvery), and
scales from lateral line to origin of dorsal fin 5–5.5 (vs. 5 or fewer).
Description. D. XVII–XIX,7–9; A. VIII–X,6–7. Scales from lateral line to first dorsal fin ray modally 2;
circumpeduncular scales usually 17–18; total vertebrae 28 (additional meristic data appear in Table 3).
Largest specimen examined 91 mm SL. The most slender Amatitlania, body depth 43–48% of SL, usually
less than 47% of SL (further morphometric data appear in Table 4). Head profile concave to straight on orbits
to convex on nape. Teeth conical. Upper symphysial teeth usually subequal to adjacent teeth. Upper lip medi-
ally narrow; lower lip tapering or not, rounded at corner, sometimes with a dorsal posteriad projection.
Pectoral and pelvic fins sometimes not reaching 1st anal-fin spine, but usually extending caudad beyond 2nd
anal-fin spine Filamentous rays of dorsal fin to end of caudal fin. One or two pores continuing lateral line on
caudal fin, subsidiary scales forming rows between other caudal-fin rays. Dorsal-fin interradial scales
arranged in one row, anal-fin scales in one or two rows, both up to 8 scales long.
Gut simple with a double medial-loop and the anal loop turned dorsad, always longer than standard length
of fish. Peritoneum with uniformly distributed melanophores among others, larger, fewer, aligned with ribs; in
young, only rostrally pigmented. Genital papilla elongated, contours parallel, end somewhat crenulated; pig-
mented on margins, base, tip, and posterior (caudal) side.
No suborbital streak discernible; stripe from snout to eye usually diffuse. Head bar darkening the whole
opercle, except for area rostral to lateral line. Eyes bluish-green. Bars on side of body more intensely black

than other Amatitlania; 4th bar Y-shaped, coalescent ventrally with 5th. Sometimes three rows of hyaline spots,
on tip of soft dorsal and base of soft anal fins. Breast olive-blackish. Axil of pectoral fin dusky; base of pecto-
ral fin whitish or with same coloration as breast. Caudal blotch present as a bar on fin, not on peduncle, nearly
all across the depth of the fin.
Distribution. Known only from the type locality, Lake Coatepeque, El Salvador (but see Remarks) (Fig.
17; photographs in Hildebrand 1925, his figs. 3 and 4).
Etymology. Coatepeque is the name of the lake where the species occurs, likely formed from the Nahuatl
cóatl=snake and tépetl=mount, hence “mount of the snake.” A noun in apposition.
Remarks. In his study of Salvadorian fishes, Hildebrand (1925) included specimens of Cichlasoma
nigrofasciatum from Lake Coatepeque; the species was “especially abundant among the rocks where, because
of the very clear water, it could be seen at a depth upward of 6 meters.” He noted that “some specimens are
much darker than others”, and that the natives called the light variety “plateada” (silver) and the dark one
“negra” (black). Am. coatepeque has much more intense bars on side of body than other Amatitlania species.
The population at Lake Chalchuapa, El Salvador (FMNH 12139), although identifiable on other respects
as Am. nigrofasciata, displays in 2 of 7 individuals the Y-shaped 4th bar characteristic of the new species. The
site is near Lake Coatepeque, and may have been connected to it in the past. The above mentioned “plateada”
and “negra” forms coexist at Chalchuapa (Hildebrand 1925).
Amatitlania kanna, new species

Figures 17, 20
Archocentrus nigrofasciatus (part. et non Günther), Allgayer 1994: 12 (misidentification).

Cryptoheros cf. nigrofasciatus, Allgayer 2001: 18 (preliminary detection).
Holotype. FMNH 59243, 83 mm SL (Fig. 20), E. H. Behre and J. Chambers, Feb. 2, 1923. San San swamp,
Atlantic Panama.
Paratypes. BMNH 1925.3.6.119 (1), 1925.3.6.120–121 (2), 1925.3.6.122–123 (2), 1925.3.6.126 (1),
FMNH 59240 (2), 59241 (1), 59242 (1), 116465 (1), MHNG 2646.78 (2),UMMZ 145716 (1).
FIGURE 20. Amatitlania kanna, holotype, FMNH 59243. Photo, P. Willink.

Diagnosis. No unique autapomorphies, but differs from all other Amatitlania by secondary caudal pores
absent (vs. present), peritoneum pigmented dorsolaterally (vs. uniformly), and quadrate bone wider than long
(vs. not wider than long). Several features shared with Am. siquia (body relatively deeper, fewer circumpe-
duncular scales), some of them synapomorphies (Schmitter-Soto, in press): gill-rakers on first arch often bifid
(vs. trapezoidal); caudal edge of urohyal sigmoid (vs. concave); anterodorsad spine on first dorsal pterygio-
phore present (vs. absent). However, it differs from its sister species as follows: dorsal-fin interradial scales
distally in two rows (vs. one); posterior edge of mesethmoid rounded (vs. straight-irregular); gut simple (S-
shaped, not folded anal and medial loops touching) (vs. shaped like an S folded ventrorostrally in adults, anal
and medial loops not touching); and caudal blotch completely on fin (vs. partly on peduncle).
Description. D. XVII–XIX,8–10 (one specimen of 19 with 11 dorsal fin rays); A. IX–X,7–8. Gill rakers
trapezoidal or bifid. Scales from lateral line to first dorsal-fin ray modally 2; circumpeduncular scales usually
16–17, modally 16; total vertebrae 27 (further meristic data appear in Table 3).
Largest specimen examined, 83 mm SL. The deepest-bodied Amatitlania, body depth 48–58% of SL, usu-
ally greater than 50% of SL (further morphometric data appear in Table 4). Head profile concave on orbits,
convex on nape. Teeth conical, pointed, slightly retrorse. Upper symphysial teeth abruptly larger than adjacent
teeth. Upper lip medially narrow; lower lip often with fleshy, hair-like papillae, not tapering nor squarish at
corner, its lower angle acute.
Pectoral fins always reaching caudad beyond 2nd anal-fin spine pelvic fins always extending beyond 5th
and often to 10th anal-fin spine. Filamentous rays of dorsal fin reaching to distal third of caudal fin. One or two
pores continuing lateral line on caudal fin, no subsidiary scales. Dorsal-fin interradial scales arranged distally
in one or two rows, up to 12 scales long; anal-fin scales usually in one row, up to 9 scales long.
Gut simple, usually slightly shorter than standard length. Peritoneum pigmented only dorsally, less so on
upper sides. Genital papilla oval, but often distally expanded; uniformly pigmented on base.
Suborbital streak present; stripe from snout to eye usually well defined. Head bar interrupted in an area
rostral to lateral line. Eyes blue with a golden rim. Bars on side of body sharp, medially and dorsally more
intense; 4th bar I-shaped. No dots discernible on soft dorsal and anal fins. Breast olive. Axil of pectoral fin
with same coloration as breast or dark, often with a dorsal spot; base of pectoral fin whitish. Caudal blotch on
fin, across lateral line; often saddled.
Distribution. From rivers Cañaveral, Cricamola, Sixaola, and other localities in Atlantic Panama (Fig.
17).
Etymology. Greek κάννα, meaning “a reed”, Río Cañaveral (= reedbed) being the first locality where the
species was detected (see Remarks). A noun in apposition.
Remarks. The species was first recognized as distinct by Allgayer (2001: 18), who called his specimens
from Río Cañaveral “Cryptoheros cf. nigrofasciatus.”
Amatitlania siquia, new species

Figures 17, 21
Cichlasoma nigrofasciatum (part. et non Günther), Bussing 1987: 213 (misidentification).

Archocentrus nigrofasciatus (part. et non Günther), Bussing 1998: 346 (misidentification).
Holotype. UMMZ 245585, 61 mm SL (Fig. 21), M. J. Allen, Aug. 10, 1932. A stream flowing out of the for-
est into Río Siquia, 7 miles N of Rama, Atlantic Nicaragua.
Paratypes. ANSP 88241 (3), 67475 (4), 140692 (1), BMNH 1925.3.6.124–125 (2), FMNH 7733 (1),
MHNG 2160.60 (1), UMMZ 145720 (2), 166474 (1), 196939 (1), 196948 (25), 196954 (7), 213938 (6).
Diagnosis. Unique autapomorphies (figs. 24e and 6d in Schmitter-Soto, in press): gut simple (S-shaped),
but folded ventrorostrally, anal loop and medial loop not touching (vs. simple, not folded); peritoneum only

rostrally pigmented (vs. not only rostrally pigmented). Further distinguished from all other Amatitlania as fol-
lows: caudal blotch about two-thirds on peduncle and one-third on fin (vs. completely on fin); posterior edge
of mesethmoid straight-irregular (vs. rounded); scales from lateral line to base of first dorsal fin ray modally
1.5 (vs. modally 2 or more); caudal vertebrae modally 14 (vs. modally 13).
FIGURE 21. Amatitlania siquia, holotype, UMMZ 245585.
Description. D. XVII–XVIII,8–10; A. IX–X,7–8. Gill rakers bifid in larger specimens, sometimes ser-
rated, or at least distally expanded, compressed. Scales from lateral line to base of first dorsal-fin ray always
1.5; circumpeduncular scales usually 15–17, modally 16; total vertebrae 26–27 (additional meristic data
appear in Table 3).
Largest specimen examined, 79 mm SL. A deep-bodied Amatitlania, body depth 48–55% of SL, usually
greater than 49% of SL (further morphometric data appear in Table 4). Head profile straight to convex. Teeth
conical, pointed, slightly or not labiolingually compressed. Upper symphysial teeth abruptly larger than adja-
cent teeth. Upper lip medially narrow; upper angle of lower lip at corner much rounded, lower angle acute,
tapering ventrad.
Pectoral fins always reaching caudad beyond 3rd, pelvic fins always beyond 5th anal-fin spine and often to
8th anal-fin spine. Filamentous rays of dorsal fin to distal third of caudal fin. Up to three pored scales continu-
ing lateral line on caudal fin, isolated subsidiary scales on larger specimens. Dorsal- and anal-fin interradial
scales arranged in one row, up to six scales long.
Gut length may reach 188% of SL. Genital papilla oval, but often medially constricted, peanut-shaped,
longer than wide, with a long protuberance on tip; isolated melanophores on base and sides.
Suborbital streak present, at least in young; stripe from snout to eye usually well defined. Interorbital
bands sometimes not so diffuse. Head bar more intense dorsally. Eyes bluish, greenish, greyish. Bars on side
of body ventrally more obsolescent and coalescent than in other species; 4th bar I-shaped. Dots sometimes dis-
cernible on bases of soft dorsal and anal fins. Breast olive. Axil of pectoral fin with a dorsal dark spot; base of
pectoral definitely white. Caudal blotch two-thirds on fin, more dorsal than ventral to lateral line, never to
ventral edge, oval.
Distribution. Both coasts of Costa Rica (from tributaries to the Golfo de Nicoya on the Pacific coast and
Río Parismina on the Atlantic) to Nicaragua (including the Great Lakes and the Mosquitian basins); also north
to Atlantic Honduras (Río Yeguaré) (Fig. 17).

