Phylogenetic Analysis and Taxonomy of The Poeciloid Fishes

Zoological Journal of the Linnean Society (2000), 130: 1–53.
With 22 figures
doi:10.1006/zjls.1999.0213, available online at http://www.idealibrary.com on
Phylogenetic analysis and taxonomy of

the poecilioid fishes (Teleostei:
Cyprinodontiformes)
MICHAEL J. GHEDOTTI∗
Natural History Museum, University of Kansas, Lawrence, KS 66045-2454, U.S.A.
Received July 1998; accepted for publication July 1999
Evidence from morphology is used to infer the phylogeny of the superfamily Poecilioidea
using other cyprinodontoid fishes as outgroups. The three equally most parsimonious trees
resulting from the phylogenetic analysis support the monophyly of the families Anablepidae
and Poeciliidae with respect to each other, but the previous taxonomy within the Poeciliinae
is not consistent with the resultant phylogenetic trees. The Poeciliidae is recognized with
three subfamilies: the Aplocheilichthyinae containing solely Aplocheilichthys spilauchen, the
Procatopodinae containing Fluviphylax (Fluviphylacini) and the African lamp-eyed killifishes
(Procatopodini), and the Poeciliinae. The inferred hierarchical relationships of included
suprageneric taxa are: ((Oxyzygonectinae, Anablepinae) (Aplocheilichthyinae ((Fluviphylacini,
Procatopodini) (Alfarini (Priapellini (Gambusini (Heterandrini (Cnesterodontini (Girardini,
Poeciliini))))))))). The tribe Alfarini is resurrected and a new tribe, the Priapellini, is described.
Tomeurus gracilis is not the most basal poeciliine, and facultative viviparity in Tomeurus is not
a plesiomorphic intermediate condition of viviparity retained from the common ancestor of
poeciliines. Facultative viviparity in Tomeurus is the result of an evolutionary loss of obligate
viviparity. Tomeurus gracilis is recognized as a member of the tribe Cnesterodontini. Lamprichthys
tanganicus and Micropanchax pelagicus are not sister taxa, and the pelagic lacustrine habits of
these two species are inferred to have evolved independently. Based on the principles of
vicariance biogeography, the origin of the Poecilioidea is inferred to have occurred before
the separation of Africa and South America.
 2000 The Linnean Society of London
ADDITIONAL KEY WORDS:—Poecilioidea – morphology – phylogeny – Poeciliidae –

Procatopodinae – Aplocheilichthyinae – Poeciliinae – Anablepidae – viviparity – gonopodium
– biogeography.
CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . . . . . 2
Taxonomic history of the Poecilioidea . . . . . . . . . . . . . . 2
Anablepidae . . . . . . . . . . . . . . . . . . . . . 3
Poeciliinae . . . . . . . . . . . . . . . . . . . . . . 3
Fluviphylax . . . . . . . . . . . . . . . . . . . . . . 5
Aplocheilichthyinae and Procatopodinae . . . . . . . . . . . . 6
∗ Current address: Biology Department D-8, Regis University, 3333 Regis Boulevard, Denver, CO
80221, U.S.A. E-mail: mghedott@regis.edu
1
0024–4082/00/090001+53 $35.00/0  2000 The Linnean Society of London
2 M. J. GHEDOTTI
Methods . . . . . . . . . . . . . . . . . . . . . . . . 8
Results . . . . . . . . . . . . . . . . . . . . . . . . 9
Transformation series . . . . . . . . . . . . . . . . . . 9
Phylogenetic analysis . . . . . . . . . . . . . . . . . . 37
Taxonomy . . . . . . . . . . . . . . . . . . . . . . 37
Discussion . . . . . . . . . . . . . . . . . . . . . . . 39
Acknowledgements . . . . . . . . . . . . . . . . . . . . 43
References . . . . . . . . . . . . . . . . . . . . . . . 44
Appendix 1: Material examined . . . . . . . . . . . . . . . . 48
Appendix 2: Data matrix . . . . . . . . . . . . . . . . . . 50
Appendix 3: Diagnostic synapomorphies . . . . . . . . . . . . . 53
INTRODUCTION
The superfamily Poecilioidea is a diverse group of cyprinodontiform fishes com-

posed of approximately 300 species (Nelson, 1994) that live in temperate and tropical
regions of the New World and Africa. This group includes well-known aquarium
species such as the guppy (Poecilia reticulata), the mollies (Poecilia spp.), and the
swordtails (Xiphophorus spp.). These familiar species and other poecilioids are important
in research fields such as genetics, oncology, ecology, and developmental biology
(Meffe & Snelson, 1989). This group also includes the anatomically distinctive
foureyed fishes (Anableps spp.) which have dorsally protruding eyes and a horizontally
divided pupil that allows simultaneous aquatic and aerial vision. Most poecilioid
fishes live in shallow brackish or freshwater environments, often associated with
vegetation and near the shoreline. However, two species are pelagic in fresh waters,
living in offshore areas of the African Great Lakes.
Approximately two thirds of poecilioid species have internal fertilization and are
viviparous, and one species (Tomeurus gracilis) has internal fertilization and is fac-
ultatively viviparous, laying fertilized eggs or giving birth to free-swimming young
(Breder & Rosen, 1966). The remaining species exhibit external fertilization and
oviparity. Based on the phylogenetic studies of Parenti (1981), Meyer and Lydeard
(1993), Parker (1997), and Costa (1998) it is likely that viviparity and internal
fertilization evolved twice within the Poecilioidea, once each in the families An-
ablepidae and Poeciliidae. The phylogenetic position of Tomeurus gracilis within the
Poeciliinae is controversial, and this species has been thought to exhibit a reproductive
pattern retained from an early stage in the evolution of poeciliine viviparity (Rosen
& Bailey, 1963; Meyer & Lydeard, 1993). The purposes of this study are to
formulate a phylogenetic hypothesis of relationships of poecilioid fishes, to provide
the foundation of a new taxonomy based on monophyletic taxa, and to use the
phylogeny to interpret patterns of the evolution of within the superfamily.
TAXONOMIC HISTORY OF THE POECILIOIDEA
Parenti (1981) first recognized the superfamily Poecilioidea with its present
composition, Anableps, Jenynsia, Oxyzygonectes, Fluviphylax, Aplocheilichthys spilauchen,
the procatopodines, and the poeciliines (=poeciliids sensu stricto) (Fig. 1A). Close
evolutionary relationships between Anableps and Jenynsia (Hubbs, 1924; Foster 1967),
Anableps and the poeciliids (Turner, 1950; Miller, 1979), and Fluviphylax and the
POECILIOID PHYLOGENY 3
procatopodines (Roberts, 1970) had been proposed previously. However, the in-
clusion of all of these taxa in a larger unit was new to Parenti’s (1981) phylogenetic
treatment of the order. Monophyly of the Poecilioidea (sensu Parenti, 1981) sub-
sequently has been supported by studies using DNA sequence data (Meyer &
Lydeard, 1993), morphological data (Costa, 1998), and combined DNA sequence
and morphological data (Parker, 1997). However, Meyer and Lydeard (1993) did
not support the monophyly of the Poeciliidae sensu Parenti (1981). Their analysis
supports a sister group relationship between the Anablepidae and the Poeciliinae as
well as a sister group relationship between Fluviphylax and the procatopodines (Fig.
1B).
Anablepidae
The Family Anablepidae currently contains three genera of fishes distributed in

Central and South America. Traditionally, the family Anablepidae was comprised
of only the viviparous foureyed fishes, genus Anableps. Members of the genus Jenynsia,
commonly called the onesided livebearers, were recognized in their own family
( Jenynsiidae) and the white eye, Oxyzygonectes dovii, was one of many oviparous taxa
placed in the family Cyprinodontidae ( Jordan, 1923; Myers, 1931; Greenwood et
al., 1966). One notable exception to this traditional classification was Hubbs (1924)
who, following previous suggestions that the Anablepidae and the Jenynsiidae were
closely related (Regan, 1911; Jordan, 1923), placed Anableps and Jenynsia together in
the family Anablepidae. The inclusion of Jenynsia in the Anablepidae was not
accepted by most subsequent authors (Myers, 1931; Rosen, 1964; Greenwood et al.,
1966). A close relationship between Anableps and Jenynsia was rejected by Turner
(1950) and Miller (1979), who proposed a close relationship between Anableps and
the Poeciliinae (=Poeciliidae sensu stricto). Parenti (1981) recognized the family
inclusive of the genera Anableps, Jenynsia, and Oxyzygonectes based on a phylogenetic
analysis of the Cyprinodontiformes (Fig. 1A). Meyer and Lydeard (1993) corroborated
a sister-group relationship between Anableps and Jenynsia using DNA sequence data
(Fig. 1B). Based on a phylogenetic analysis of all described anablepid taxa using
morphological data, Ghedotti (1998) recognized the family with its current limits
and supported the phylogenetic relationships within the family proposed by Parenti
(1981).
Poeciliinae
The poeciliine fishes are familiar to many people and include common aquarium
fishes such as the guppy (Poecilia reticulata), the mollies (Poecilia sp.), the swordtails
(Xiphophorus sp.), and the mosquitofishes (Gambusia sp.). The poeciliine fishes have
long been recognized as a monophyletic group based on the possession of a viviparous
mode of reproduction and a gonopodium composed of anal-fin rays three, four,
and five in adult males. The taxonomic history of genera within this primarily
Neotropical group is complex, with many generic names being proposed and
synonymized. However, the placement of the knife livebearers, genus Alfaro, and
the facultatively viviparous species Tomeurus gracilis has been controversial. Regan
(1913) completed the first comprehensive revision of the Poeciliinae using gonopodial
4 M. J. GHEDOTTI
Oxyzygonectes
Jenynsia* Anablepidae
Anableps*
Poeciliinae* Poeciliinae
Fluviphylax Fluviphylacinae
Aplocheilichthys
Lamprichthys
Pantanodon Aplocheilichthyinae
'Aplocheilichthys'
Hypsopanchax
A Procatopus
47 Aplocheilichthys Aplocheilichthyinae
Fluviphylax Fluviphylacinae
43 58 Jenynsia* Anablepidae
57 Anableps*
Tomeurus*
88
81 Cnesterodon*
Poeciliinae
53 Poecilia*
Xiphophorus*
B
Figure 1. Phylogenetic relationships of the superfamily Poecilioidea as depicted in: A, Parenti (1981);
B, Meyer & Lydeard (1993). Bootstrap values are given above branches. Groups supported by bootstrap
replicate values below 50% are indicated by dashed lines.
characters and provided a synopsis and key to the genera. Subsequently, Hubbs
(1924) recognized four subfamilies (Tomeurinae, Gambusiinae, Poeciliinae, and
Poeciliopsinae) and nine tribes within the Poeciliidae (=Poeciliinae sensu Parenti,
1981). Hubbs (1924) stated that the Gambusiinae are the least specialized poeciliids
whereas Tomeurus gracilis is the most specialized. In 1926, Hubbs moved Alfaro from
the Tomeurinae to the Poeciliinae leaving Tomeurus as the sole genus in the
Tomeurinae. Howell and Hubbs (1936) recognized Alfaro as distinct from both
Tomeurus and the Poeciliinae and classified this genus it in its own subfamily, Alfarinae.
This taxonomic decision was supported by Rosen (1952) and Rosen and Gordon
(1953).
In 1963, Rosen and Bailey published their classic revision of poeciliid fishes. This
work separated the Poeciliidae into two subfamilies, Tomeurinae for Tomeurus and
Poeciliinae for all other poeciliids including Alfaro, which was recognized in the tribe
Poeciliini. This work remains the systematic and taxonomic authority on the group.
Taxonomic studies and phylogenetic analyses subsequent to Rosen and Bailey (1963)
have provided insight into the relationships of smaller groups of poeciliine fishes
(e.g., Rosen, 1967, 1979; Rosa & Costa, 1993; Meyer, Morrissey & Schartl, 1994;
Mojica, Meyer & Barlow, 1997; Rodriguez, 1997), but these studies did not alter
the higher level classification of the family. Parenti (1981) recognized these fishes as
a subfamily (Poeciliinae) within a more inclusive family (Poeciliidae) including the
oviparous Aplocheilichthyinae and Fluviphylacinae. Parenti and Rauchenberger
(1989) modified the classification of Rosen and Bailey (1963) to reflect the change
in taxonomic rank proposed by Parenti (1981) (Table 1).
T 1. Classification of the subfamily Poeciliinae proposed by Parenti and Rauchenberger (1989)
(based largely on Rosen & Bailey, 1963). ∗Genera with species included in this study
Subfamily Poeciliinae Garman, 1895