Etymology. Siquia is the name of the river chosen as type locality. The name means “avocado” in the
Miskito dialect Ulwa (Salamanca 2002). A noun in apposition.
Rocio, new genus
Heros, Regan 1903: 417 (part.).

Cichlasoma, Pellegrin 1904: 167 (part.).
Parapetenia, Allgayer 1989: 26 (part.).
‘Cichlasoma’, Kullander 1996: 151 (incertae sedis).
Archocentrus, Schmitter-Soto 1998: 154 (part.).
Nandopsis, Burgess 2000: 48 (part.).
Type species. Heros octofasciatus Regan, 1903, by original designation.
Diagnosis. Three strict synapomorphies (figs. 2e, 6e and 7e in Schmitter-Soto, in press): main gill rakers on
first arch with a mediad projection at base; posterior edge of mesethmoid with an indentation and a posteriad
spine; posterior edge of supraoccipital crest undulated, with a deep concavity. Many homoplastic synapomor-
phies: first dorsal fin ray not divided; ascending premaxillary process to anterior orbit rim (also in Amatitla-
nia); lower symphysial teeth usually bicuspidate (also in Cryptoheros); narrowest point of dentigerous arm of
premaxilla at caudal tip; scale rows between lateral line and base of first dorsal fin ray modally 3.5 or more
(also in Ar. spinosissimus); first bar on side of body Y-shaped but diffuse (also in some Archocentrus, some
Cryptoheros, and Tomocichla); longitudinal stripe from orbit to lateral blotch (also in Ar. spinosissimus and P.
loisellei).
Description. D. XVII–XIX,8–11; A. VIII–IX (rarely VI, VII, or X),7–9 (rarely 6); pectoral 14–16. Lower
gill rakers 6–8 (rarely 9); main gill-rakers with low ridges directed posteriad. Scales weakly ctenoid (almost
cicloid) to strongly ctenoid. Interradial scale rows in dorsal and anal fins, up to 8 scales long, usually in one
row. Scales from lateral line to first dorsal fin spine 3.5–5.5; pored lateral-line scales (not counting scales
overlapping between the two segments of the lateral line) 27–31, prolonged by up to two pored scales on cau-
dal fin; scales from lateral line to first dorsal fin ray 2.5–4, rarely 4.5; scales between vent and interpelvic
scale 7–15; circumpeduncular scales 17–22. Procurrent rays of caudal fin two; anal creases modally 14; verte-
brae 15–17 caudal, 28–30 total. Said to reach 250 mm SL (Page & Burr 1991), usually much smaller. Body
rather oval in young, almost Archocentrus-like; more elongate in adults, almost Parachromis-like. Premaxil-
lary dorsal process to anterior rim of orbit. Lower jaw extending beyond upper jaw; frenum present in lower
lip; lips not medially narrow. Teeth moderately embedded; upper symphysial canine or conical, unicuspid or
with a lingual cusp, abruptly larger than adjacent teeth; lower symphysial teeth lower than adjacent teeth. Ori-
gin of pelvic fin behind origin of dorsal fin. Caudal fin rounded-truncate. Gut simple all life long. Peritoneum
almost immaculate, except for some melanophores dorsally on anteriormost ribs. Genital papilla oval, longer
than wide, tip bluntly triangular in males, in females opening oval and not much crenulated, sunken; pig-
mented just on base, on margins, or not at all. Two usually well-marked interorbital bands; a sharp vertical bar
on head, across all of opercle and nape, opercular spots part of it; cheeks, opercular and gular regions often
speckled. A longitudinal stripe from snout through orbit and opercle to lateral spot on 3rd or 4th lateral bar; 8
bars on sides (sometimes rather indistinct), first bar diffusely Y-shaped, 4th and sometimes also 3rd bar medi-
ally more intense. Bars on side of body not extending onto dorsal and anal fins; 3–6 rows of dots on bases of
soft dorsal and anal; no dorsal ocellus or abdominal blotch in mature females; ocellated spot on caudal fin,
completely dorsal to lateral line.
Distribution. Atlantic versant, from Honduras to Mexico, as far north as Veracruz (Fig. 22).
Species composition. Three species: R. octofasciata, R. ocotal, R. gemmata.

Etymology. The gender is feminine: Rocío is my wife’s name. The Spanish word “rocío” means “morn-
ing dew”, an image evoked by the resplendent spots on cheek and sides of some species, notably R. gemmata.
FIGURE 22. Distribution of the species in the genus Rocio. “+” signs, R. octofasciata; circle, R. ocotal; squares, R. gem-
mata. A symbol may represent more than one collecting site.
Rocio octofasciata (Regan, 1903), n. comb.

Figures 22–24
Heros cyanoguttatus (part. et non Baird & Girard), Evermann & Goldsborough 1902: 157 (misidentification).
Heros octofasciatus Regan, 1903: 417 (original description).
Cichlasoma octofasciatum, Meek 1904: 218 (new combination).
Cichlasoma hedricki Meek, 1904: 208 (junior synonym).
Cichlosoma biocellatum Regan, 1909: 234 (junior synonym).
Parapetenia octofasciata, Allgayer 1989: 26 (new combination).
‘Cichlasoma’ octofasciatum, Kullander 1996: 151 (incertae sedis).
Archocentrus octofasciatus, Schmitter-Soto 1998: 154 (new combination).
Nandopsis octofasciata, Burgess 2000 : 48 (new combination [as octofasciatum]).
Holotype. MHNG 665.55, 39 mm SL (Fig. 23), A. C. Buller, Feb. 1, 1866. “Río de Sarabia” (Cosamaloapan,
according to the original label), Coatzacoalcos drainage, Veracruz, Mexico. No paratypes, but originally
mixed, unlabeled, in same jar with eight more specimens (see Remarks).
Diagnosis. No unique autapomorphies. Spots on sides smaller than scales, aligned in ca. 15 regular series
(vs. not clearly aligned); abdomen predominantly whitish or greyish in life (also in R. gemmata, vs. reddish in

R. ocotal); ventral angle of articular, acute (vs. right); first neural spine oriented rostrad (vs. caudad); cir-
cumpeduncular scales as few as 17 (vs. always more than 19); distance from caudal esophageal loop in gut to
esophagus always greater than 24% gut length (vs. less than 16%).
FIGURE 23. Rocio octofasciata, holotype, MHNG 665.55. Photo, C. Ratton.
Description. D. XVII–XIX,8–10, modally 10 (7 of 174 with 11 dorsal fin rays); A. VIII–IX,7–9, modally
8 (2 of 177 with 6 or 7 anal-fin spines); pectoral 14–16. Gill rakers rounded, distally expanded or trapezoidal,
sometimes bifid and even trifid, sometimes serrated. Scale rows on cheek modally 6 (4–7, 12 of 68 specimens
with 7 scale rows); predorsal scales modally 12; pored lateral-line scales (not counting scales overlapping
between the two segments of the lateral line) 27–30; scales from lateral line to origin of dorsal fin 4–5.5;
scales from lateral line to base of first dorsal-fin ray 3–4 (further meristic data appear in Table 3).
Largest specimen examined, 217 mm SL, but reported to 250 mm SL (Page & Burr 1991). Body usually
slender, deeper in young (39–51% of SL). Head length 33–42% of SL; orbital diameter usually 21–25% of
head length (up to 31% of head length in juveniles—further morphometric data appear in Table 4). Head pro-
file convex, straight, concave on orbits. Maxilla reaching only a horizontal line, not a vertical, from orbit. Cor-
ners of lower lip slightly tapering, if at all.
Pectoral and pelvic fins nearly always reaching caudad beyond 1st or 2nd anal-fin spine (not reaching in 2
of 127 specimens). Filamentous rays of dorsal fin to mid-caudal fin. Scales between dorsal and anal fin rays in
one row, up to 6 scales long.
Gut simple, anal and anterior esophageal loops adjacent; gut length shorter or subequal to standard length
of fish. Genital papilla thick, cylindrical, notched or triangular-tipped, sometimes sunk, longer than broad,
medially swollen, somewhat crenulate at tip, sometimes rather vase-shaped; pigmented just in basal half and
margins, if at all.
Stripe from snout to eye diffuse or absent; suborbital streak narrow, with pointed ends (not always visible
in preserved specimens). Eye color varied: yellowish, greyish, bluish, coppery, reddish. Bars on sides some-
times diffuse, sporadically dorsally or medially coalescent. Lateral blotch oval to squarish, joining 3rd and 4th
bars, sometimes not discernible from longitudinal stripe. About 15 rows of light spots on sides, centered in
each scale, not always visible; breast region olive. Axil of pectoral fin with same coloration as breast or
dusky; base of pectoral fin paler than surrounding region. For life colors, see Fig. 24.