Supertribe Tomeurini Hubbs, 1924
Genus Tomeurus Eigenmann, 1909∗
Supertribe Poeciliini Garman, 1895
Tribe Poeciliini Garman, 1895
Genus Alfaro Meek, 1912∗
Genus Poecilia Bloch & Schneider, 1801∗
Genus Priapella Regan, 1913∗
Genus Xiphophorus Heckel, 1848
Tribe Cnesterodontini Hubbs, 1924
Genus Phallotorynus Henn, 1916∗
Genus Phalloceros Eigenmann, 1907∗
Genus Phalloptychus Eigenmann, 1907
Genus Cnesterodon Garman, 1895∗
Tribe Scolichthyini Rosen, 1967
Genus Scolichthys Rosen, 1967
Tribe Gambusini Hubbs, 1924
Genus Brachyraphis Regan, 1913
Genus Gambusia Poey, 1854∗
Genus Belonesox Kner, 1860
Tribe Girardini Hubbs, 1924
Genus Girardinus Poey, 1854∗
Genus Quintana Hubbs, 1934
Genus Carlhubbsia Whitley, 1951
Tribe Heterandrini Hubbs, 1924
Genus Priapichthys Regan, 1913
Genus Neoheterandria Henn, 1916
Genus Heterandria Agassiz, 1853∗
Genus Poeciliopsis Regan, 1913∗
Genus Phallichthys Hubbs, 1924∗
Tribe Xenodexini Hubbs, 1950
Genus Xenodexia Hubbs, 1950
Fluviphylax
The genus Fluviphylax includes the smallest known cyprinodontiform fishes, with
a maximum length of approximately 22.0 mm (Weitzman, 1982; Weitzman & Vari,
1988). The members of this diminutive oviparous, Neotropical genus have a relatively
short taxonomic history, with the first species described in 1955. Myers and Carvalho
in Myers (1955) described the first species as Potamophylax pygmaeus. Whitley (1965)
pointed out that Potamophylax was preoccupied in Insecta and proposed Fluviphylax
as a replacement name. The osteology of Fluviphylax was reviewed by Roberts (1970),
and he proposed its placement in a monotypic subfamily, Fluviphylacinae. He
suggested that Fluviphylax was related to either the Procatopodinae or the Fundulinae
and noted a similarity between the cephalic squamation of Fluviphylax and Pantanodon.
The Fluviphylacinae was placed within an expanded Poeciliidae by Parenti (1981)
(Fig. 1). This relationship was supported and clarified by Costa (1996) who placed
Fluviphylax in a new tribe, Fluviphylacini, within the Aplocheilichthyinae, described
three new species (F. simplex, F. zonatus, and F. obscurum), and proposed a generic
phylogeny based on morphological data.
6 M. J. GHEDOTTI
Aplocheilichthyinae and Procatopodinae
The subfamilies Aplocheilichthyinae and Procatopodinae contain more than 100

oviparous species distributed in sub-Saharan Africa and the Nile Valley. The species-
level taxonomy of these groups is poorly understood and it is likely that many species
remain undescribed. Regan (1911) first suggested a close relationship among many
of the fishes currently recognized in the Aplocheilichthyinae or the Procatopodinae.
He expanded the genus Aplocheilichthys to include many species previously placed in
the genus Aplocheilus and suggested a close relationship among Aplocheilichthys, Pro-
catopus, and Lamprichthys. Fowler (1916) described two new subfamilies, Procatopinae
and Lamprichthyinae, for Procatopus and Lamprichthys (a distinctive genus of pelagic
fishes endemic to Lake Tanganyika) respectively. Myers (1924b) disagreed with this
subfamilial taxonomy and stated that Procatopus was not distinctive enough to be
given subfamily rank as Fowler (1916) had done.
In 1928, Myers described the tribe Aplocheilichthyini, for the genera Aplocheilichthys,
Procatopus, Plataplocheilus, Hypsopanchax, Platypanchax, and Cynopanchax and recognized
Lamprichthys in the subfamily Lamprichthyinae. Myers (1938) reviewed the taxonomy
of the genera included in his Aplocheilichthyini. In 1955, he introduced the subfamily
name Procatopodinae as a replacement for Aplocheilichthyini without explanation.
Procatopodinae probably was proposed as a corrected formation of the family
name Procatopinae of Fowler (1916). Myers (1955) did not mention the subfamily
Lamprichthyinae. This could suggest that he placed Lamprichthys within the Pro-
catopodinae. However, a reprint of this paper at the California Academy of Sciences
that was annotated by Dr Myers indicates that the exclusion of Lamprichthyinae
from this work was an oversight (Parenti, pers. com.). Use of the senior name
Procatopodinae for a subfamily including Procatopus, Aplocheilichthys, and other genera
was followed by many subsequent authors (e.g. Whitehead, 1962; Foster, 1967;
Clausen, 1967). Many authors (Sethi, 1960; Foster, 1967; Clausen, 1967) recognized
a close relationship among Aplocheilichthys spilauchen, the procatopodines, and Lam-
prichthys. Clausen (1967) placed Lamprichthys within the subfamily Procatopodinae
and suggested that Lamprichthys is most closely related to Plataplocheilus and Procatopus.
The relationships of the genus Pantanodon have been controversial for much of its
history. Myers (1955) created a new subfamily, Pantanodontinae, to contain Pantanodon
podoxys, a toothless cyprinodontiform, which he described in the same work. White-
head (1962) also placed Pantanodon in Pantanodontinae and suggested that it was an
aberrant derivative of the Procatopodinae. Rosen (1965) subsequently moved a
tooth-bearing species, Oryzias madagascariensis, from the genus Oryzias to the genus
Pantanodon, and suggested elevating the Pantanodontinae from subfamilial to familial
rank because its relationship to other cyprinodontiforms was unclear. Foster (1967)
supported a relationship between Pantanodon and Oryzias. However, Clausen (1967)
suggested a relationship between the Pantanodontinae and the Procatopodinae
(especially Procatopus).
Parenti (1981) recognized Aplocheilichthyinae Myers (1928) as the valid family
group name for Aplocheilichthys, Lamprichthys, Pantanodon, Hypsopanchax, Procatopus,
Plataplocheilus, and Cynopanchax (instead of Procatopodinae Fowler, 1916, as corrected
by Myers, 1955) (Table 2). At the generic level Parenti (1981) placed the genus
Hylopanchax in the synonymy of Procatopus and noted that the large genus Aplocheilichthys
as then recognized was not monophyletic (Fig. 1A). In 1982, Huber proposed a new
taxonomy for these fishes and described a new genus, Laciris, to contain Aplocheilichthys
T 2. Classification of the taxa included in the subfamily Aplocheilichthyinae as proposed by

Parenti (1981) and Huber (1982). The type species follows the generic names in parentheses
PARENTI, 1981
Subfamily Aplocheilichthyinae Myers, 1928
Genus Aplocheilichthys Bleeker, 1863
Subgenus Aplocheilichthys Bleeker, 1863 (A. typus Bleeker, 1863, a junior synonym of Poecilia spilauchen
Dumeril, 1861)
Subgenus Micropanchax Myers, 1924a (Haplochilus schoelleri Boulenger, 1904, a junior synonym of H. loati
Boulenger, 1901)
Subgenus Lacustricola Myers, 1924b (Haplochilus pumilis Boulenger, 1906)
Subgenus Poropanchax Clausen, 1967 (Aplocheilichthys macropthalmus Meinken, 1932)
Subgenus Congopanchax Poll, 1971 (Aplocheilichthys myersi Poll, 1952)
Genus Lamprichthys Regan, 1911 (Haplochilus tanganicus Boulenger, 1898)
Genus Pantanodon Myers, 1955 (P. podoxys Myers, 1955)
Genus ‘Aplocheilichthys’ (no type, includes Haplocheilus johnstoni Günther, 1893)
Genus Procatopus Boulenger, 1904 (P. nototaenia Boulenger, 1904)
Genus Hypsopanchax Myers, 1924b (Haplochilus platysternus Nichols and Griscom, 1917)
Genus Cynopanchax Ahl, 1928 incertae sedis (Haplocheilichthys bukobanus Ahl, 1928)
Genus Plataplocheilus Ahl, 1928 incertae sedis (Haplocheilichthys ngaensis Ahl, 1928)
HUBER, 1982 (Article dated 1981 but validly published in 1982. See Lazara, 1983)
Genus Aplocheilichthys Bleeker, 1863
Genus Micropanchax Myers, 1924a
Subgenus Micropanchax Myers, 1924a
Subgenus Lacustricola Myers, 1924b
Subgenus Poropanchax Clausen, 1967
Subgenus Congopanchax Poll, 1971
Subgenus Cynopanchax Ahl, 1928
Genus Lamprichthys Regan, 1911
Genus Pantanodon Myers, 1955
Genus Procatopus Boulenger, 1904
Subgenus Procatopus Boulenger, 1904
Subgenus Plataplocheilus Ahl, 1928
Genus Hypsopanchax Myers, 1924b
Genus Laciris Huber, 1982 (Aplocheilichthys pelagicus Worthington, 1932)
pelagicus, a pelagic species endemic to Lake Edward. Huber (1982) recognized

Plataplocheilus as a subgenus within Procatopus and elevated Micropanchax from a
subgenus of Aplocheilichthys to generic rank (Table 2). Huber (1982) recognized
Aplocheilichthys as a monotypic genus containing A. spilauchen, and placed all other
species previously in Aplocheilichthys in the genus Micropanchax.
Little taxonomic or systematic work above the species level has occurred since
these two revisions. Lazara (1983) reviewed the taxonomy of the Aplocheilichthyinae
(sensu Parenti, 1981) and Wildekamp (1995) reviewed the composition of the genus
Aplocheilichthys. Lazara (1983) and Wildekamp (1995) did not adopt most of the
taxonomic conclusions of Huber (1982). Most recently, Costa (1996) placed the
Neotropical genus Fluviphylax within the Aplocheilichthyinae (sensu Parenti, 1981)
and recognized two tribes within the subfamily, Fluviphylacini and Aplocheilichthyini,
to contain Fluviphylax and all other aplocheilichthyines (sensu Parenti, 1981) re-
spectively.
The previous use of the family group name Aplocheilichthyinae Myers (1928) for
a group composed of Aplocheilichthys, Pantanodon, Lamprichthys, Procatopus, and the other
African lamp-eyed killifishes (e.g. Parenti, 1981; Huber, 1982; Lazara, 1983; Nelson,
1994; Costa, 1996) was in error because Procatopodinae Fowler (1916) has priority.
8 M. J. GHEDOTTI
However, I recognize both family group names as separate, subfamilial-rank taxa.