FIGURE 24. Rocio octofasciata live. Locality: Arroyo Tres Garantías, Quintana Roo, Mexico. Photo, H. Bahena.
Distribution. From Río Ulúa, Honduras (Lee et al. 1980), to tributaries of Río Actopan, Veracruz, Mex-
ico (Greenfield & Thomerson 1997) (Fig. 22). Introduced in several localities in northern Veracruz (Obregón
et al. 1994).
Remarks. Contra Regan’s (1903) original designation (and S. Fisch–Müller’s opinion, pers. com.), Mah-
nert (1976: 44) treated the nine specimens originally in MHNG 665.55 as syntypes. However, there is little
doubt that the largest individual is the holotype. The confusion was likely created by Regan (1904, 1905) him-
self, when he added material to his successive redescriptions, somewhat more accessible than the original
description.
Cichlosoma biocellatum, based on an aquarium specimen, is a synonym, its type locality (“Río Negro at
Mañaos [sic], Brazil”) certainly in error.
Cichlasoma hedricki was synonymized by Regan (1905), barely one year after Meek’s (1904) description,
without explicit reasons. The action was followed by most authors. Topotypes of C. hedricki (from the upper
Papaloapan) tend to be deeper-bodied than other R. octofasciata, and the presence of a prominent lingual cusp
in their symphysial teeth (fig. 15a in Schmitter-Soto, in press) tends to be more constant, but the forms merge
imperceptibly towards the type locality of R. octofasciata in the Coatzacoalcos, the drainage immediately to
the south of Papaloapan.
In print, Meek (1904) called the type locality of C. hedricki “Obispo, Veracruz;” however, his manuscript
label in the paratype jar at UMMZ (176671) reads “Río Obispo, Oaxaca.” The eight paratypes in this lot
(labelled as such by Meek himself) should probably be added to the 44 specimens mentioned by Eschmeyer
(2005); however, on the other hand, the original description mentions only one “type,” thus rendering the lit-
erature (and paratype holdings at UMMZ and FMNH) in error.
Rocio ocotal, new species

Figures 22, 25
Cichlasoma (Parapetenia) sp., Miller 1957: 241 (preliminary detection).

Cichlasoma octofasciatum (part. et non Regan), Stawikowski & Werner 1998 (misidentification).

Holotype. UMMZ 245583, 70 mm SL (Fig. 25), R. A. Paynter, Jul. 21, 1954. Laguna Ocotal, “Lacandona
region,” Chiapas, Mexico.
FIGURE 25. Rocio ocotal, holotype, UMMZ 245583.
Paratypes. UMMZ 171140 (5, 49–96 mm SL). Paratopotypes.

Diagnosis. No unique autapomorphies, but distinguished from the other Rocio species as follows: abdo-
men reddish in life (Miller 1957, Stawikowski & Werner 1998) (vs. whitish–greyish); pelvic fins usually fall-
ing short of anal-fin origin (vs. nearly always reaching caudad beyond 1st or 2nd anal-fin spine); lingual cusp in
lower symphysial teeth absent (vs. usually present); isolated secondary pores (i.e., in addition to the pored
scales on the extended caudal fin) present (vs. none or sporadic); spots on scales on side of body absent (vs.
present); dentary pores 4 or 5 (vs. always 4) (fig. 16 in Schmitter-Soto, in press).
Description. D. XVIII–XIX,9–10; A. VIII–IX,6–9, modally 7; pectoral 16. Gill rakers elongated, pointed
or digitiform. Scales cycloid on shoulder (rostrad to origin of dorsal fin, dorsad to base of pectoral fin). Scale
rows on cheek modally 5; predorsal scales modally 14; pored lateral-line scales (not counting scales overlap-
ping between the two segments of the lateral line) 29–30; scales from lateral line to origin of dorsal fin 3.5–
4.5; scales from lateral line to base of first dorsal-fin ray 2.5–3.5 (additional meristic data appear in Table 3).
Largest specimen examined, 96 mm SL. Body usually slender, depth 41–46% of SL. Head length 35–41%
of SL; orbital diameter 21–23% of head length (further morphometric data appear in Table 4). Head profile
convex, straight above orbits. Maxilla reaching only a horizontal line, not a vertical, from orbit. Corners of
lower lip straight, not curved downward, tapering, rounded. Sometimes five instead of four dentary pores.
Pectoral and pelvic fins often falling short of first anal-fin spine. Filamentous rays of dorsal fin to mid-
caudal fin. Scales between between dorsal fin rays, distally in two rows, up to 5 scales long.
Gut simple, anal and anterior esophageal loops adjacent; gut length shorter than standard length of fish;
distance from last loop in gut to esophagus always less than 16%. Genital papilla a little longer than broad,
rounded, sunk, cylindrical, erect, somewhat crenulate at tip; pigmented just in basal half and margins.
Stripe from snout to eye diffuse or absent. Eyes reddish. Bars on sides rather indistinct, especially in two
almost completely black specimens. Lateral blotch rather rounded. 12–16 rows of light spots on sides, cen-
tered in each scale, not always visible; breast region bronze-yellowish or blackish. No clear dots or streaks on
anal fin. Axil of pectoral fin with same coloration as breast or dusky; base of pectoral fin pale; isolated melan-
ophores on pectoral ray inner (posterior) bases. Abdomen reddish in life (Stawikowski & Werner 1998).
Distribution. Probably endemic to Laguna Ocotal, a rather isolated, highland water body in the Lacantún-
Usumacinta drainage, Chiapas, Mexico (Fig. 22—pictures in Miller 1957).

Etymology. From the Spanish “ocotal,” meaning “an ocote forest,” “ocote” being a species of Pinus; the
name of the lake where the species lives. A noun in apposition.
Remarks. The view that the Ocotal population is in fact a new species was already expressed nearly 50
years ago (Miller 1957). This is one of the many species in several families that the late Dr. Miller detected as
new, but did not get to describe (Hendrickson et al. 2002).
Rocio gemmata Contreras-Balderas & Schmitter-Soto, new species

Figures 22, 26
Archocentrus aff. octofasciatus, Schmitter-Soto 1998: 156 (preliminary detection).
Holotype. ECOCH 4054, 64 mm SL (Fig. 26), the author, Jun. 11, 1999. Nameless cenote 12 km N of Leona
Vicario, Quintana Roo, Mexico.
FIGURE 26. Rocio gemmata, holotype, ECOCH 4054. Photo, H. Bahena.
Paratypes. ECOCH 1468, 3145, collected at the same karstic sinkhole by the author and H. C. Gamboa-
Pérez, and UANL 15046 (4), collected at Laguna Leona Vicario by S. Contreras-Balderas, co-discoverer of
the species.
Diagnosis. Unique autapomorphies (fig 14e in Schmitter-Soto, in press): spots on sides, larger than scales
and not clearly aligned (vs. smaller than scales and rather well-aligned); stripe from snout to eye interrupted
(vs. continuous); quadrate bone with a spine (vs. without a spine). In addition, maxilla reaching both a vertical
and a horizontal line from orbit (vs. just to the ventral rim); cheek-scale rows modally 7 (vs. 6 or fewer); inter-
radial scales on dorsal fin in one row (vs. distally in two rows); dorsal and anal fins not bearing filaments (vs.
bearing filaments); anal and medial gut-loops not adjacent, well separated by the stomach and the liver (vs.
always adjacent); rostral end of maxilla notched or at least concave (vs. convex, with no notch); caudal ocel-
lus blue in life (vs. white).
Description. D. XVIII,9–10; A. VIII–IX,7–8; pectoral 15–16. Gill rakers trapezoidal, bifid in larger spec-
imens, their posterior ridge serrated. Scale rows on cheek 7; predorsal scales 14–15; pored lateral-line scales
(not counting scales overlapping between the two segments of the lateral line) 28–30; scales from lateral line
to origin of dorsal fin 4–4.5; scales from lateral line to base of first dorsal-fin ray 3.5 (further meristic data
appear in Table 3).

The smallest species of the genus Rocio; largest specimen examined, 70 mm SL. Body depth 41–46% of
SL. Head length 35–41% of SL; orbital diameter 25–30% of head length (further morphometric data appear in
Table 4). Head profile convex, straight above orbits. Lower lip at corner of mouth slightly curved downward,
tapering.
Pectoral fins often falling short of first anal-fin spine, pelvic fins always reaching at least first anal-fin
spine. Dorsal and anal fins not bearing filaments. Just one row, up to 4 scales long, of interradial scales on
both dorsal and anal fin rays.
Gut simple, anal and anterior esophageal loops not adjacent; gut length may be greater than standard
length; distance from last loop in gut to esophagus always less than 16% gut length. Genital papilla rounded,
may be smaller than creased area of anus; pigmented only on base.
Suborbital streak wide, blunt-ended (usually not visible in preserved specimens). Stripe from snout to eye
diffuse, rather a darkening at rim of orbit. Large, iridescent, metallic green-blue speckles on cheek in life,
turning blackish in ethanol. Eyes bronze-bluish. Bars on sides sometimes doubled medially; lateral blotch
oval, sometimes ocellated. About six unordered rows of spots on sides, larger than scales; breast bronze-
blackish with green-blue tinge in life. Dots on fins large, dark blue. Axil of pectoral fin with a dorsal spot;
base of pectoral fin whitish.
Distribution. Endemic to cenotes and small inland lakes in northern Quintana Roo, eastern Yucatan Pen-
insula, Mexico (Fig. 22).
Etymology. Latin gemmata, meaning “bejeweled,” in reference to the large, bright green and blue cheek
and opercle spots in life. An adjective.
Remarks. The authorship of this species is joint with S. Contreras-Balderas, who independently collected
and recognized it as distinct.
Genus Hypsophrys Agassiz
Hypsophrys Agassiz, 1859: 408 (original description).