Aplocheilichthys spilauchen is the sole member of the Aplocheilichthyinae Myers (1928)
and Fluviphylax together with all other species previously placed in the Aplo-
cheilichthyinae (sensu Parenti, 1981) is recognized as the Procatopodinae Fowler
(1916).
METHODS
Specimens examined in this study are listed in Appendix 1. Standard institutional

symbolic codes for specimens are as listed in Leviton et al. (1985). Specimens
examined were either whole alcohol preserved specimens or cleared and stained for
bone and cartilage following the procedure of Dingerkus and Uhler (1977). Dissection
of cleared and stained specimens followed the method outlined by Weitzman (1974),
except that the branchial basket was removed before the suspensorium. Specimens
were examined and illustrated using a Zeiss stereomicroscope with an attached
camera lucida.
Polarity of transformation series was determined by outgroup comparison.
Crenichthys baileyi, Cubanichthys pengelleyi, Cyprinodon variegatus, Fundulus chrysotus,
Profundulus labialis, Valencia hispanica, and V. letourneuxi were chosen as outgroups
based on the ordinal level phylogenies proposed by Parenti (1981), Meyer and
Lydeard (1993), and Costa (1998). Transformation series that vary solely among
outgroups are not within the scope of this work and were not included. Therefore,
outgroup topology was constrained based on the topologies proposed in Parenti
(1981), Meyer and Lydeard (1993), and Costa (1998) (Fig. 2). A separate analysis
was run for each outgroup topology. Ingroup taxa were chosen as representatives
of genera or other higher taxa based on availability and taxonomic significance
(Table 3).
Data were compiled and formatted as a data matrix of 36 taxa and 99
transformation series (Appendix 2). Character states were coded as integers and
‘0’ was arbitrarily assigned to the character state in Profundulus labialis. Question
marks were used for inapplicable coding and are discussed in each transformation
series account. Parsimony analysis using the heuristic search option of PAUP
v3.1.1 (Swofford, 1993) was used to analyse the data matrix. Polymorphic
transformation series were recognized as such using the polymorphic character
state option. Five hundred random addition sequence replicates using the tree
bisection-reconstruction and branch-swapping options were performed to minimize
the possibility that the heuristic search would yield a result that represented a
less than most parsimonious set of trees. Decay indices (Bremer, 1988) were
calculated using the program TreeRot (Sorensen, 1996). The distribution
of character states was examined using both ACCTRAN and DELTRAN
optimization.
Transformation series with character states that uniquely or unambiguously
support a node in all equally most parsimonious phylogenetic trees are considered
diagnostic. Features may still be considered diagnostic if they are reversed or
independently acquired in other groups. Autapomorphic transformation series and
transformation series varying solely among outgroup taxa were not included in the
phylogenetic analysis.
Profundulus labialis
Fundulus chrysotus
Valencia hispanica
Valencia letourneauxi
Cyprinodon variegatus
Cubanichthys pengelleyi
Crenichthys baileyi
A INGROUP
Fundulus chrysotus
Crenichthys baileyi
B INGROUP
Fundulus chrysotus
Crenichthys baileyi
Valencia hispanica
Valencia letourneauxi
C INGROUP
Figure 2. Constrained outgroup topologies: A, based on Parenti (1981); B, based on Meyer & Lydeard
(1993); C, based on Costa (1998). Valencia was not included in analyses using the outgroup topology
illustrated in B because this taxon was not included in Meyer & Lydeard’s (1993) analysis.
RESULTS
Transformation series
Transformation series are grouped into traditional anatomical units and assigned
numbers, as in Appendix 2. Each account includes: (1) a description of the character
state with the coding used in the data matrix, (2) a discussion of the historical usage
of each transformation series in reference to poecilioid taxa (if relevant), and (3) a
description of the distribution of either the apomorphic or plesiomorphic character
state(s). Taxonomic names follow the taxonomy proposed in this work (Table 4).
When a character state is said to be present in all members of a group (e.g. all
poeciliines) the statement refers only to those representative taxa included in this
study unless stated otherwise. The consistency index (ci) and retention index (ri) for
each transformation series were calculated using the outgroup topology proposed
by Costa (1998) (Fig. 2C). These values appear in parentheses at the end of each
account separated by a colon (ci: ri). When both indices are equal, only one number
was provided.
Neurocranium
1. Mesethmoid: (0) ossified (Fig. 3A,D,E); (1) cartilaginous (Fig. 3C). Parenti (1981)
discussed a cartilaginous mesethmoid as diagnostic of the subfamily Aplo-
cheilichthyinae, including Aplocheilichthys spilauchen. However, all specimens of A.
spilauchen I examined have an ossified mesethmoid (Fig. 3A). The mesethmoid of all
procatopodines examined is cartilaginous. Parenti (1981) also used a cartilaginous
10 M. J. GHEDOTTI
T 3. The 27 ingroup taxa included in this study placed in a classification based on Parenti (1981)
and Parenti and Rauchenberger (1989). Subgenera of Aplocheilichthys are placed in parentheses
Superfamily Poecilioidea
Family Anablepidae
Subfamily Oxyzygonectinae
Oxyzygonectes dovii
Subfamily Anablepinae
Anableps dowi
Jenynsia unitaenia
Jenynsia multidentata
Family Poeciliidae
Subfamily Aplocheilichthyinae
Aplocheilichthys (Aplocheilichthys) spilauchen
Aplocheilichthys (Micropanchax) loati
Aplocheilichthys (Lacustricola) pumilis
Aplocheilichthys (Poropanchax) macropthalmus
Aplocheilichthys schioetzi
Aplocheilichthys pelagicus
Lamprichthys tanganicus
Pantanodon madagascariensis
‘Aplocheilichthys’ johnstoni
Procatopus nototaenia
Hypsopanchax zebra
Hypsopanchax cf. modestus
Subfamily Fluviphylacinae
Fluviphylax obscurum
Subfamily Poeciliinae
Supertribe Tomeurini
Tomeurus gracilis
Supertribe Poeciliini
Alfaro cultratus
Priapella compressa
Poecilia sphenops
Phallotorynus victoriae
Phalloceros caudimaculatus
Cnesterodon decemmaculatus
Gambusia affinis
Girardinus metallicus
Heterandria formosa
Poeciliopsis latidens
Phallichthys amates
mesethmoid to diagnose the suborder Aploceiloidei. However, Tigano and Parenti

(1988) suggested that the bone interpreted by Parenti (1981) as a vomer in aplo-
cheiloids was more correctly a compound bone composed of a coossified dermal
vomer and ventral mesethmoid. The mesethmoid is cartilaginous in all pro-
catopodines examined and Cyprinodon variegatus. (0.50:0.90)
2. Anterior margin of frontals: (0) straight or slightly cleft medially (Fig. 3E); (1)
extend anteriorly between nasals (Fig. 3A,C,D). The anterior margin of the frontals
extends anteriorly in Oxyzygonectes dovii, Aplocheilichthys spilauchen, Alfaro cultratus, Priapella
compressa, and all procatopodines examined except Fluviphylax pygmaeus and Hyp-
sopanchax zebra. (0.20:0.70)
3. Parietals: (0) large and elongate, contacting the sphenotic anteriorly (Fig. 3D);
(1) short and compact, restricted to the epiotic region, not contacting the sphenotic
anteriorly (Fig. 3E); (2) absent (Fig. 3A,C). Parenti (1981) recognized the absence of
T 4. Proposed classification of the superfamily Poecilioidea based on the phylogenetic hypothesis
presented in Figure 19. Genera with members in this study are indicated by an asterisk (∗). Approximate
number of Recent species are in parentheses
Superfamily Poecilioidea
Family Anablepidae Bonaparte, 1831
Subfamily Oxyzygonectinae Parenti, 1981
Oxyzygonectes ∗(1)
Subfamily Anablepinae Bonaparte, 1831
Anableps ∗(3), Jenynsia ∗(9)
Family Poeciliidae Garman, 1895
Aplocheilichthys ∗(1)
Subfamily Procatopodinae Fowler, 1916
Tribe Fluviphylacini Roberts, 1970
Fluviphylax ∗(4)
Tribe Procatopodini Fowler, 1916
Micropanchax ∗(35) with three subgenera (Micropanchax, Lacustricola, and Poropanchax),
‘Micropanchax’ (currently unnamed genus)∗(4), Platypanchax ∗(2), Lamprichthys ∗(1), Pantanodon
∗(2), Hypsopanchax ∗(5), Procatopus ∗(3), Cynopanchax (1), Plataplocheilus (4)
Subfamily Poeciliinae Garman, 1895
Tribe Alfarini Hubbs, 1924
Alfaro ∗(2)
Tribe Priapellini new tribe
Priapella ∗(3)
Tribe Gambusini Hubbs, 1924
Brachyraphis (9), Gambusia ∗(45), Belonesox (1)
Tribe Heterandrini Hubbs, 1924
Priapichthys (7), Neoheterandria (4), Heterandria ∗(1), Pseudoxiphophorus (8), Poeciliopsis ∗(21),
Tribe Girardini Hubbs, 1924
Girardinus ∗(8), Quintana (1), Carlhubbsia (2)
Tribe Poeciliini Garman, 1895
Poecilia ∗(25), Pamphorichthys (6), Limia (20), Xiphophorus (17), Phallichthys ∗(4)
Tribe Cnesterodontini Hubbs, 1924
Phallotorynus ∗(3), Phalloceros ∗(1), Phalloptychus (2), Cnesterodon ∗(4), Tomeurus ∗(1)
Tribe Scolichthyini Rosen, 1967
Scolichthys (2)
Tribe Xenodexini Hubbs, 1950
Xenodexia (1)
parietals as synapomorphic for all cyprinodontids except Cubanichthys and potentially