Heros, Günther 1864: 153 (part.).
Neetroplus Günther, 1867: 603 (junior synonym).
Cichlasoma, Pellegrin 1904: 167 (part.)
Copora Fernández-Yépez, 1969: 3 (junior synonym).
Type species. Hypsophrys unimaculatus Agassiz, 1859 = Heros nicaraguensis Günther, 1864 (see Remarks).
Diagnosis. No strict synapomorphies, but unique characters in combination: total number of gill rakers on
first arch modally 11 (also in Parachromis); posterior end of dentigerous arm of dentary rounded or squarish
(also in Parachromis, Archocentrus, and others); no parhypurapophysis (also not in Amphilophus); secondary
caudal pores forming rows (also in Cr. spilurus + cutteri); caudal emarginate (a unique reversal); peritoneum
only anterodorsally pigmented (also in Rocio).
Description. D. XVIII–XIX, modally 10 rays; A. VII–VIII, modally 7 or 8 rays. First dorsal-fin ray not
divided. Just one row of interradial scales in dorsal and anal fins, up to 5 or 6 (dorsal) and 7 or 8 (anal) scales
long. Predorsal scales 15–20; pored lateral-line scales (not counting scales overlapping between the two seg-
ments of the lateral line) 30–33, prolonged by two pored scales on caudal fin; scales from lateral line to first
dorsal fin ray always 2.5; circumpeduncular scales modally 17. Small to medium-sized heroines, to 165 mm
SL (Kullander 2003). Body fusiform to rather oval, depth 38–46% of SL; caudal peduncle slender, longer than
deep, least depth 13–15% of SL. Maxilla and premaxilla falling short of orbit. Lower jaw receding (contra
Günther 1869, who considered the jaws of both H. nicaraguensis and H. nematopus “equal in front”); frenum
present in lower lip. Dorsal and anal fins bearing filaments. Anal loop and rostral esophageal loop of gut, adja-

cent. Genital papilla oval, in females opening oval and not much crenulated. Gill rakers trapezoidal, denticu-
late, with no basal process. Secondary pored scales on caudal fins forming rows between the rays. Four or
more procurrent rays of caudal fin. Caudal vertebrae modally 16, total vertebrae 29–30. Stripe from snout to
eye absent or diffuse; no interorbital bands; suborbital streak present; no speckles on cheek; no dorsal ocellus
or abdominal black blotch in mature females; absent or diffuse longitudinal band on side of body; 6–7 bars on
sides, diffuse, except for 3rd bar; 4th bar and sometimes also 3rd bar medially more intense, forming a lateral
spot, round or vertically oval; no bars on fins; base of pectoral fin whitish or of same color as breast.
Distribution. Río Santa Clara, Costa Rica, through Great Lakes of Nicaragua north to Río Putkrukira-
Coco, Nicaragua (Fig. 27).
FIGURE 27. Distribution of the species in the genus Hypsophrys. Squares, H. nicaraguensis; triangles, H. nematopus. A
symbol may represent more than one collecting site.
Species composition. Two species, H. nicaraguensis and H. nematopus.

Remarks. Neetroplus is hereby synonymized, so Hypsophrys is no longer monotypic. The other species
assigned to Neetroplus by some recent authors, Cr. panamensis, is here regarded to belong in Cryptoheros (see
above).
The availability of the name, the identity of the type species, the status of Copora and other issues were
dealt with by Kullander and Hartel (1997). Further comments below.

Hypsophrys nicaraguensis (Günther, 1859)
Figures 27–28
Hypsophrys unimaculatus Agassiz, 1859: 408 (nomen nudum, but see Remarks).
Heros nicaraguensis Günther, 1864: 153 (original description).
Heros balteatus Gill in Gill & Bransford, 1877: 184 (junior synonym).
Cichlasoma nicaraguense, Pellegrin 1904: 167 (new combination).
Cichlasoma spilotum Meek, 1912: 73 (junior synonym).
Hypsophrys nicaraguensis, Kullander 1996: 195 (new combination).
Copora nicaraguense, Fernández-Yépez 1969: 3 (new combination).
Holotype. BMNH 1867.9.23.37, 141 mm SL (Fig. 28), J. M. Dow. Lake Nicaragua, Nicaragua. No paratypes.
FIGURE 28. Hypsophrys nicaraguensis, holotype, BMNH 1867.9.23.37. Photo, BMNH, London.
Diagnosis. Unique autapomorphies (Schmitter-Soto, in press): first dorsal fin ray spiniform; pharyngeal
jaws 19 teeth rows wide, 11 rows long; gill rakers on lower limb of first arch modally 9. Easily distinguished
from the other species in the genus by the pointed jaw teeth and the strongly convex head profile (“Cory-
phaena-like”—Günther 1869), among many other differences.
Description. D. XVIII–XIX (modally XIX),9–11; A. VII–VIII,7–9 (modally VIII,8); pectoral 15–16.
First dorsal fin ray not divided. Gill rakers on lower limb of first arch 9–10; gill rakers long, arched, may be
bifid. Subsidiary pored scales on caudal fin in two long rows between rays. Scale rows on cheek 4–5 (contra
Günther 1869, who counted 6); pored lateral-line scales (not counting scales overlapping between the two
segments of the lateral line) 31–33; scales from lateral line to origin of dorsal fin 5–5.5; scales from vent to
interpelvic scale 9–12; anal creases modally 12 (additional meristic data appear in Table 3).
Largest specimen examined, 141 mm SL, but grows at least to 165 mm SL (Kullander 2003). Body depth
44–46% of SL (further morphometric data appear in Table 4). Teeth moderately embedded, at least lateral
teeth; symphysial teeth small, conical, narrow, slightly retrorse; upper symphysial teeth not abruptly larger
than adjacent teeth, lower subequal. Lips not medially narrow; lower lip at corner of mouth square-rounded or
slightly tapering.
Pelvic fins inserted behind origin of dorsal fin. Pectoral fins often falling short of first anal-fin spine; pel-

vic fins always reaching at least to 3rd anal-fin spine. Caudal fin profile emarginate to subtruncate. Scales
moderately ctenoid, rather deciduous.
Gut simple; gut length ca. 80% of SL. Genital papilla rounded, wider than long, almost triangular, may be
smaller than creased area of anus; immaculate or a few melanophores on basal margins.
No stripe from snout to eye. Eyes greenish. Six bars on sides, diffuse; a conspicuous, rounded lateral
blotch, on 4th bar; a diffuse longitudinal bar from orbit to caudal fin, in which there may be a faint blotch on
fin, above lateral line (often absent). Dots on fins (often not noticeable in young). Rows of spots on sides 13–
15, smaller than scales; breast olive-yellowish. Axil of pectoral fin darkened, especially dorsally; base of pec-
toral fin whitish.
Distribution. Ríos Sapoá, Pizote, and Chirripó-Matina, Costa Rica, north to lakes Managua and Nicara-
gua and Río Coco, Nicaragua (Fig. 28).
Remarks. The list of objective synonyms highlights the uncertain generic affinities of the species. How-
ever, its relationship to H. nematopus is supported not only by the present morphological study, but also by
several recent molecular phylogenies (Martin & Bermingham 1998; Hulsey et al. 2004; Concheiro Pérez et al.
2007).
The entire original description of H. unimaculatus Agassiz, 1859 from Lake Nicaragua gives no other
detail but that it “resembles Chrysophrys” (= Sparus aurata Linnaeus), a Mediterranean and north Atlantic
coastal fish (Eschmeyer 2005). Nevertheless, Kullander and Hartel (1997) explained how this simple state-
ment leaves no doubt that the species involved is H. nicaraguensis (the only species in Lake Nicaragua to
have an oval lateral blotch similar to that of Chrysophrys).
Because there are no types of H. unimaculatus and the name constitutes a senior synonym unused after
1899 (ICZN 1999), I shall file a petition to ICZN to conserve the younger specific epithet nicaraguensis.
Hypsophrys nematopus (Günther, 1867), n. comb.

Figures 27, 29
Neetroplus nematopus Günther, 1867: 603 (original description).

Neetroplus fluviatilis Meek, 1912: 74 (junior synonym).
Neetroplus nicaraguensis Gill in Gill & Bransford, 1877:186 (non Günther, junior synonym).
Holotype. BMNH 1865.7.20.35, 94 mm SL (Fig. 29), J. M. Dow. Lake Managua, Nicaragua. No paratypes.
FIGURE 29. Hypsophrys nematopus, holotype, BMNH 1865.7.20.35. Photo, BMNH, London.

Diagnosis. Many unique autapomorphies (fig. 6c in Schmitter-Soto, in press): Teeth properly incisor,
even in young; caudal edge of mesethmoid with a sui generis shape; caudal edge of supraoccipital crest
straight, inclined rostrad; supraoccipital crest height 27% of postethmoid skull length; width of posterior pre-
maxillary expansion greater than 41% of length of same bone; arms of articular equal in length; gut S-folded
ventrorostrally in adults, anal and medial loops nearly touching. Further distinguished from the other species
in the genus by the abrupt, steep head profile.
Description. D. XVIII–XIX (modally XVIII),9–10; A. VII–VIII,6–7 (modally VII,7); pectoral 16–17.
First dorsal fin ray not divided. Gill rakers on lower limb of first arch 8; gill rakers distally broadened, ventrad
curved. Subsidiary pored scales on caudal fin in short rows. Scale rows on cheek 5–6; pored lateral-line scales
(not counting scales overlapping between the two segments of the lateral line) 30–32; scales from lateral line
to origin of dorsal fin 6.5–7, the dorsalmost pair reduced; scales from vent to interpelvic scale 13–17; anal
creases modally 15 (further meristic data appear in Table 3).
Largest specimen examined, 94 mm SL, maximum size 140 mm SL (Conkel 1993). Body depth 38–46%
of SL (further morphometric data appear in Table 4). Teeth not embedded; symphysial teeth equal in level to
adjacent teeth. Upper lip medially narrow; lower lip at corner of mouth tapering.
Pelvic fins inserted far behind origin of dorsal fin. Pectoral fins never reaching 1st anal-fin spine; pelvic
fins always reaching at least 5th anal-fin spine in adults. Caudal fin slightly but definitely emarginate, contra
Günther (1869), who described it as truncate (but depicted it as emarginate in his plate LXXIV, fig. 4). Scales
moderately ctenoid in young, weakly so in adults.
Gut simple in juveniles, but the basic S-shape becoming ventrorostrally folded with growth, and anal loop
displaced caudad until almost touching median loop (fig. 24i in Schmitter-Soto, in press). Gut length ca. 80%
of SL. Genital papilla rounded, semicircular, broader than long, or squarish; sunk; pigmented on margins.
A diffuse stripe from snout to eye; an indistinct vertical bar on head. Eyes blue-silver. Seven bars on sides,
diffuse, except 3rd, which constitutes a large, vertically elongated lateral blotch (this bar may be light on a dark
background instead of dark on light in some color phases); no longitudinal stripe; no blotch on caudal fin, just
base of fin darkened, same as last scales on peduncle. No dots on fins, but often hyaline areas near tip of soft
dorsal and anal fins. No spots on scales; breast dark olive. Axil of pectoral fin dusky or darkened; base of pec-
toral fin whitish or with same coloration as breast.
Distribution. Río Santa Clara, Costa Rica, north to lakes Nicaragua and Jiloá, Nicaragua (Fig. 27).
Remarks. The “incisor” teeth of Cr. panamensis are not a true convergence, but a different character state
(Rogers 1981; Schmitter-Soto, in press).
Discussion
The traditional “working diagnosis” of Archocentrus was based strongly on meristics, especially on the high
number of dorsal and anal fin elements, with formulae D. XVII–XX,8–11 and A. VII–XII,6–9. There have
been also morphometric attributes, albeit rather general: the pectoral fin reaching or exceeding the anal fin ori-
gin; body somewhat deep; snout short, mouth rather small, maxilla not reaching the orbital rim, neither verti-
cally nor horizontally; teeth in the outer row subequal in size anteriorly in both jaws (Regan 1905; Kullander
1996; Greenfield & Thomerson 1997; Miller et al. 2005). No synapomorphies were known for Archocentrus,
Cryptoheros, or Hypsophrys, but some of these traditionally diagnostic characters have turned out to be syna-
pomorphic, e.g. the number of anal-fin spines for Archocentrus; others, as expected, were convergent, e.g. the
length of the maxilla (Schmitter-Soto, in press). Still others, such as body depth, are useful for alpha-level tax-
onomy, but not for phylogeny reconstruction (Zelditch et al. 2005).
Allgayer (2001) had already expressed the view that Archocentrus should be restricted to Ar. centrarchus
and Ar. spinosissimus. Burgess (2000) judged that Ar. multispinosus should be added too, and Allgayer (2001)