synapomorphic of Fluviphylax, Aplocheilichthys spilauchen and the procatopodines. Pa-
rietals are present but reduced in size in Hypsopanchax zebra, Tomeurus gracilis, Phalloceros
caudimaculatus, Poecilia sphenops, Girardinus metallicus, and Phallichthys amates. Parietals
are absent in Aplocheilichthys spilauchen, Cnesterodon decemmaculatus, Poeciliopsis latidens,
and all procatopodines except Hypsopanchax zebra. Phallotorynus victoriae is polymorphic,
with some individuals lacking parietals and others having small parietals (1&2). (0.40:
0.70)
4. Epiotic (epioccipital) processes in adults: (0) short, not extending beyond first
vertebra (Fig. 3D); (1) absent (Fig. 3C,E); (2) long, extending beyond first vertebra (Fig.
3A). Parenti (1981) used expanded epiotic processes as an anablepid synapomorphy.
Ghedotti (1998) did not recognize expanded epiotic processes as uniquely sy-
napomorphic of the Anablepidae and noted their presence in Anableps, Oxyzygonectes,
three species of Jenynsia, and Aplocheilichthys spilauchen. Epiotic processes are absent
in all procatopodines examined, Tomeurus gracilis, Cnesterodon decemmaculatus, Phallotorynus
victoriae, Cubanichthys pengelleyi, Cyprinodon variegatus, Valencia, and Crenichthys baileyi. Long
12 M. J. GHEDOTTI
premaxilla 5-1
vomer
maxilla
mesethmoid lateral ethmoid
nasal frontal
parasphenoid
frontal
pterosphenoid
dermosphenotic sphenotic
sphenotic prootic
pterotic
pterotic
supraoccipital 10-1
exoccipital exoccipital
epiotic process intercalar bassioccipital
A 4-2 B
13-1 13-0
1-0
1-1
14-0
2-1
2-0
3-2 3-0
4-1
4-0 3-1
C D E
6-0
6-1 5-0
8-1 8-0
9-0
10-0
F 43-0 G 43-1 H
Figure 3. Neurocrania in: A, Aplocheilichthys spilauchen, dorsal view, USNM 304139; B, Aplocheilichthys
spilauchen, ventral view, USNM 304139; C, Micropanchax pelagicus, dorsal view, FMNH 26956; D, Alfaro
cultratus, dorsal view, KU 11122; E, Tomeurus gracilis, dorsal view, UMMZ 167944; F, Micropanchax
pelagicus, ventral view, FMNH 26956; G, Alfaro cultratus, ventral view, KU 11122; H, Tomeurus gracilis,
ventral view, UMMZ 167944. Cartilage is indicated by uniform shading. Rostral cartilage is not
illustrated. Maxillae and premaxillae are not illustrated in ventral view. Arrows indicate character
states discussed in the text. Scale bars=1 mm.
epiotic processes are present in Oxyzygonectes dovii, Anableps dowi, Jenynsia unitaenia,
Aplocheilichthys spilauchen, Girardinus metallicus, Poecilia sphenops, and Fundulus chrysotus.
(0.20:0.25)
5. Dorsolateral processes of vomer: (0) do not contact lateral ethmoids (Fig. 3G,H);
(1) contact lateral ethmoids (Fig. 3B). Ghedotti (1998) noted contact between the
lateral ethmoids and the vomer in Aplocheilichthys spilauchen, Anableps, and all Jenynsia
except J. pygogramma. Species with unossified vomers (Fig. 3F) are coded with a
question mark (?). The vomer contacts the lateral ethmoids in all Anableps, Jenynsia,
Aplocheilichthys spilauchen, Valencia, and Fundulus chrysotus. (0.25:0.50)
6. Vomer: (0) ossified (Fig. 3B,G,H); (1) unossified (Fig. 3F). Parenti (1981) used an
unossified vomer to diagnose both Fluviphylax and a group composed of Micropanchax
johnstoni (‘Aplocheilichthys’ johnstoni sensu Parenti, 1981), Pantanodon, Hypsopanchax, and
Procatopus. The vomer is unossified in Micropanchax macropthalmus, M. johnstoni, M.
pelagicus, ‘M.’ schioetzi, Pantanodon madagascariensis, Hypsopanchax zebra, Procatopus no-
totaenia, and Fluviphylax obscurum. The vomer may be partially ossified in some
individuals of M. loati and this species was coded as polymorphic (0&1) for this
transformation series. (0.50:0.71)
7. Parasphenoid in region of orbit: (0) straight; (1) strongly curved ventrally. Many
procatopodine species have large orbits, however, the parasphenoid is strongly
curved ventrally and is coded so only in Micropanchax macropthalmus and Lamprichthys
tanganicus. (0.50:0.00)
8. Ascending processes of parasphenoid in adults: (0) long, contact pterosphenoids
(Fig. 3B,G); (1) short, do not contact pterosphenoids (Fig. 3F,H). Roberts (1970)
cited the presence of short anterolateral processes (=ascending processes) of the
parasphenoid as evidence of relationship between Fluviphylax and the procatopodines.
Costa (1996) recognized this character state as a synapomorphy of the Aplo-
cheilichthyinae sensu Costa (1996) (Aplocheilichthys spilauchen, Fluviphylax, and the
procatopodines). Short anterolateral processes of the parasphenoid are present in
Fluviphylax obscurum, Tomeurus gracilis, Cnesterodon decemmaculatus, Cubanichthys pengelleyi,
Cyprinodon variegatus, and all procatopodines examined except Platypanchax cf. modestus,
Hypsopanchax zebra, and Procatopus nototaenia. (0.33:0.64)
9. Prootic bridge over lateral canal and trigeminofacialis chamber of prootic: (0)
narrow (Fig. 3); (1) broad (Ghedotti, 1998: Fig. 4B,C); (2) very narrow, forming a
distinct vertical flange. Ghedotti (1998) recognized a broad prootic bridge as
synapomorphic of the Anablepidae with a reversal in Jenynsia alternimaculata. The
prootic bridge is broad in all anablepids examined and Crenichthys baileyi. The prootic
bridge forms a distinct vertical flange in Pantanodon madagascariensis, Hypsopanchax
zebra, and Procatopus nototaenia. (0.66:0.74)
10. Intercalar (opisthotic): (0) small or absent, if present, restricted to point of
attachment of the lower limb of posttemporal (Fig. 3F–H); (1) large, elongate,
extending laterally beyond point of attachment of the lower limb of posttemporal
(Fig. 3B). Ghedotti (1998) discussed the previous treatment of this element in
Cyprinodontiformes (Rosen & Bailey, 1963; Dyer & Chernoff, 1996) and recognized
enlarged intercalars in Aplocheilichthys spilauchen and all anablepids except Jenynsia
pygogramma. I found well-developed intercalars, often extending onto the adjacent
pterotic, in Aplocheilichthys spilauchen, Pantanodon madagascariensis, Hypsopanchax zebra,
Procatopus nototaenia, and all anablepids examined. (0.33:0.71)
Jaws and suspensorium

11. Teeth in outer row on dentary and premaxilla, at least in embryos and young
individuals: (0) unicuspid with distinct point and strongly compressed; (1) unicuspid
14 M. J. GHEDOTTI
and conical; (2) bicuspid; (3) tricuspid; (4) rounded and strongly compressed, without
a distinct point. Tooth form has been used extensively in the identification and
classification of cyprinodontiform fishes (Günther, 1866; Garman, 1895; Regan
1911; Ahl, 1924, 1928). The phylogenetic importance of tooth form was minimized
by other precladistic authors in favor of reproductive characteristics (e.g. Regan,
1913; Rosen & Bailey, 1963). Parenti (1981) used tricuspid teeth in the outer row
as a synapomorphy of the Anablepidae. Ghedotti (1998) who recognized a reversal
to conical teeth in Anableps anableps and A. microlepis confirmed this synapomorphy.
Aplocheilichthys spilauchen, all procatopodines examined, Gambusia affinis, Alfaro cultratus,
Tomeurus gracilis, Valencia, and Fundulus chrysotus have conical outer teeth. Crenichthys
bailey has bicuspid outer teeth. Jenynsia unitaenia, J. multidentata, and Cyprinodon variegatus
have tricuspid outer teeth. Oxyzygonectes dovii, young Anableps dowi, and Cubanichthys
pengelleyi have outer teeth that are variably tricuspid or compressed and are coded
as polymorphic (0&3). The lateral cusps on the tricuspid teeth are less prominent
in these three species. (0.70:0.75)
12. Teeth in inner rows on dentary and premaxilla, at least in embryos and young
individuals: (0) unicuspid and conical or slightly compressed; (1) tricuspid; (2) absent.
Oxyzygonectes dovii, Jenynsia unitaenia, and J. multidentata have tricuspid inner teeth.
Phallotorynus victoriae, Girardinus metallicus, and Cyprinodon variegatus have only one row
of teeth and lack inner teeth. Young Anableps dowi and Cubanichthys pengelleyi have
inner teeth that are variably tricuspid or compressed and are coded as polymorphic
(0&3). The lateral cusps on the tricuspid teeth are less prominent in these two
species. (0.67:0.50)
13. Medial surface of ascending process of premaxilla: (0) approximately straight
(Fig. 3A,D,E); (1) angled laterally at proximal end, forming triangular space between
proximal ends of ascending processes (Fig. 3C). The medial surface of the ascending
processes is angled laterally in Micropanchax pelagicus, Cnesterodon decemmaculatus, Phal-
loceros caudimaculatus, Phallotorynus victoriae, Girardinus metallicus, Poecilia sphenops, and
Phallichthys amates. Anableps dowi lacks ascending processes of the premaxilla and was
coded as unknown for this transformation series (?). (0.33:0.67)
14. Lateral surface of ascending process of premaxilla: (0) angled medially near
proximal end, forming convex profile (Fig. 3); (1) not angled medially near proximal
end with straight or concave profile. The lateral surface of the ascending processes
is not angled medially near the proximal end in Micropanchax pumilis, ‘M.’ schioetzi,
Platypanchax sp., Lamprichthys tanganicus, Cubanichthys pengelleyi, and Cyprinodon variegatus.
Anableps dowi lacks ascending processes of the premaxilla and was coded as unknown
for this transformation series (?). (0.33:0.60)
15. Teeth on alveolar arm of premaxilla: (0) absent (Fig. 4A,B); (1) present (Fig.
4C,D). Teeth were considered to be present on the alveolar arm when the posterior
teeth on the premaxilla are angled anteriorly due to their position on the more
ventrally directed alveolar arm. Parenti (1981) recognized that the presence of teeth
near the distal tip of the alveolar arm of the premaxilla supports a sister group
relationship between Hypsopanchax and Procatopus. The teeth on the alveolar arm are
most extensive in Procatopus nototaenia, extending to near the distal tip of the alveolar
arm. Pantanodon madagascariensis, Hypsopanchax zebra, and Poeciliopsis latidens have teeth
on only the more proximal portion of the alveolar arm. The teeth on the alveolar
arm are least extensive in Poeciliopsis latidens. (0.50:0.67)
Figure 4. Lateral view of left premaxilla, maxilla, and lacrymal in: A, Aplocheilichthys spilauchen, USNM
304139; B, Micropanchax johnstoni, USNM 246173; C, Procatopus nototaenia, CAS 155515; D, Pantanodon
madagascariensis, AMNH 20526. Arrows indicate character states discussed in the text. Scale bars=
1 mm.
16. Dorsal process of maxilla: (0) small, more narrow anteriorly than at its base; (1)
expanded medially, more broad anteriorly than at its base; (2) greatly reduced,
forming only low ridge or absent. Medially expanded dorsal processes of the maxillae
are present in Anableps dowi, Aplocheilichthys spilauchen, Cubanichthys pengelleyi, Cyprinodon
variegatus, and all procatopodines examined except Hypsopanchax zebra and Procatopus
nototaenia. The dorsal processes are greatly reduced or absent in Pantanodon ma-
dagascariensis, Crenichthys baileyi, and Fundulus chrysotus. (0.25:0.54)
17. Ventral process of anterior maxilla: (0) elongate, extending anteriorly and
medially; (1) greatly reduced, consisting of rounded ventromedial bulge. The ventral
process is greatly reduced in Micropanchax johnstoni and Platypanchax cf. modestus. (0.50:
0:00)
18. Anterior indentation in ventral maxilla: (0) absent, ventral maxilla appears
straight (Fig. 4C,D); (1) present, ventral maxilla appears expanded (Fig. 4A,B).
Parenti (1981) recognized that the reduction of the spatulate distal (=ventral) arm
of the maxilla supports a sister group relationship between Hypsopanchax and Procatopus.
The ventral maxilla is straight, lacking a spatulate portion, in Micropanchax johnstoni,
Platypanchax cf. modestus, Lamprichthys tanganicus, Pantanodon madagascariensis, Hypsopanchax
zebra, Procatopus nototaenia, Phalloceros caudimaculatus, Phallotorynus victoriae, Girardinus
metallicus, Cubanichthys pengelleyi, Cyprinodon variegatus, Crenichthys baileyi, and Profundulus
labialis. (0.13:0.50)
19. Dentary near symphysis: (0) narrow, lacking distinct ventral extension of anterior
dentary (Fig. 5A,B,D,E); (1) deep, distinct ventral extension of anterior dentary
16 M. J. GHEDOTTI
coronomeckelian
dentary
anguloarticular
Meckel's cartilage retroarticular

22-0
A
23-0
B C 24-2
23-1 22-1 24-0 19-1
19-0
21-1
D 21-0 E
Figure 5. Medial view of right lower jaw in: A, Profundulus labialis, KU 18163; B, Aplocheilichthys spilauchen,
CAS 163440; C, Micropanchax johnstoni, USNM 246173; D, Alfaro cultratus, KU 11122; E, Cnesterodon
decemmaculatus, KU 22411. Cartilage is indicated by uniform shading. Arrows indicate character states
discussed in the text. Scale bars=1 mm.
present (Fig. 5C). Parenti (1981) recognized a robust dentary as a synapomorphy

for a clade composed of Micropanchax johnstoni and its close relatives, Hypsopanchax,
and Procatopus. The dentary near the symphysis is deep, with a distinct ventral
expansion in Micropanchax johnstoni, M. loati, M. pelagicus, and ‘M.’ schioetzi. (0.50:0.67)
20. Lateral extension of anterior dentary: (0) absent; (1) present, bearing teeth. The
lateral expansion of the dentary is usually accompanied by a less prominent lateral
expansion of the premaxilla that together form a broad, straight biting surface.
Pantanodon madagascariensis possesses lateral expansions of the dentary but does not
exhibit lateral expansions of the premaxilla. The dentary is expanded laterally in
Pantanodon madagascariensis, Girardinus metallicus, Poecilia sphenops, and Phallichthys amates.
(0.50:0.67)
21. Medially directed flange on posteroventral dentary and anteromedially directed
flange on ventral anguloarticular: (0) absent (Fig. 5A–D); (1) present (Fig. 5E). The
flanges on these two bones were placed in one transformation series because they
do not seem to exhibit independent distributions. The dentary and anguloarticular
have these distinctive medial flanges in all poeciliines examined except Alfaro cultratus,
Priapella compressa, Gambusia affinis, and Tomeurus gracilis. (0.50:0.86)
22. Anterior cleft in anguloarticular: (0) large, extending posteriorly beyond posterior
border of Meckel’s cartilage (Fig. 5A); (1) small, not extending posteriorly beyond
posterior border of Meckel’s cartilage (Fig. 5B–E); (2) absent. Small anterior clefts in
the anguloarticulars are present in Oxyzygonectes dovii, Anableps dowi, all procatopodines
examined except Pantanodon madagascariensis, and all poeciliines examined. Jenynsia
multidentata and Aplocheilichthys spilauchen were coded as polymorphic (0&1). An anterior
anguloarticular cleft is absent in Pantanodon madagascariensis and Fluviphylax obscurum.
(0.67:0.75)
23. Ventral process of anguloarticular: (0) short, ventral process of anguloarticular
does not extend anterior to where anguloarticular overlaps dentary (Fig. 5A); (1)
long, ventral process of anguloarticular extends anterior of where anguloarticular
overlaps dentary (Fig. 5B–E). Parenti (1981) recognized an elongate retroarticular
as synapomorphic for the superfamily Poecilioidea. Costa (1998) modified the
character state description used by Parenti (1981) to recognize the cooccurrence of
a long retroarticular and a long ventral process of the anguloarticular as sy-
napomorphic of Poecilioidea. The condition of the ventral process of the an-
guloarticular and the retroarticular are treated as separate transformation series
here because they vary independently. The ventral process of the anguloarticular is
long in all poecilioids examined. (1)
24. Retroarticular: (0) short, not extending anteriorly to anterior margin of coronoid
process of anguloarticular (Fig. 5A,B,D,E); (1) long and robust, extending anterior
to anterior margin of coronoid process of anguloarticular; (2) long and slender,
extending anterior to anterior margin of coronoid process of anguloarticular (Fig. 5C).
See transformation series 23 for discussion of historical treatment. The retroarticular is
long and somewhat slender in Micropanchax macropthalmus, M. pumilis, M. johnstoni, M.
loati, and M. pelagicus. The retroarticular is long and robust in Gambusia affinis,
Anableps, and all Jenynsia examined except J. unitaenia. (0.50:0.71)
25. Medial process of lacrymal: (0) long and narrow, more anteriorly directed (Fig.
4A,C); (1) short and broad, more medially directed (Fig. 4B,D). Short and broad
medial processes of the lacrymals are present in Cyprinodon variegatus and all poeciliids
examined except Aplocheilichthys spilauchen, Micropanchax pumilis, Lamprichthys tanganicus,
Hypsopanchax zebra, Procatopus nototaenia, and Poecilia sphenops. (0.17:0.67)
26. Dorsoposterior region of lacrymal bordering orbit: (0) broad and dorsally
indented, forming process anterior to orbit (Fig. 4A); (1) broad, without indentation,
forming continuous bony shelf between orbit and lacrymal canal (Fig. 4B,C); (2)
narrow with lacrymal canal close to orbit (Fig. 4D). The dorsoposterior region of
the lacrymal is broad and continuous in all procatopodines examined except some
individuals of Procatopus nototaenia (coded as polymorphic, 0&1) The dorsoposterior
region of lacrymal is narrow in Pantanodon madagascariensis, Fluviphylax pygmaeus,
Tomeurus gracilis, Cnesterodon decemmaculatus, Phalloceros caudimaculatus, Phallotorynus victoriae,
and Cubanichthys pengelleyi. (0.30:0.61)
27. Dorsoposterior process of autopalatine: (0) short, forming only low bump, or
absent (Fig. 6B); (1) long, prominent, and dorsoposteriorly directed (Fig. 6C,D); (2)
long, prominent, and medially directed (Fig. 6A). Long and dorsoposteriorly directed
processes of the palatines are present in Micropanchax loati, M. pelagicus, ‘M.’ schioetzi,
Lamprichthys tanganicus, Pantanodon madagascariensis, Hypsopanchax zebra, Procatopus noto-
taenia, Gambusia affinis, Alfaro cultratus, Priapella compressa, Girardinus metallicus, and Poecilia
18 M. J. GHEDOTTI
Figure 6. Medial view of right suspensorium and opercular series in: A, Jenynsia unitaenia, USNM
300299; B, Aplocheilichthys spilauchen, USNM 304139; C, Micropnchax pelagicus, FMNH 62956; D, Poecilia
sphenops, KU 22711. Arrows indicate character states discussed in the text. Scale bars=1 mm.
sphenops. Long and medially directed processes of the palatines are present in all
anablepids examined, Cubanichthys pengelleyi, and Cyprinodon variegatus. (0.22:0.53)
28. Anterior margin of dermopalatine: (1) straight (Fig. 6A,B,D); (2) convex with
distinct anterior angle on dorsal-most third (Fig. 6C). This distinct convexity on the
dermopalatine is present in Micropanchax macropthalmus, M. johnstoni, M. loati, M.
pelagicus, Platypanchax cf. modestus, Pantanodon madagascariensis, Gambusia affinis, Tomeurus
gracilis, Cnesterodon decemmaculatus, Phalloceros caudimaculatus, Phallotorynus victoriae, Phal-
lichthys amates, and Fluviphylax obscurum. (0.13:0.50)
29. Posteroventral process of hyomandibula: (0) absent or indistinct (Fig. 6B–D); (1)
present and prominent, usually associated with rounding or abrupt change in shape
of anterior shelf of preoperculum (Fig. 6A). Ghedotti (1998) recognized a prominent
posteroventral process of the hyomandibula as synapomorphic of the Anablepidae.
The posteroventral process of the hyomandibula is prominent in all anablepids
examined, Cyprinodon variegatus, Valencia, and Fundulus chrysotus. (0.33:0.71)
30. Region on dorsal margin of interopercle anterior to area overlain medially by
posterior ceratohyal: (0) with small convex flange (Fig. 6B); (1) straight, without
distinct flange or concavity (Fig. 6A,C); (2) with small concavity (Fig. 6D). The
margin of the interopercle is straight in Micropanchax macropthalmus, M. pumilis, M.
loati, M. pelagicus, Hypsopanchax zebra, Procatopus nototaenia, Cubanichthys pengelleyi, and
all anablepids examined except Anableps dowi. The margin of the interopercle has a
distinct concavity in Girardinus metallicus, Poecilia sphenops, and Phallichthys amates. (0.20:
0.53)
31. Dorsal process of subopercle: (0) short, approximately as tall as the width at its
base (Fig. 6A,D); (1) long, taller than the width at its base (Fig. 6B,C). The dorsal
ventral anterior posterior

hypohyal ceratohyal ceratohyal
32-0 branchiostegal
ray
A
34-0
32-1 34-1 32-2
B C
19-0
Figure 7. Ventrolateral view of left hyoid arch in: A, Alfaro cultratus, KU 11122; B, Aplocheilichthys
spilauchen, USNM 304139; C, Micropanchax pelagicus, FMNH 26956. Cartilage is indicated by uniform
shading. Arrows indicate character states discussed in the text. Scale bars=1 mm.
process of the subopercle is long in all anablepids examined except Jenynsia unitaenia
as well as in Aplocheilichthys spilauchen, Micropanchax pumilis, M. johnstoni, M. loati,
Lamprichthys tanganicus, Hypsopanchax zebra, Cubanichthys pengelleyi, Cyprinodon variegatus,
Valencia, and Fundulus chrysotus. (0.17:0.67)
Branchial skeleton
32. Number of branchiostegal rays: (0) six (Fig. 7A); (1) five (Fig. 7B); (2) four (Fig.
7C). In individuals with six branchiostegal rays, the anterior two branchiostegal rays
are in contact with the slender anterior portion of the anterior ceratohyal, three
branchiostegal rays are in contact with the ventromedially expanded portion of the
anterior ceratohyal, and the posterior branchiostegal ray is in contact with the
posterior ceratohyal (Fig. 7A). In individuals with fewer than six branchiostegal rays,
one or both of the anterior two branchiostegal rays are absent (Fig. 7B,C). Jenynsia
multidentata, Jenynsia unitaenia, Aplocheilichthys spilauchen, Micropanchax johnstoni, ‘M.’
schioetzi, Platypanchax sp., Pantanodon madagascariensis, Hypsopanchax zebra, Procatopus
nototaenia, Fluviphylax obscurum, Valencia letourneuxi, Fundulus chrysotus, Crenichthys baileyi,
and all poeciliines examined except Alfaro cultratus, Priapella compressa, Gambusia affinis,
Poecilia sphenops, and Phallichthys amates have five branchiostegal rays. Micropanchax
macropthalmus, M. pumilis, M. loati, M. pelagicus, and Lamprichthys tanganicus have four
branchiostegal rays. (0.18:0.37)
33. Basihyal width: (0) slender, anterior ossified margin approximately twice as wide
as posterior part (Fig. 8A,B); (1) broad, anterior ossified margin greater than twice
as wide as posterior part (Fig. 8C). A wide, anteriorly ossified basihyal is capped by
an equally wide cartilage. The basihyal is wide in Anableps, Jenynsia, Platypanchax cf.
modestus, Hypsopanchax zebra, Procatopus nototaenia, Cubanichthys pengelleyi, and Cyprinodon
variegatus. (0.20:0.50)
20 M. J. GHEDOTTI
basihyal 33-0
ventral
hypohyal 33-1
34-0
34-1
urohyal anterior ceratohyal

A B C
Figure 8. Ventral view of anterior hyoid arch in: A, Aplocheilichthys spilauchen, USNM 304139; B, Alfaro
cultratus, KU 11122; C, Hypsopanchax zebra, USNM 191521. Cartilage is indicated by uniform shading.
Arrows indicate character states discussed in the text. Scale bars=1 mm.
34. Anterior process of anterior ceratohyal extending ventral to ventral hypohyal:

(0) present (Fig. 8A); (1) absent (Fig. 8B,C). Parenti (1981) and Costa (1998) recognized
the absence of an anterior process of the anterior ceratohyal (only one condyle on
the anterior ceratohyal) as synapomorphic of the Poeciliidae. The anterior process
of the anterior ceratohyal is absent in all poeciliids examined except Aplocheilichthys
spilauchen. (1)
35. Anterior margin of first hypobranchial: (0) mostly straight (Fig. 9A); (1) concave,
forming distinct anterolateral point (Fig. 9B). Pantanodon madagascariensis lacks hy-
pobranchials and was coded with a question mark (?) for this transformation series.
A concave anterior margin is present in Oxyzygonectes dovii, Cyprinodon variegatus,
Fundulus chrysotus, and all poeciliines examined except some individuals of Tomeurus
gracilis (coded as polymorphic, 0&1). (0.40:0.77)
36. Teeth on second and third hypobranchials: (0) absent (Fig. 9); (1) present.
Ghedotti (1998: fig. 10) illustrated the presence of these teeth in Anableps and
Oxyzygonectes. The hypobranchial teeth in Valencia hispanica are irregularly distributed,
with some individual hypobranchials asymmetrically lacking teeth in some individuals.
Pantanodon madagascariensis lacks hypobranchials and was coded as a question mark
(?) for this transformation series. Teeth are present on the second and third
hypobranchials in Oxyzygonectes dovii, Anableps dowi, and Valencia hispanica. (0.33:0.00)
37. Third hypobranchial: (0) cartilage continuous posteriorly, without distinct car-
tilaginous points of articulation with the second basibranchial and third cer-
atobranchial (Fig. 9B); (1) two distinct cartilaginous heads facing second basibranchial
medially and third ceratobranchial posterolaterally, no cartilage on posteromedial
surface (Fig. 9A). Pantanodon madagascariensis lacks hypobranchials and was coded as
a question mark (?) for this transformation series. The third hypobranchial has
distinct cartilaginous points of articulation with the second basibranchial and the
bb1 35-0 35-1

hb1
bb2 hb2
hb3 37-0
37-1
39-0
39-1 cb1
cb2
cb3
cb4
A B
cb5
Figure 9. Dorsal view of ventral branchial skeleton in: A, Hypsopanchax zebra, USNM 191521; B, Alfaro
cultratus, KU 11122. Cartilage is indicated by uniform shading. Gill rakers are not illustrated.
Abbreviations are as follow: bb=basibranchial, hb=hypobranchial, cb=ceratobranchial. Arrows
indicate character states discussed in the text. Scale bars=1 mm.
third ceratobranchial in Platypanchax cf. modestus, Hypsopanchax zebra, and Alfaro cultratus.
(0.33:0.00)
38. Teeth on fourth ceratobranchial: (0) present (Fig. 9A,B); (1) absent (Fig. 9C).
Most commonly cyprinodontoids have at least one row of conical teeth on the
anterior portion of the fourth ceratobranchial. These teeth are absent in Pantanodon
madagascariensis, Cyprinodon variegatus, Girardinus metallicus, Poecilia sphenops, Phallichthys
amates, Phallotorynus victoriae, Phalloceros caudimaculatus, and Cnesterodon decemmaculatus.
(0.25:0.57)
39. Anterior process of fifth ceratobranchial: (0) narrow, as narrow or more narrow
than anterior fourth ceratobranchial (Fig. 9B); (1) wide, wider than anterior fourth
ceratobranchial (Fig. 9A). The anterior process of the fifth ceratobranchial is wide
in Platypanchax cf. modestus, Hypsopanchax zebra, and Procatopus nototaenia. (0.33:0.33)
40. Interarcual cartilage: (0) present (Fig. 10); (1) absent. Parenti (1981) recognized
the absence of an interarcual cartilage as synapomorphic for all procatopodines
except Lamprichthys. Costa (1996) recognized the absence of an interarcual cartilage
as synapomorphic for Fluviphylax. The interarcual cartilage is absent in Micropanchax
macropthalmus, ‘M.’ schioetzi, Platypanchax cf. modestus, Lamprichthys tanganicus, Pantanodon
madagascariensis, Hypsopanchax zebra, Procatopus nototaenia, Fluviphylax obscurum, and Tom-
eurus gracilis. (0.33:0.75)
41. Third and fourth pharyngobranchial tooth plates: (0) separate (Fig. 10A); (1)
fused (Fig. 10B). In the derived state (1) the third pharyngobranchial has a long and
narrow anterodorsal process and the fourth pharyngobranchial tooth plate is notably
rounded posteriorly. In all cases where the third and fourth pharyngobranchial
tooth plates are fused, the opposing portions of the fifth ceratobranchials are
expanded medially and meet at the midline (Fig. 9C). However, they are not fused,
except in large individuals of Crenichthys baileyi. Because the fifth ceratobranchial
teeth oppose the posterior pharyngobranchial tooth plates and the two conditions
do not seem to occur independently, the expansion of the fifth ceratobranchials are
22 M. J. GHEDOTTI
iac
eb1 pbt2
pbt3
eb1
eb4
pbt4 41-1
41-0
eb3
A B C D
Figure 10. A, Dorsal view of dorsal branchial skeleton of Micropanchax pelagicus, FMNH 26956; B,
ventral view of pharyngobranchial toothplates of Micropanchax pelagicus, FMNH 26956; C, dorsal view
of dorsal branchial skeleton of Cnesterodon decemmaculatus, KU 22411; D, ventral view of pharyngobranchial
toothplates of Cnesterodon decemmaculatus, KU 22411. Cartilage is indicated by uniform shading. Ab-
breviations are as follow: eb=epibranchial; iac=interarcual cartilage; pbt=pharyngobranchial tooth-
plate. Arrows indicate character states discussed in the text. Scale bars=1 mm.
not recognized as a separate transformation series. In Crenichthys baileyi, the fifth

ceratobranchials and pharyngobranchial tooth plates bear many large, blunt teeth.
The apomorphic condition is present in Pantanodon madagascariensis, Crenichthys baileyi,
and all poeciliines examined except Tomeurus gracilis, Alfaro cultratus, Priapella compressa,
and Gambusia affinis. (0.25:0.67)
Vertebrae and ribs

42. Neural arch of first vertebra: (0) open dorsally (Fig. 11A); (1) closed dorsally at
tips of neuropophyses forming median, vertical flange (Fig. 11C); (2) closed dorsally
at base of neuropophyses forming horizontal ‘roof’ over spinal cord (Fig. 11B).
Parenti (1981) and Costa (1998) discussed the occurrence of an open neural arch
in cyprinodontoids. Costa (1998) also mentioned the presence of a horizontal
expansion between the bases of neuropophyses in some unspecified cyprinodontoids.
The first neural arch is closed with a median, vertical flange in Aplocheilichthys
spilauchen, Crenichthys baileyi, and all procatopodines examined except Pantanodon
madagascariensis. The first neural arch is closed by a horizontal flange at the bases of
the neuropophyses in all poeciliines examined except Cnesterodon decemmaculatus and
Tomeurus gracilis. (0.50:0.85)
43. Base of neural arch of first vertebra: (0) not in contact with exoccipitals
(exoccipital condyles present, see Fig. 3B,F, Fig. 11C); (1) in contact with exoccipitals
(exoccipital condyles absent, see Fig. 3G,H, Fig. 11A,B). Parenti (1981) recognized
the derived state in poeciliines, cyprinodontids, and Pantanodon. I confirm this
distribution. The base of the neural arch contacts the exoccipitals in Pantanodon
madagascariensis, the poeciliines, Cubanichthys pengelleyi, and Cyprinodon variegatus. (0.33:
0.86)
44. Ligastyle in adult males: (0) absent; (1) present and ossified; (2) present as an
unossified ligament. In most poeciliines, the hemal arch on the thirteenth vertebra
is resorbed during development forming a free, rod-like, usually ossified hemal spine
called the ligastyle dorsal to the gonopodial suspensorium (Turner, 1941; Kamel,
1947; Rosa-Molinar et al., 1994). An ossified ligastyle is present in all examined
42-0
supraoccipital
crest
neural
spine
1st neural
arch 2nd neural
arch
exoccipital
2nd vertebral
centrum
1st vertebral pleural rib

centrum
A
42-1
42-2
43-1 43-0
B C
Figure 11. Dorsolateral view of posterior neurocranium and first two vertebrae in: A, Alfaro cultratus,
KU 11122; B, Cnesterodon decemmaculatus, KU 22411; C, Hypsopanchax zebra, USNM 191521. Cartilage
is indicated by uniform shading. Arrows indicate character states discussed in the text. Scale bars=
1 mm.
poeciliines except Poecilia sphenops and Tomeurus gracilis in which the ligamentous
ligastyle is unossified. In Tomeurus gracilis there is a small ossification located just
anterior to the hemal spine on the seventeenth vertebra. The posterior location of
this ossified element suggests that it may not be the ligastyle and may be the free
hemal spine from the sixteenth vertebra. (0.67:0.90)
45. Hemal arches on fourteenth and fifteenth vertebrae in adult males: (0) absent
or angled posteriorly, similar to more posterior hemal arches; (1) enlarged and
angled anteriorly, involved in suspension of anal fin (gonapophyses present). Rosen
and Bailey (1963) discussed and illustrated the structure and distribution of gon-
apophyses in many poeciliines. Gonapophyses are present in all poeciliines examined
except Cnesterodon decemmaculatus and Tomeurus gracilis. The gonapophyses are least
elaborate and most similar to more posterior hemal arches in Alfaro cultratus. (0.50:
0.89)
46. Parapophyses on fifteenth vertebra in adult males: (0) on vertebral centrum as
in anterior vertebrae if present; (1) on hemal arch, below vertebral centrum.
24 M. J. GHEDOTTI
Cnesterodon decemmaculatus and Tomeurus gracilis lack a hemal arch on the fifteenth
vertebra in adult males (Rosen & Bailey, 1963: fig. 13). Alfaro cultratus has both
parapophyses and hemal arches on the fifteenth vertebral centrum (Rosen & Bailey,
fig. 17). All other poeciliines have parapophyses on the hemal arch of vertebra
fifteen in adult males. (0.50:0.88)
47. First hemal arch: (0) narrowly expanded, similar to more posterior hemal arches;
(1) broadly expanded and surrounding posterior portion of swimbladder, notably
wider than more posterior hemal arches. Parenti (1981) recognized expanded hemal
arches as a synapomorphy for aplocheilichthyines. The first hemal arch is expanded
in Aplocheilichthys spilauchen, the procatopodines, and Cubanichthys pengelleyi. (0.33:0.83)
48. Third hemal arch: (0) narrowly expanded, similar to more posterior hemal
arches; (1) broadly expanded and surrounding posterior portion of swimbladder,
notably wider than more posterior hemal arches. Sethi (1960) and Clausen (1967)
recognized three or more expanded hemal arches as evidence of a relationship
between Procatopus and Lamprichthys. The third hemal arch is expanded in Pantanodon
madagascariensis, Hypsopanchax zebra, Procatopus nototaenia, and Lamprichthys tanganicus.
(0.50:0.67)
49. Pleural ribs associated with hemal arches: (0) absent; (1) present. Some taxa
examined (Cnesterodon decemmaculatus, Micropanchax johnstoni, M. pumilis, and Platypanchax
cf. modestus) had only very reduced pleural ribs consisting of small, elongate ossifications
associated with the anterior hemal arch. These taxa were recognized as having the
derived character state (1). Parenti (1981) recognized pleural ribs on hemal arches
as a synapomorphy of Poeciliidae. Costa (1998) did not find the derived character
state in the poeciliid taxa he examined. Pleural ribs on hemal arches are found in
Aplocheilichthys spilauchen, Micropanchax loati, M. pelagicus, Procatopus nototaenia, Fluviphylax
obscurum, Cubanichthys pengelleyi, Cyprinodon variegatus, and all poeciliids examined except
Tomeurus gracilis and Poeciliopsis latidens. (0.14:0.60)
50. Distal tips of pleural ribs associated with sixth through tenth vertebrae in males:
(0) narrow and straight, extend ventrally (Rosen & Bailey, 1963: fig. 17); (1) narrow
and strongly angled anteriorly, extend under three or four more anterior vertebrae
(Rosen & Bailey, 1963: fig. 26); (2) thickened compared to other vertebrae and
somewhat curved anteriorly, extend under one or two more anterior vertebrae
(Rosen & Bailey, 1963: fig. 49). The sexually modified pleural ribs in Tomeurus gracilis
are narrow, horizontally positioned and far anterior (character state 1). Proximally
these pleural ribs are unossified and the distal tips do not have direct contact with
the parapophyses in adult males. In juvenile and transforming males the proximal
portion of these pleural ribs is continuous (though cartilaginous), connecting the
distal tips to the parapophyses. Rosen and Bailey (1963: 41) recognized the presence
of modified male pleural ribs in tribe Cnesterodontini (sensu Rosen & Bailey, 1963),
Neoheterandria, some Girardinus, and some Poeciliopsis. Rosen and Bailey (1963: 36) also
discussed the disconnected, horizontal ribs of Tomeurus. Narrow and strongly anteriorly
angled pleural ribs in males are present in Tomeurus gracilis, Cnesterodon decemmaculatus,
Phalloceros caudimaculatus, Phallotorynus victoriae, and Girardinus metallicus. Expanded and
somewhat anteriorly curved pleural ribs are present in adult male Poeciliopsis latidens
and Phallichthys amates. (0.40:0.57)
Figure 12. Lateral view of left pectoral girdles in: A, Jenynsia multidentata, KU 22416; B, Lamprichthys
tanganicus, AMNH 38421; C, Procatopus nototaenia, CAS 155515; D, Micropanchax pelagicus, FMNH 62956.
Pectoral-fin rays and distal radials are not illustrated. Supracleithra are illustrated in inset boxes.
Arrows indicate character states discussed in the text. Scale bars=1 mm.
Pectoral fin and girdle