himself included Herotilapia in his subtribe Archocentrina (= Archocentrus + Amatitlania + [Cryptoheros
without Cr. panamensis], a group for which Schmitter-Soto [in press], found no support). This opinion,
although only recently published explicitly, is a venerable one: Regan (1908) called Herotilapia “evidently
closely allied” to Archocentrus. Other authors, e.g. Bussing (1976), have considered Herotilapia “derived”
from Archocentrus.
However, most molecular phylogenetic hypotheses (Martin & Bermingham 1998; Farias et al. 2004; Hul-
sey et al. 2004; Concheiro Pérez et al. 2007 —all based on the cytochrome b gene) have found Herotilapia
quite unrelated from Archocentrus and allies. A possible explanation for this pattern may lie on the intrinsic
limitations of cytochrome b. As stated by Martin & Bermingham (1998), “heroine cichlids… are unfortu-
nately refractory to phylogenetic inference using cytochrome b sequences.” Long branches are common in
“dwarf” Neotropical cichlids, such as species of Archocentrus and allies, and the cytochrome b gene presents
saturation at third-codon position (Farias et al. 2004). Other genes should provide interesting independent evi-
dence to settle the issue.
My understanding of Cryptoheros diverges from Allgayer’s (2001), who included also what is here called
Amatitlania, and excluded Cr. panamensis. On the other hand, Kullander (2003), who did not recognize Cryp-
toheros, decided to call this species Archocentrus panamensis, recognizing that its affinities do not lie with
Neetroplus (nor with Hypsophrys). Am. nigrofasciata had been considered a Cryptoheros by Allgayer (2001)
and an Archocentrus by Kullander (2003).
Neetroplus has had a rather unfortunate usage history. Based on a very distinctive but overrated dental
character (like the tricuspid teeth of Herotilapia), monotypic at first, it became an artificial group with the
inclusion of several incisor-toothed cichlids, such as N. carpintis Jordan & Snyder, 1899 (= Herichthys
carpintis), N. bocourti Vaillant & Pellegrin, 1902 (= ‘Cichlasoma’ bocourti), and N. panamensis (= Cr. pana-
mensis). Among this assemblage, H. nematopus is the only species to have incisor teeth as juvenile (Rogers
1981).
The synonymization of Neetroplus with Hypsophrys has the disadvantage of obliterating a widely used
name (Neetroplus nematopus). However, a classification should also convey information about phylogeny,
and this purpose is not served well by recognizing two monotypic sister genera instead of calling their clade
by just one generic name. An example of this stance is the synonymization of Garmanella with Jordanella
(Cyprinodontidae), in “…the interest of having generic categories define derived groups, rather than recog-
nize individual differences…” (Parenti 1981). Moreover, the sister-group relationship of H. nicaraguensis and
H. nematopus is supported by most proposed phylogenies (e.g. Martin & Bermingham 1998; Hulsey et al.
2004; Concheiro Pérez et al. 2007).
It should be acknowledged that a classification alternative to the one here proposed, and equally congru-
ent with the phylogeny presented by Schmitter-Soto (in press), might include all species dealt with in this
work in the genus Hypsophrys (or Amphilophus, or Parachromis, all three names having been proposed in the
same work, on the same page—Kullander & Hartel 1997). It is my opinion, notwithstanding, that the classifi-
cation proposed here is more informative, and also less disruptive with recent usage. On the other hand, since
the aim of this work is to review the species formerly assigned to Archocentrus, the choice of taxa was not
ideal for such higher level decisions.
I follow the Evolutionary Species Concept (ESC). However, the ESC is not an operational concept; to be
able to recognize species under the ESC (i.e., every lineage that “maintains its identity from other such lin-
eages and which has its own evolutionary tendencies and historical fate”: Wiley 1978), I applied here both the
“autapomorphic” and the “diagnosable” versions of the Phylogenetic Species Concept (PSC) (Mayden 1997):
every species should display at least one derived character of its own (Rosen 1979), but, if a non-homoplastic
(unique) autapomorphy is not found, then “the smallest diagnosable cluster of individual organisms within
which there is a [presumed] parental pattern of ancestry and descent” (Cracraft 1983) is to be recognized as a
species. Thus, I agree with Lucena et al. (1992), who point out that “to consider two distinguishable popula-

tions as a single species [implies] loss of information about phylogeny and biogeography.”
The phylogeny and biogeography of the species formerly assigned to Archocentrus are discussed by
Schmitter-Soto (in press).
Acknowledgments
Sven Kullander and Paul Loiselle performed very deep and constructive reviews of this work, which resulted
in a significant improvement. The paper also benefited very much from the detailed editorial labour of Carter
R. Gilbert and Larry M. Page.
This work was done while on sabbatical stay at the University of Michigan Museum of Zoology, Division
of Fishes, under a Fulbright grant (which included a travel award to present advances of this work during the
2004 ASIH meetings at Norman, Oklahoma).
I thank most warmly my hosts, Bill Fink and Jerry Smith, for giving me the opportunity to enjoy not only
their facilities at UMMZ, but also the stimulating environment they have created. An important part of that
world is the “Fish Club,” whose cichlidologist members Prosanta Chakrabarty and Ron Oldfield discussed
with me several interesting issues germane to this project.
Doug Nelson helped tremendously with loan management. Live photos are by Humberto Bahena. Type
photographs are by H. Bahena (Cr. chetumalensis), Rick Feeney and Jeff Seigel (Cr. sajica), Claude Ferrara
(Ar. spinosissimus, C. immaculatum, Cr. altoflavus, Cr. nanoluteus), Kyle Luckenbill (Cr. cutteri), C. Ratton
(R. octofasciata), Melanie Stiassny (Cr. myrnae), Phil Willink (Am. kanna) and the Photographic Unit of the
Museum of Natural History, London (Ar. centrarchus, Ar. multispinosus, Cr. septemfasciatus, Am. nigrofasci-
ata, H. nicaraguensis, H. nematopus). Janneth Padilla produced the maps; H. Bahena processed the photo-
graphs. Many people, most constantly Héctor Gamboa-Pérez and Roberto Herrera, participated with me in
field work to collect species deposited at ECOCH.
Not only for loan of material under their care (including permission to clear and stain some specimens),
but also for hospitality and enlightening exchange of ideas, I wholeheartedly thank Mary Anne Rogers and
Mark Westneat (FMNH), Guy Duhamel and Javier Gregorio (MNHN), Oliver Crimmen and Patrick Campbell
(BMNH), Sonia Fisch-Müller and Claude Weber (MHNG), John Lundberg and Kyle Luckenbill (ANSP),
Rocío Rodiles-Hernández and Alfonso González-Díaz (ECOSC), J.T. Williams (USNM), Oldřich Řičan and
Sven Kullander (NRM), Salvador Contreras-Balderas (UANL, co-discoverer of R. gemmata), Reeve Bailey
(who detected Am. siquia decades before I was born, but nobly declined any coauthorship in its description),
Jean Huber, Darrin Hulsey, Melanie Stiassny, and Martha Valdez-Moreno.
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Appendix 1. Material examined. Primary types mentioned first, followed by secondary types and then non-type mate-
rial, ordered alphabetically by collection codon and numerically by catalog number. Number of specimens in parenthe-
ses. Abbreviations: C&S, cleared and stained; skel., skeletonized; XR, radiographed; dig., digitized. Institutional codons
follow ASIH (2005). More details (e.g. collectors, dates, etc.) available from the author on request.
Amatitlania
Am. coatepeque: UMMZ 245584 (holotype, dig.), UMMZ 181823 (18 paratypes, 2 C&S, 16 dig.), FMNH 12136
(10 paratypes, dig.), El Salvador, island in Lake Coatepeque; UMMZ 202805 (4 paratypes, dig.), El Salvador, Santa Ana,
E coast of Lake Coatepeque at Monterrey.
Am. kanna: FMNH 59243 (holotype, dig.), FMNH 116465 (paratype, dig.), Panama, San San swamp; BMNH
1925.3.6.119 (1 paratype), Panama, Río Sixaola at Isla Grande, creek and swamp; BMNH 1925.3.6.120-121 (2
paratypes), Panama, swamp near hwy bridge, San San; BMNH 1925.3.6.122-123 (2 paratypes), Panama, Río Cricamola
near Konkintu; BMNH 1925.3.6.126 (1 paratype), Panama, Río Nigua into Bahía Almirante; FMNH 59240 (2, 1 C&S,
paratypes), Panama, tributary of Río Cricamola near “Conquanta” [Konkintu]; FMNH 59241 (1 paratype), Panama,
creek and pond, tributaries of Quebrada Nigua, Bahía Almirante; FMNH 59242 (1 paratype), Panama, Isla Grande creek
and swamp into Rìo Sixaola; MHNG 2646.78 (2 paratypes), Panama, Chiriquí Grande; UMMZ 145716 (1, XR, dig.,
paratype), Panama, Río Cricamola at San San; ANSP 156817 (1), Panama, Bocas del Toro, Río San San; UMMZ 145231
(1), Isla Grande creek and swamp into Rìo Sixaola; Panama, UMMZ 145715 (1), Panama, a tributary of Río Cricamola
above Konkintu; UMMZ 145729 (1), Panama, creek and pond, tributaries of Quebrada Nigua, Bahía Almirante.
Am. nigrofasciata: BMNH 1865.4.29.76 (lectotype), BMNH 1865.4.29.77 (12 paralectotypes), BMNH
1865.4.29.78 (1 paralectotype, skel.), Guatemala, Lake Amatitlán; ANSP 77834 (104), Guatemala, outlet of “Amatitan”
[Lake Amatitlán]; ANSP 101836 (212), Guatemala, Lake Atitlán; BMNH 1961.7.3.1 (1), Guatemala, Lake Atitlán;
FMNH 12137 (2), El Salvador, El Ángel; FMNH 12139 (7, dig.), El Salvador, Lake “Calchuapa” [Chalchuapa]; MNHN
0000-5729 (1), Guatemala, Lake Amatitlán; UMMZ 143958 (10), Guatemala, Escuintla, Río Guacalote; UMMZ 166471
(30), UMMZ 188223 (14), Guatemala, Sololá, Lake Atitlán; UMMZ 188245 (11, 2 C&S, 11 dig.), Honduras, Morazán,
Quebrada de La Chocona; UMMZ 188151 (2), Honduras, tributary to Río Nacaome; UMMZ 188228 (1), Guatemala,
Lake Amatitlán; UMMZ 194127 (101), Guatemala, Suchitepéquez, Río La Primavera, 6 km ESE of Eca Cocales;
UMMZ 197392 (23, 2 C&S, 16 dig.), Guatemala, Jutiapa, river 7.5 km SE of Asunción Mita; UMMZ 199558 (26), Hon-
duras, tributary to Río Patuca; UMMZ 202781 (1), El Salvador, San Salvador, Lago Ilopango at Apula; UMMZ 202812
(9), El Salvador, headwaters of Río Sucio, 1 km SSE of Armenjá.
Am. siquia: UMMZ 245585 (holotype), UMMZ 196948 (25 paratypes, 2 C&S, 16 dig.), UMMZ 196954 (7
paratypes), Nicaragua, Río Siquia, 7 mi above Rama; ANSP 88241 (3 paratypes), Costa Rica, “Esperata” [Esparta];
ANSP 67475 (4 paratypes), Nicaragua, “great falls” on Río Pis Pis; ANSP 140692 (1 paratype), Costa Rica, Puntarenas,
Río Ahogados; FMNH 7733 (1 paratype), Costa Rica, Turrubares; UMMZ 145720 (2 paratypes, 2 dig.), Costa Rica,
Talamanca, Río Bioliri, tributary to Río Lari; UMMZ 166474 (1 paratype), Costa Rica, Laguna del Misterio; UMMZ
196939 (1 paratype, dig.), Nicaragua, creek tributary of Río Siquia, 7 mi upstream from Rama; BMNH 1925.3.6.124-125
(2 paratypes), Costa Rica, Talamanca, Río Bioliri, tributary to Río Lari; MHNG 2160.60 (1 paratype), Nicaragua, Lake
Managua at Momotombo; UMMZ 213938 (6 paratypes), Nicaragua, Rivas, Río Cañas Gordas; ANSP 93215 (2), Costa
Rica, “Esperata” [Esparta]; ANSP 168328 (57), Costa Rica, Alajuela, Lake and Río Cuarto; BMNH 1929.8.9.20-29 (10),
Costa Rica, Liberia; BMNH 1968.1.12.2-4 (3), Nicaragua; BMNH 1968.1.12.32-33 (2), Costa Rica, “Squirres”
[Siquirres]; UMMZ 165774 (1), Nicaragua, Matagalpa; UMMZ 166472 (2), Costa Rica, Laguna del Misterio; UMMZ
180616 (1), Nicaragua, Lago Managua at Matearé; UMMZ 188120 (3), Honduras, El Paraíso, Río Cato in Valle de
Jamastrán; UMMZ 188144 (6, 4 dig.), Honduras, Morazán, Lagunita S of road Tegucigalpa-Danlí; UMMZ 188257 (7),
UMMZ 188258 (1), Nicaragua, Zelaya, Río Huahuashán at Corozo Camp; UMMZ 188293 (4), Honduras, Morazán, Río
Yeguaré at Tegucigalpa; UMMZ 190191 (18, 2 C&S, 14 dig.), Costa Rica, Guanacaste, 5 km NW of Las Cañas, Río
Corobicí; UMMZ 190199 (16, 6 dig.), Costa Rica, Guanacaste, Río Piedras, 12 mi SE of Liberia; UMMZ 196939 (1,
dig.), UMMZ 199627 (129), Nicaragua, Río Kurnog, tributary of Laguna Póhara [Pahara?], SE of Bilwaskarma; UMMZ
199637 (86, 6 dig.), Nicaragua, 6 km E of Waspam, Río Putkrukira, tributary of Río Coco; UMMZ 199650 (51, 6 dig.),
Nicaragua, 35 km S of Waspam, Río Likus, tributary to Río Wawa; UMMZ 199661 (35), Nicaragua, Río Coco, 300 m
upstream of Waspam; UMMZ uncat. (22), Costa Rica, Guanacaste, Tempisque, 2 mi S of Hacienda Tenorio.

Archocentrus
Ar. centrarchus: BMNH 1905.3.27.2 (labelled “cotype”), Nicaragua, Lake Nicaragua; FMNH 5982 (62, XR), Nica-
ragua, Granada, Laguna Jenicero [Jenízaro]; UMMZ 180612 (1, XR), Nicaragua; UMMZ 180643 (1, XR), Costa Rica,
Limón, Laguna Tortuguero, 2.5 mi from inlet; UMMZ 224131 (50, 1 C&S, 1 partly skel., 16 dig.), Nicaragua, Río Sapoá
into Lake Nicaragua.
Ar. multispinosus: BMNH 1865.7.20.34 (holotype), Nicaragua, Lake Managua; UMMZ 180621 (1, dig.), Nicaragua,
Lake Nicaragua at Granada; UMMZ 199539 (26, 2 C&S, XR, 16 dig.), Honduras, Río Patuca, canal to Brus Laguna;
UMMZ 213939 (5, dig.), Nicaragua, Rivas, Río Cañas Gordas; UMMZ 228679 (6), Honduras, Gracias a Dios, Río Rapa.
Ar. spinosissimus: MNHN A-0352 (lectotype), MNHN 2005-0780 (3 paralectotypes), Guatemala, Izabal, Río
Polochic; UMMZ 146070 (1, 1 partly skel.), Guatemala, Alta Verapaz; UMMZ 146086 (4, 1 partly skel.), Guatemala,
Izabal, Río Polochic; UMMZ 190737 (25), UMMZ 197249 (7, 1 C&S, XR, 7 dig.), Guatemala, Izabal, 19 km SW of
Livingston, Río Paujilá on El Golfete.
Cryptoheros (Panamius)
Cr. panamensis: FMNH 7601 (holotype), Panama, Canal Zone, Río Mandinga at “Bas Obispo”; FMNH 8112 (1),
Panama, Gorgona; NRM 13097 (3), Panama, Río Ipetí; NRM 37100 (4), Panama, Río Mandinga; UMMZ 170964 (3,
XR, dig.), Panama, road to Madden Dam, a tributary of Lake Gatún.
Cryptoheros (Cryptoheros)
Cr. chetumalensis: ECOCH 5467 (holotype, dig.), ECOCH 1005 (11 paratypes), ECOCH 1693 (8 paratypes), Mex-
ico, Quintana Roo, Arroyo Aguadulce, tributary of Río Hondo, at Sabidos, about 20 km upstream from Chetumal;
ECOCH 1465 (2 paratypes), Mexico, Quintana Roo, Arroyo Ucum, tributary of Río Hondo, at Ucum; ECOCH 1536 (1
paratype), Mexico, Quintana Roo, outlet of Laguna Encantada into Río Hondo; ECOCH 1559 (4 paratypes), Mexico,
Quintana Roo, Laguna Kaná; ECOCH 1593 (3 paratypes), ECOCH 2328 (7 paratypes), Mexico, Quintana Roo, Laguna
Ocom; ECOCH 1900 (13 paratypes), Mexico, Quintana Roo, Laguna Caobas; UMMZ 210888 (9 paratypes, 2 C&S, 8
dig.), Mexico, Quintana Roo, Río Hondo at La Unión; BMNH 1974.1.3.753-755 (3), British Honduras [Belize]; UMMZ
159308 (4), UMMZ 167692 (16, 16 dig.), UMMZ 167696 (56, 2 C&S, 16 dig.), Belize, Cayo, Belize River; UMMZ
187977 (21), Guatemala, Petén, Río Sarstún; UMMZ 197224 (16, 2 C&S, 15 dig.), Guatemala, Río Nimblajá, 1 km
above mouth into Río Sarstún.
Cr. cutteri: ANSP 53930 (holotype, dig.), ANSP 53931 (1), ANSP 53932 (1), ANSP 53933 (1, XR) (paratypes,
dig.), Honduras, Atlántida, Río Lancetilla; ANSP 56143 (1), Honduras, near La Ceiba; BMNH 1933.1.17.1-2 (2),
BMNH 1933.3.28.22-24 (3), aquarium material from Honduras; MNHN 2001-1168 (1), Honduras, Lake Yojoá; UMMZ
113404 (5), Honduras, river off Tela; UMMZ 155878 (2), Honduras, Comayagua, Río Celán, tributary of Río Humuya;
UMMZ 173195 (9, 2 C&S, 9 dig.), Honduras, Atlántida, Río Lancetilla; UMMZ 173235 (1), Honduras, Santa Bárbara,
W coast of Lake Yojoá; UMMZ 173274 (4), Honduras, Cortés, brook in Veracruz; UMMZ 173291 (5), Honduras,
Cortés, Río de Masca; UMMZ 173301 (16), Honduras, Cortés, Segundo Río Tulián; UMMZ 188219 (58), Honduras,
Comayagua, Río Celán (Río Selguapa), tributary of Río “Humuyu” [Humuya]; UMMZ 188134 (5, dig.), UMMZ 188136
(5, 2 C&S, 5 dig.), Honduras, Cortés, E coast of Lake Yojoá; UMMZ 188214 (15), Honduras, Comayagua, Río Humuya;
UMMZ 188219 (58), Honduras, Comayagua, Río Celán;UMMZ 193847 (10), Honduras, Colón, Río Sico, 30 km SW of
Iriona; UMMZ 193858 (2), Honduras, Atlántida, creek 34 km WSW of La Ceiba; UMMZ 193983 (31, 6 dig.), Guate-
mala, Zacapa, Río Achuelo, W of Gualán; UMMZ 194294 (21, 6 dig.), Guatemala, Zacapa, Río Matasano; UMMZ
197302 (4, 4 dig.), Guatemala, Zacapa, Quebrada Juilín; UMMZ 199605 (37), Honduras, Río Sikri, S of Laguna
Sikalanka; UMMZ 199678 (111, 2 C&S, 14 dig.), Honduras, tributary of Caribbean Sea at mouth, 5 km W of Trujillo;
UMMZ 213669 (2, dig.), Honduras, Colón, Río Guinea, E of Santa Fe; UMMZ 219146 (1), Honduras, Río Aguán, 44.6
mi W of Trujillo; UMMZ 228665 (2), Honduras, Atlántida, Río Jutiapa.
Cr. spilurus: BMNH 1864.1.26.52 (lectotype, dig.), BMNH 1864.1.26.53-55 (3 paralectotypes, dig.), Guatemala,
Lake “Isabel” [Izabal]; BMNH 1865.4.29.74 (1), Guatemala, Río Motagua; MNHN 0000-9845 (3), Guatemala, “Le
Mullins”; UMMZ 146096 (2), Guatemala, Alta Verapaz, El Canal, tributary of Río Polochic; UMMZ 190591 (32, 6
dig.), Guatemala, Izabal, Río del Amatillo; UMMZ 190625 (16, 6 dig.), Guatemala, Izabal, Río Samahyi, 10 mi SE of
Modesto Méndez; UMMZ 190642 (23, 6 dig.), Guatemala, Izabal, Río Sauce 4 km E of El Estor; UMMZ 190669 (1)