51. Position of the pectoral fins: (0) low, dorsal insertion of pectoral fin below
midline; (1) high, dorsal insertion of pectoral fin at or above midline. Sethi (1960)
and Clausen (1967) recognized high pectoral fins as evidence of relationship between
Lamprichthys and Procatopodines (sensu Myers, 1955). Parenti (1981) and Costa (1998)
recognized high pectoral fins as a synapomorphic reversal in poeciliids to the
condition in non-cyprinodontiform atherinomorphs. The pectoral fins are high in
all poeciliids examined. (1)
52. Ventral arm of posttemporal: (0) ossified (Fig. 12A,C); (1) unossified (Fig. 12B,
D). The ventral arm of the posttemporal is unossified in Cnesterodon decemmaculatus,
Phalloceros caudimaculatus, Phallotorynus victoriae, Valencia, Fundulus chrysotus, and all
procatopodines examined except Hypsopanchax zebra, Procatopus nototaenia, Pantanodon
madagascariensis, and ‘Micropanchax’ schioetzi. (0.13:0.50)
53. Bony flange on supracleithrum in contact with posttemporal: (0) present (Fig.
12A); (1) absent (Fig. 12B–D). The small bony flange on the supracleithrum
may be homologous with the supracleithral sensory canal that is found in other
acanthomorph fishes. Although Stiassny (1993) considered the loss of this canal to
be synapomorphic of a mugilid and atherinomorph clade, this flange is positionally
similar to the supracleithral sensory canal in the angle of the flange and its contact
26 M. J. GHEDOTTI
with the posttemporal. This suggests that the canal was either independently
acquired in cyprinodontoids or that the canal is reduced rather than lost in some
atherinomorphs. A supracleithral flange is absent in all poeciliids examined, Valencia,
and Crenichthys baileyi. Tomeurus gracilis lacks a supracleithrum and was coded with a
question mark (?). (0.33:0.71)
54. Supracleithrum shape: (0) elongate, length approximately twice width (Fig. 12A,
B); (1) short, often disc-like, length distinctly less than twice width (Fig. 12C,D).
Stiassny (1993) contrasted the reduced supracleithra of mugilids and atherinomorphs
with the elongate supracleithra of percomorphs. However, my use of the descriptor
elongate for the plesiomorphic condition in this transformation series is not syn-
onymous with Stiassny’s (1993) usage. All cyprinodontiforms examined have a
reduced supracleithrum as discussed in Stiassny (1993) and this transformation series
refers to the shape of these reduced supracleithra. A short, disc-like supracleithrum
is present in all poeciliids examined except Micropanchax loati, Lamprichthys tanganicus,
Platypanchax cf. modestus, and Pantanodon madagascariensis. Tomeurus gracilis lacks a
supracleithrum and was coded with a question mark (?). (0.17:0.69)
55. Posterior extension of dorsal enclosure of cleithrum: (0) present with distinct
ventral notch (Fig. 12A); (1) present and straight or slightly curved ventrally (Fig.
12B–D); (2) absent. The posterior extension of the dorsal enclosure of the cleithrum
has a distinct notch in Oxyzygonectes dovii, Jenynsia unitaenia, J, multidentata, Cubanichthys
pengelleyi, Crenichthys baileyi, and Profundulus labialis. A posterior extension of the dorsal
enclosure of the cleithrum is absent in Anableps dowi. (0.40:0.50)
56. Dorsal proximal pectoral radial: (0) free, unfused to scapula (four free radials)
(Fig. 12A,B,D); (1) fused with scapula (three free radials) (Fig. 12C). Rosen (1965)
mentioned that Pantanodon and Procatopus both have only three free radials. The
dorsal proximal pectoral radial is fused to the scapula in Pantanodon madagascariensis,
Procatopus nototaenia, Cnesterodon decemmaculatus, and Tomeurus gracilis. This radial is
sometimes fused with the scapula in Gambusia affinis, Poeciliopsis latidens, and Cyprinodon
variegatus and was coded as polymorphic (0&1) in these taxa. (0.60:0.33)
57. Posteroventral coracoid: (0) straight with prominent, posterior point (Fig. 12A);
(1) rounded, with posterior point obscured (Fig. 12B–D). The apomorphic character
state is present in all poeciliids except some individuals of Fluviphylax obscurum. (0.67:
0.89)
58. Dorsal (primary) postcleithrum: (0) present; (1) absent. Parenti (1981), Costa
(1998), and Ghedotti (1998) indicated that Anableps and all poeciliids except Aplo-
cheilichthys spilauchen lack a primary (dorsal) postcleithrum. (0.50:0.90)
59. Ventral postcleithrum width: (0) slender, similar in width to adjacent first pleural
rib (Ghedotti, 1998: fig. 13A); (1) broad, with a laminar flange that is wider than
adjacent first pleural rib (Ghedotti, 1998: fig. 13B). All anablepids examined,
Micropanchax macropthalmus, Platypanchax cf. modestus, Lamprichthys tanganicus, Valencia,
and Crenichthys baileyi have broad ventral postcleithra. (0.20:0.56)
Pelvic fin and girdle

60. Number of pelvic-fin rays: (0) six; (1) fewer than six; (2) greater than six. Most
atherinomorph fishes have six pelvic-fin rays (Parenti, 1981) and the number of
Figure 13. Ventral view of pelvic girdle in: A, Oxyzygonectes dovii, KU 18174; B, Cnesterodon decemmaculatus,
male, KU 22411; C, Procatopus nototaenia, CAS 155515; D, Tomeurus gracilis, male, UMMZ 167944; E,
Tomeurus gracilis, female, UMMZ 167944. Pelvic-fin rays are not illustrated. Cartilage at anterior tips
of pelvic girdles is indicated by uniform shading. Arrows indicate character states discussed in the text.
Scale bars=1 mm.
pelvic fin rays is nearly invariant within species with six pelvic-fin rays. Phallotorynus
victoriae, Phalloceros caudimaculatus, and Cnesterodon decemmaculatus all have five pelvic-fin
rays (occasionally four in C. decemmaculatus). Tomeurus gracilis has from two to four
pelvic-fin rays, and Cyprinodon variegatus has seven pelvic-fin rays. (1)
61. Distinct ventral flange on central part of pelvic bone in males: (0) absent (Fig.
13A,C,D,E); (1) present (Fig. 13B). The ventral flange is present in all poeciliines
examined except Tomeurus gracilis. (0.50:0.90)
62. Medial processes of pelvic bones in males: (0) overlap medially (Fig. 13A,B,C);
(1) fused medially, forming single pelvic element (Fig. 13C); (2) separated (Ghedotti,
1998: fig. 14B). The medial processes of the left and right pelvic bones are fused
together in male Pantanodon madagascariensis, Tomeurus gracilis, Poeciliopsis latidens, and
Phallichthys amates. The medial processes of the pelvic girdle in Anableps dowi are
separated. (0.40:0.00)
63. Posterior process of pelvic bone: (0) present (Fig. 13A); (1) absent (Fig. 13B,D,
E); (2) elongate, approximately twice as long as medial process (Fig. 13C). Posterior
processes are absent in poeciliines and Anableps dowi and are elongate in Hypsopanchax
zebra, Procatopus nototaenia, and Cyprinodon variegatus. (0.50:0.86)
64. Position of pelvic girdle in adults: (0) posterior, lateral arm of pelvic girdle at
or posterior to pleural rib associated with seventh vertebra; (1) anterior, lateral arm
of pelvic girdle anterior to pleural rib associated with seventh vertebra. Parenti
28 M. J. GHEDOTTI
(1981) recognized an anteriorly positioned pelvic girdle as synapomorphic for

poeciliids. Costa (1998) confirmed this and also recognized the presence of an
anterior pelvic girdle in cyprinodontids. An anteriorly positioned pelvic girdle is
present in all poeciliids, cyprinodontids, and some individuals of Jenynsia multidentata.
(0.67:0.83)
65. Position of pelvic girdle in adult males: (0) same as in females, lateral arm of
the pelvic girdle approximately at or posterior to pleural ribs associated with fifth
vertebra; (1) slightly more anterior than in females, lateral arm of the pelvic girdle
at pleural ribs associated with third or fourth vertebrae; (2) far anterior, under
pectoral girdle, lateral arm of the pelvic girdle at or more anterior than pleural rib
associated with second vertebra. Differences in position between the male and female
pelvic girdle only occur in a subset of the taxa with anterior pelvic girdles (T.S. 64,
character state 1). A slightly more anterior pelvic girdle is found in male Girardinus
metallicus, Poecilia sphenops, Phallichthys amates, and Poeciliopsis latidens. The pelvic girdle
is under the pectoral girdle in male Tomeurus gracilis, Cnesterodon decemmaculatus,
Phalloceros caudimaculatus, and Phallotorynus victoriae. (1)
66. Pelvic bones: (0) not in contact with pleural ribs; (1) overlapping pleural ribs
ventrolaterally. The pelvic bones overlap the pleural ribs in Anableps dowi, Aplo-
cheilichthys spilauchen, Valencia, and all procatopodines examined except Pantanodon
madagascariensis. (0.20:0.71)
Anal fin and skeletal supports