Guatemala, Izabal, Río Tarnejá, 10 km SW of Livingston; UMMZ 190691 (122), Guatemala, Izabal, Río San Marcos;
UMMZ 190709 (3), Guatemala, Río San Francisco II, 15 km S of Puerto Barrios; UMMZ 190735 (38), Guatemala, Río
Juaquitún, tributary to Río Secón; UMMZ 190754 (3, 3 dig.), Guatemala, Alta Verapaz, Río Polochic, 1.5 W of Panca-
jché; UMMZ 197198 (62, 2 C&S, XR), UMMZ 197265 (34, 10 dig.), Guatemala, Izabal, Río Dulce N of mouth of Río
Ciénaga; UMMZ 197317 (20), Guatemala, Izabal, Río Trincheras; UMMZ 198872 (2); UMMZ 225000 (2, 2 dig.), Gua-
temala, Izabal, Río Dulce at mouth on Lago “Isabel” [Izabal]; UMMZ uncat. (7), Guatemala, Río Blanco; UMMZ uncat.
(1), Guatemala, Lago Izabal. [Cichlasoma spinosissimum var. immaculatum: MNHN 9846 (2 syntypes), Guatemala, Río
Polochic.]
Cryptoheros (Bussingius)
Cr. altoflavus: MNHN 2001-1163 (holotype, dig.), MNHN 2001-1164 (1, XR), MNHN 2001-1167 (1) (paratypes),
Panama, río Cañaveral; ANSP 156820 (1), Panama, Bocas del Toro, Río Changuinola; BMNH 1925.3.6.127-131 (6),
Panama, Río Cricamola near Konkintu; UMMZ 145722 (4, 1 C&S, 4 dig.), UMMZ 145723 (2, dig.), Panama, Guibarí
Creek, tributary to Río “Cricamol” [Cricamola] at Konkintu.
Cr. myrnae: UMMZ 217739 (20, 2 C&S, 8 dig., paratypes), Costa Rica, Limón, Río Cocolis, tributary to Río Sixa-
ola, 3.5 km SE of Shiroles; ANSP 163131 (1), ANSP 163179 (1), ANSP 163181 (1), ANSP 163195 (1), Costa Rica,
Limón, Bri Bri; ANSP 168312 (1), Costa Rica, Limón, headwaters of Río Estrella, 13 km upstream from Pandora;
FMNH 6261 (1), Costa Rica, Turrubares; UMMZ 145708 (1, dig.), UMMZ 145711 (1), Costa Rica, creek tributary to
Río Tiliri-Sixaola; UMMZ 180677 (3, 1 C&S, 3 dig.), Costa Rica, Limón, Río Sixaola.
Cr. nanoluteus: MNHN 1993-260 (holotype, dig.), MNHN 1993-261 (1), MNHN 1993-263 (1, XR) (paratypes),
ANSP 104377 (9), Panama, Bocas del Toro, brooks of Río Guarumo at Chiriquicito; MHNG uncat. (2), Panama, Río
Priki; NRM 25942 (2), Panama, Chiriquí Grande, Río Guarumo (Guabo); NRM 35880 (1), Panama, Bocas del Toro, Río
Guaromo [Guarumo].
Cr. sajica: USNM 211617 (20 paratypes), Costa Rica, Puntarenas, a tributary of Río Sierpe, 2 km S of Palmar Sur;
MHNG 2447.21 (1), Costa Rica, Puntarenas, Río Ceiba; UMMZ 194210 (4), UMMZ 194239 (28, 2 C&S, 8 dig.), Costa
Rica, Puntarenas, a tributary to Río Ceiba, just above mouth.
Cr. septemfasciatus: BMNH 1909.3.13.82 (lectotype, dig.), BMNH 1909.3.13.83-90 (17 paralectotypes), Costa
Rica, Río Iroquois; ANSP 45401 (2), Costa Rica, “Guapilis” [Guápiles]; UMMZ 180613 (1), Nicaragua, Lago Nicaragua
at Granada; UMMZ 180678 (3), Costa Rica, Turrialba, a tributary of Río Reventazón; UMMZ 190367 (120, 1 C&S, 16
dig.), Costa Rica, Limón, 1 km E of Guápiles, Río Santa Clara and tributary; UMMZ 199872 (2), Costa Rica, Heredia,
Río Puerto Viejo, ½ mi upstream of Sarapique.
Hypsophrys
H. nicaraguensis: BMNH 1867.9.23.37 (holotype, dig.), Nicaragua, Lake Nicaragua; FMNH 5995 (10, 6 dig.), Nic-
aragua, Lake Managua; FMNH 5996 (12, XR), UMMZ 181826 (2, 1 skel.), Nicaragua, Jiloá; UMMZ 199639 (8, 1 C&S,
4 dig.), Nicaragua, 6 km E of Waspam, Río Putkrukira, tributary of Río Coco. [Cichlasoma spilotum: UMMZ 188994
(3), Costa Rica, Victoria; Heros balteatus: BMNH 1905.3.27.3 (cotype), Nicaragua, Lake Nicaragua.]
H. nematopus: BMNH 1865.7.20.35 (holotype, dig.), Nicaragua, Lake Managua; UMMZ 166475 (2), UMMZ
181824 (2), UMMZ 197507 (20, 2 C&S, 10 dig.), Nicaragua, Lake Jiloá.
Rocio
R. gemmata: ECOCH 4054 (holotype, dig.), ECOCH 1468 (1 paratype, C&S, dig.), ECOCH 3145 (1 paratype, dig.),
Mexico, Quintana Roo, nameless cenote 12 km N of Leona Vicario; UANL 15046 (4 paratypes), Mexico, Quintana Roo,
Laguna Leona Vicario.
R. ocotal: UMMZ 245583 (holotype, dig.), UMMZ 171140 (5 paratypes, XR, dig.), Mexico, Chiapas, Lacandon
region, Laguna Ocotal.
R. octofasciata: MHNG 665.55 (holotype, dig.), MHNG 2666.83 (8), Mexico, Veracruz, Cosamaloapan; BMNH
1890.10.10.96-99 (2), Mexico, Río de Sarabia; BMNH 1905.12.6.777-784 (3), Mexico, Oaxaca, Río Obispo; BMNH
1913.6.21.181-190 (3), Mexico, “Vera Cruz”; BMNH 1935.9.9.18 (1), Mexico, “Puerto” [Veracruz?]; ECOCH 5468 (1
skel.), Mexico, Quintana Roo, Arroyo de Pedro A. Santos; ECOSC 693 (10), Mexico, Chiapas, Río Cedros; ECOSC

1060 (2), ECOSC 1105 (2), ECOSC 1187 (2), Mexico, Chiapas, Laguna Carranza; FMNH 82094 (1), Belize, Orange
Walk, Ramgoat Creek; FMNH 82104 (1), Belize, Cayo, Aguacate Lagoon; FMNH 82277 (2), Belize, Orange Walk, pond
on dry Wamil Creek near Gallon Jug; FMNH 83136 (1), Belize, Río Hondo at San Antonio; FMNH 97685 (1), Belize,
Corozal, pond 6 mi N of Orange Walk; MHNG 673.67 (5), Mexico, Veracruz, Cosamaloapan; MHNG 2395.28 (15),
Mexico, Chiapas, “Lac des Lacandons”, near Bonampak; MHNG 2395.36 (6), Mexico, Campeche, “marais près de Pal-
izada”; MNHN 0000-5731 (3), Mexico, Veracruz; MNHN 1894-242 (1), “Jamaïque” [wrong locality]; MNHN 1904-
0036 (1), British Honduras [Belize]; MNHN 1910-0200 (1), “Brésil” [wrong locality]; UMMZ 97678 (5, 4 dig.), Mex-
ico, Veracruz, Río Santa Fe, tributary of Río Papaloapan; UMMZ 102133 (6, 1 C&S, 6 dig.), Mexico, Yucatan, Miramar
spring at “Talcha” [Telchac]; UMMZ 108567 (2), UMMZ 124187 (3), UMMZ 124257 (1), Mexico, Oaxaca, laguna near
Río Papaloapan; UMMZ 124200 (2, 2 dig.), UMMZ 124221 (1, 1 dig.), UMMZ 124272 (1), Mexico, Oaxaca, Arroyo
Zacatispan, tributary of Río Papaloapan; UMMZ 124238 (2), UMMZ 124265 (1), Mexico, Oaxaca, ponds 4-5 km S of
Papaloapan; UMMZ 143964 (52, 2 C&S, 40 dig.), Guatemala, Petén, Laguna de Petenxil, outlet; UMMZ 143967 (43),
UMMZ 181804 (2, 2 dig.), Mexico, Veracruz, 6 mi NW of Alvarado; UMMZ 178539 (2), Mexico, Oaxaca, Río Sarabia;
UMMZ 181301 (53), Mexico, Veracruz, Río Viejo, 1 mi E of Achiotal; UMMZ 187779 (31, 3 skel.), Mexico, Veracruz,
Estero de Tatagapilla, 4.5 km WSW of Tenochtitlan; tributary of Río Chiquito, Coatzacoalcos drainage; UMMZ 184694
(1, XR), Mexico, Campeche, Laguna Pom; UMMZ 186659 (1), Mexico, Veracruz, ditch on hwy to Medellín, Río Jamapa
drainage; UMMZ 190157 (9, 2 C&S, 9 dig.), British Honduras [Belize], estuary near Belize City; UMMZ 190859 (1, 1
skel.), Mexico, Campeche, Xpujil; UMMZ 190867 (16, XR, 7 dig.), Mexico, Campeche, Zohlaguna; UMMZ 190959 (1,
1 C&S), Mexico, Campeche; UMMZ 194699 (2), Mexico, Veracruz, creek between Jesús Carranza and Veracruz, under
railroad bridge; UMMZ 194704 (1), Mexico,Veracruz, 5 mi N of Jesús Carranza; UMMZ 194715 (1), Mexico, Veracruz,
tributary of Arroyo Santa Lucrecia, E of Jesús Carranza; UMMZ 194795 (3), Mexico, Veracruz, Arroyo Amate, 8.5 km
E of Minatitlán; UMMZ 194872 (4), Mexico, Veracruz, 3 km W of Coatzacoalcos; UMMZ 196387 (48, 6 dig.), Mexico,
Veracruz, spring 6.7 mi ENE of intersection of hwy 175 and hwy to Tierra Blanca; UMMZ 196567 (4, 1 C&S, 4 dig.),
Mexico, Yucatan, cenotes 2.5 km S of Sisal; UMMZ 197245 (1), Guatemala, Izabal, Río Paujilá, 19 km SW of Living-
ston; UMMZ 201745 (2, dig.), Mexico, Yucatan, cenote near Tizimín; UMMZ 202707 (12, 2 C&S, 12 dig.), Mexico,
Tabasco, 4 km N of Teapa, Río Muerto, tributary to Río Teapa; UMMZ 202871 (12), Belize, Gabourel Creek; UMMZ
202918 (6, XR, dig.), Belize, “chorro” 4 mi N of Corozal; UMMZ 202922 (6, 3 dig.), Belize, estuary near Belize City;
UMMZ 209326 (6, dig.), Mexico, Chiapas, 5 km N of Lacanjá; UMMZ 209358 (16, 1 dig.), Mexico, Chiapas, creek near
Palenque [at Nututún?]; UMMZ 209369 (1, dig.), Mexico, Chiapas, 24 km E of Palenque; UMMZ 209590 (12), Mexico,
Veracruz, tributary of Río Tonalá; UMMZ 209654 (8), Mexico, Veracruz, channel 5.7 SW of bridge over Río Papaloa-
pan; UMMZ 209661 (48, 6 dig.), Mexico, Veracruz, 3.2 mi S of toll bridge over Río Papaloapan; UMMZ 210772 (15),
UMMZ 210789 (8, dig.), Mexico, Oaxaca, oxbow S of Río Papaloapan; UMMZ 210986 (18, 2 C&S, 16 dig.), Mexico,
Veracruz, 10 km S of Playa Vicente; UMMZ 223375 (3), Mexico, Veracruz, Río La Antigua. [Cichlasoma biocellatum:
BMNH 1905.12.5.30 (holotype), “Río Negro at Mañaos, Brazil” (wrong locality, probably aquarium specimens); C.
hedricki: FMNH 4673 (holotype and 10 paratypes), FMNH 4579 (paratypes), Mexico, Veracruz, Veracruz; FMNH 4597
(25), Mexico, Veracruz, Pérez; MNHN 1905-0480,0481 (2), UMMZ 167472 (2, 2 dig.), UMMZ 176671 (8 paratypes, 1
skel., 7 dig.), Mexico, Oaxaca, Río Obispo, tributary of Río Papaloapan.]
Additional material
Amphilophus citrinellus: BMNH 1864.1.26.201–203 (3 syntypes), Nicaragua, Lake Nicaragua; UMMZ 180617 (7, 2
C&S, 6 dig.), UMMZ 188309 (7, 1 C&S, 1 skel., 6 dig.), Nicaragua, Lake Managua at Matearé; UMMZ 197508 (10, 10
XR), Nicaragua, Jiloá [Heros basilaris: BMNH 1905.3.27.4 (cotype), Nicaragua, Lake Nicaragua.]
Caquetaia spectabilis: UMMZ 190822 (1, partly skel.), aquarium material, “probably from Guyana”; UMMZ
215564 (6, 6 C&S, dig.), Guyana, Rupununi, Sawariwara River.
Oreochromis mossambicus: UMMZ 190378 (45), Guatemala, Escuintla, Río Michatoya; UMMZ 199400 (1 C&S),
UMMZ 199401 (2 C&S, 1 partly skel.), aquarium material; UMMZ 213374 (12, 1 C&S, 8 dig.), USA, Idaho, Barney
springs.
Parachromis dovii: UMMZ 166473 (5, 1 C&S, 2 skel.), Costa Rica, Laguna del Misterio; UMMZ 188256 (1, 1
C&S), UMMZ 197399 (41, 2 C&S), UMMZ 199651 (28, 1 C&S, 10 dig.), Nicaragua, 35 km S of Waspam, Río Likus,
tributary to Río Wawa.

Parachromis loisellei: UMMZ 203897 (holotype and 7 paratypes, 1 C&S, 1 skel.), Costa Rica, Río Sixaola; ANSP
163184 (1), Costa Rica, Shiroles; ANSP 163661 (1), Costa Rica, Upala; UMMZ 145739 (4, 1 C&S, 1 skel.), Panama,
San San; UMMZ 203898 (8), ; UMMZ 223304 (8, 2 C&S, 8 dig.), Honduras, Colón, pond 13.2 km E of Trujillo.
Tomocichla sieboldii: UMMZ 145733 (5 XR), Panama; UMMZ 167296 (XR), Costa Rica; UMMZ 194211 (7),
Costa Rica, San José, Río San Isidro, tributary to Río Chirripo, E of San Isidro del General; UMMZ 194240 (31, 15 XR,
2 C&S, 1 partly skel., 10 dig.), Costa Rica, Puntarenas, tributary, E bank of Río Ceiba.
Petenia splendida: UMMZ 144128 (1), Guatemala, Petén, Ek Kixil; UMMZ 189987 (2 skel.), Guatemala, Quiché,
Arroyo Cancaná, ca. 1 km above mouth into Río Chixoy; UMMZ 196461 (1 skel.), Mexico, Campeche, Río Candelaria;
UMMZ 196476 (22, 5 dig.), UMMZ 196665 (2 skel., 5 dig.), Mexico, Quintana Roo, Laguna Caobas; UMMZ 223251
(1, 1 C&S), Mexico, Campeche, Laguna de Términos.

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ZOOTAXA

A systematic revision of the genus Archocentrus (Perciformes: Cichlidae),

FIRST PUBLISHED IN 2007 BY

© 2007 Magnolia Press

ISSN 1175-5326 (Print edition)

2 · Zootaxa 1603 © 2007 Magnolia Press SCHMITTER-SOTO

A systematic revision of the genus Archocentrus (Perciformes: Cichlidae),

Accepted by L. Page: 22 Jun. 2007; published: 28 Sept. 2007 3

Key words: taxonomy, cichlids, Middle America

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Basonym Author and year Included in Archo- Present status or name

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Material and methods

Species Author and year Original genus Reference

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Key to Archocentrus and allied genera

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Key to species of genus Hypsophrys

Key to species of genus Rocio

Key to species of genus Amatitlania

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Key to species of genus Archocentrus

Key to subgenera of genus Cryptoheros

16a(7b) Teeth truncate and labiolingually compressed, incisor-like...................................................................

Key to species of genus Cryptoheros (subgenus Bussingius)

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Key to species of genus Cryptoheros (subgenus Cryptoheros)

Genus Archocentrus Gill

Type species. Heros (Archocentrus) centrarchus Gill, 1877, by monotypy.

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centrarchus 3 5(3) 2(5) (2) (1)

spinosissimus 1 1(2) 1(1) 7(4) 1(2) (1) (1)

multispinosus 5 4(1) 1(7) (1)

panamensis 3(1) (1) (1)

spilurus (1) 5(5) 4(6) (10) 1(1) (1)

chetumalensis 1(1) 1(1) 1(1) 1(1) 1(1)

cutteri 2 3(3) 3(5) 4(2) 1(4) 1 (1)

septemfasciatus 3 5 3(6) 1(5) (2) (1)

altoflavus 1 1 1(1) (1) (1)

myrnae 6 3(2) 1(4) (1)

nanoluteus 3(1) (1) (1)

sajica 1 5(1) 3(4) 1(5)

nigrofasciata 2(1) 4(2) 8(3) 5(11) 3(2) 1(4)

coatepeque 1 5(1) (2) 4(5) 1(4) 1(2)

kanna 2 2(3) 1 4(2) (3) (5) (1)

siquia (1) 5 7(1) 6(9) 3(1) 1(6) (5) (2) (1)

octofasciata 1(2) 2(1) 13(14) 13(36) 5(54) (13) (7) (1)

ocotal 3(2) (2) 1 1(1) (1)

gemmata 1 1(1) (1) 1(1)

nicaraguensis 2 1 2 4(4) 2(3) 1(3)

nematopus 3 (1) (1) (1)

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Number of scales in longest interradial row on soft dorsal fin

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Number of scales in longest interradial row on soft anal fin

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Dorsal-fin spines Dorsal-fin rays

XV XVI XVII XVIII XIX XX 7 8 9 10 11 12

octofasciata 26 122 16 13 48 106 7

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Anal-fin spines Anal-fin rays

VI VII VIII IX X XI XII XIII 6 7 8 9