Anal-fin radial terminology follows Patterson (1992). Radials are numbered based
on the anal-fin ray with which they are associated. For example, the second proximal
radial is the radial associated in series with the second anal-fin ray and not the
second proximal radial when counting proximal radials from anterior to posterior.
Directional terminology in reference to the gonopodium is for a gonopodium
pointing ventrally.
67. Male anal fin: (0) generally unmodified from female condition; (1) forming rod-
like intromittent organ supported by anal-fin rays three, four, and five (Fig. 14C,
D); (2) forming tubular intromittent organ supported primarily by four or more
anal-fin rays (Fig. 14A,B). The modification of the male anal-fin rays in poeciliines
has long been used to diagnose this group (e.g. Garman, 1895; Eigenmann, 1907;
Regan, 1911, 1913). Regan (1911), Jordan (1923), Hubbs (1924), Parenti (1981),
and Ghedotti (1998) suggested a close relationship between Jenynsia and Anableps
based in whole or in part on their shared possession of tubular intromittent organs.
(1)
68. Male anal-fin ray length: (0) short, all rays less than or equal to one third
standard length (short gonopodium); (1) long, some rays greater than one third
standard length (long gonopodium). Chambers (1987) discussed the distribution of
short and long gonopodia in poeciliine fishes. Chambers (1987) and Dawes (1991)
suggested that shorter gonopodia are correlated with more complex reproductive
behavior involving more elaborate male courtship displays. Long gonopodia are
present in all poeciliines examined except Alfaro cultratus, Priapella compressa, Gambusia
affinis, and Poecilia sphenops. (0.50:0.83)
69. Number of anal-fin rays: (0) 14 and greater; (1) 9–14; (2) eight and fewer. Sethi
78-0
11l
10l
9l
1l
1r
8l
A 2l
2r
3l
4l
6l 7l
70-0 84-0
80-1 10
9
78-1 8
1
2
84-1 5 4 7
B 75-1 3 6 77-1
10 9 8 7
6
5p 5a 75-0
79-0 4p
1 72-0
C 2
3 4a
70-1
79-1
77-2
80-0 8 7 6
5p 5a
4p
72-1
1 2
3 4a
D
Figure 14. Left lateral view of the anal fin and skeletal supports of adult males in: A, Anableps dowi,
sinistral specimen, KU 18689; B, Jenynsia multidentata, sinistral specimen, KU 21307; C, Gambusia affinis,
KU 22033; D, Cnesterodon decemmaculatus, KU 22411. The numbers refer to anal-fin ray number. Letters
refer to the left (l) or right (r) halves of the indicated anal-fin ray in Anableps dowi or anterior (a) or
posterior (p) branches of the indicated ray in Gambusia affinis and Cnesterodon decemmaculatus. Arrows
indicate character states discussed in the text. Scale bars=1 mm.
(1960) suggested that an increased number of anal-fin rays in Procatopus, Hypsopanchax,

and Lamprichthys provided evidence of a close relationship among these genera.
Micropanchax pelagicus, ‘M.’ schioetzi, Platypanchax cf. modestus, Lamprichthys tanganicus,
Pantanodon madagascariensis, Hypsopanchax zebra, Procatopus nototaenia, and Profundulus
labialis have 14 or more anal-fin rays. Fluviphylax obscurum has eight anal-fin rays.
Valencia hispanica and Crenichthys baileyi have 12–14 anal-fin rays and were coded as
polymorphic (0&1). (0.57:0.63)
30 M. J. GHEDOTTI
70. Length of anal-fin ray two in adult males: (0) long, longer than one quarter
length of anal-fin ray three (Fig. 14A); (1) short, shorter than one quarter length of
anal-fin ray three (Fig. 14B–D). This anal-fin ray is short in Jenynsia and the
poeciliines. (0.50:0.92)
71. Anal-fin ray three in adult females: (0) branched; (1) unbranched (Ghedotti,
1998: fig. 15). Miller (1979) used the lack of branching in the third anal-fin ray as
evidence of relationship between Anableps and the Poeciliinae (=Poeciliidae sensu
stricto). Ghedotti (1998) recognized an unbranched third anal-fin ray as evolving
independently in the ancestor of the Anablepinae (reversed in Jenynsia multidentata)
and the ancestor of the Poeciliinae. The third anal-fin ray is unbranched in Anableps
dowi, Jenynsia unitaenia, and all poeciliines examined. Some individuals of Pantanodon
madagascariensis have an unbranched third anal-fin ray; therefore, this species is coded
as polymorphic (0&1). (0.50:0.85)
72. Distal tip of anal-fin ray three in males: (0) simple, lacking bony or membranous
cirri (Fig. 14C); (1) with laterally paired bony cirri at distal tip (Fig. 14D, 15); (2) with
laterally paired membranous cirri subdistally. Rosen and Bailey (1963) mentioned the
presence of laterally paired cirri in most cnesterodontines and Girardinus and described
the complex structure of the gonopodium in Tomeurus gracilis. The paired ossified
cirri in Cnesterodon decemmaculatus are elongate, directed posteriorly, and closely
apposed to each other distally (Fig. 14D, 15A). The paired ossified cirri in Phalloceros
caudimaculatus are shorter than in C. decemmaculatus, directed posteriorly, and the
lateral halves are separated distally forming a forked process (Rosen & Bailey, 1963:
fig. 31C,D). The paired ossified cirri in Phallotorynus victoriae form a process composed
of a posteriorly directed, forked process similar to that seen in Phalloceros caudimaculatus.
However, in Phallotorynus victoriae the forked process is attached to a more distal
component composed of a large, paired broad structure (Rosen & Bailey, 1963: fig.
31A,B). The paired ossified cirri in Tomeurus gracilis form a broad forked process that
is strongly curved dorsally (Fig. 15B). The paired membranous cirri in Girardinus
metallicus form a forked unossified process (Rosen & Bailey, 1963: fig. 47A). (1)
73. Fleshy palp on ventral surface of anal-fin ray three in males: (0) absent; (1)
present (Rosen & Bailey, 1963: fig. 25). Rosen and Bailey (1963) used the presence
of this palp in Alfaro and Poecilia as evidence that Alfaro should be placed in the tribe
Poeciliini. Clark and Aronson (1951) and Chambers (1987) described the anatomy
of the palp and note that it is a highly vascularized and well innervated structure.
Rodriguez (1997) suggested that the presence of a palp was independently derived
in Alfaro and an ancestor of a clade including Poecilia, Pamphorichthys, and Limia. A
palp is present in Alfaro cultratus and Poecilia sphenops. (0.50:0.00)
74. Serrae on posterior margin of posterior branch of anal-fin ray four in adult
males: (0) absent (Fig. 15B); (1) present (Fig. 15A). Howell and Hubbs (1936: 9) cited
absence of serrae on this anal-fin ray in Alfaro as evidence that the genus is primitive
and originated ‘‘earlier in the phylogeny of the viviparous cyprinodonts’’. Serrae
are present on the posterior branch of anal-fin ray four in adult males in all
poeciliines examined except Alfaro cultratus and Tomeurus gracilis. (0.50:0.89)
75. Length of anal-fin ray five in adult males: (0) long, approximately as long as
anal-fin ray three (Fig. 14A,C,D); (1) short, less than three quarters length of anal-
fin ray three (Fig. 14B). Anal-fin ray five is shortened to some degree in both Jenynsia
examined, less so in J. unitaenia than in J. multidentata. (1)
Figure 15. Left lateral view of gonopodial tip in: A, Cnesterodon decemmaculatus, KU 22411; B, Tomeurus
gracilis, UMMZ 167944. The numbers refer to anal-fin ray number and the letters refer to the anterior
(a) or posterior (p) branches of the indicated ray. Arrows indicate character states discussed in the text.
Scale bars=1 mm.
76. Posteriorly directed claw on distal tip of anterior branch of anal-fin ray five in
adult males: (0) absent (Fig. 15B); (1) present (Fig. 15A). A posteriorly directed claw
on the distal tip of anal-fin ray five in adult males is present in Gambusia affinis and
Cnesterodon decemmaculatus. Rosen and Bailey (1963) mentioned the presence of this
claw in Gambusia, Belonesox, Cnesterodon, some Poecilia species, and some Xiphophorus
species. (0.50:0.00)
77. Anal-fin ray six in adult males: (0) branched and slender distally; (1) unbranched
and slender distally (Fig. 14A,B); (2) branched and thickened distally with branches
variably fused (Fig. 14C,D). Anal-fin ray six is unbranched in adult male Jenynsia
and Anableps. It is thickened in all poeciliines examined except Alfaro cultratus. (1)
78. Seventh middle anal-fin radial in males and females: (0) similar in size to
adjacent middle radials (Fig. 14A,C,D); (1) enlarged with respect to adjacent middle
anal-fin radials (Fig. 14B). Parenti (1981) referred to this bone as the sixth middle
radial because it is the sixth most anterior middle radial. I refer to it here as the
seventh middle anal radial because it is associated in series with the seventh anal-
fin ray. Parenti (1981) and Ghedotti (1998) used the apomorphic character state (1)
to diagnose the genus Jenynsia. The seventh middle anal radial is enlarged in both
Jenynsia species examined. (1)
79. Position of proximal anal-fin radials in adult males: (0) anteriorly inclined or
vertical (Fig. 14A–C; 16B–D); (1) posteriorly inclined (Fig. 14D; 16A). Proximal
anal-fin radials in adult males are posteriorly inclined in Alfaro cultratus, Tomeurus
gracilis, and Cnesterodon decemmaculatus. (0.50)
80. Anterior proximal anal-fin radial in adult males: (0) separate from subsequent
32 M. J. GHEDOTTI
Figure 16. Left lateral view of proximal and middle anal radials two through six in adult males in: (a)
Alfaro cultratus, KU 11122; (b) Poeciliopsis latidens, KU 17008; (c) Poecilia sphenops, KU 22711; (d) Phalloceros
caudimaculatus, UMMZ 206083. The numbers refer to radial numbers. Abbreviations are as follow:
vf= ventral flange on proximal radial five, df= dorsal flange on proximal radial five. Arrows indicate
character states discussed in the text. Scale bars=1 mm.
anal-fin radial (Fig. 14C,D; 16); (1) fused to subsequent proximal anal-fin radial
forming a structure composed of five or six fused proximal anal-fin radials (Fig.
14A,B). The apomorphic character state is present in Anableps and Jenynsia. (1)
81. Dorsal portion of fourth and fifth proximal anal-fin radials in adult males: (0)
separate (Fig. 16A,D); (1) dorsal tips touching or fused forming oblong opening (Fig.
16C); (2) completely fused (Fig. 16B). The fourth and fifth proximal anal-fin radials
are completely fused in Gambusia affinis, Poeciliopsis latidens, and Heterandria formosa and
partially fused, forming an oblong opening in Phallichthys amates and Poecilia sphenops.
(0.67)
82. Lateral processes on dorsal portion of fifth proximal anal-fin radial in adult
males: (0) absent (Fig. 16A); (1) present and do not extend dorsal to central arm of
fifth proximal anal-fin radial (Fig. 16B,D); (2) present and lateral tips of processes
extend dorsal to central arm of fifth proximal anal-fin radial (Fig. 16C). Processes
are present in all poeciliines examined except Alfaro cultratus and are dorsally directed
in Gambusia affinis and Poecilia sphenops. These processes are most well developed in
Poeciliopsis latidens. (0.67:0.89)
83. Lateral flanges on ventral portion of fifth proximal anal-fin radial in adult males:
(0) absent (Fig. 14A,B); (1) present and continuous, without dorsal cleft (Fig. 14C,
D; 16A–C); (2) present and cleft dorsally, forming separate dorsally directed processes
(Fig. 16D). Rosen and Bailey (1963) discussed this transformation series and noted
that gonactinosts (proximal anal-fin radials three through five in males) in Gambusia
and Belonesox are columnar, lacking lateral flanges. All other poeciliines have plate-
like gonactinosts with lateral flanges. Lateral flanges on the fourth proximal anal-
fin radial are present in adult males of all poeciliines examined except Gambusia
affinis and are cleft dorsally in Phallotorynus victoriae, Phalloceros caudimaculatus, and
Girardinus metallicus. (0.50:0.78)
84. Ventral portion of seventh, eighth, and ninth proximal anal-fin radials in adult
males: (0) laterally compressed, with anterior and posterior flanges (Fig. 14A,B); (1)
not laterally compressed, without anterior and posterior flanges (Fig. 14C,D). The
seventh, eighth, and ninth proximal anal-fin radials in adult males lack anterior and
posterior flanges in all poeciliines examined. (1)
85. Lateral processes on ventral portion of seventh, eighth, and ninth proximal
anal-fin radials in adult males: (0) absent (Fig. 14A,B); (1) present, not contacting
successive posterior processes and oriented parallel to each other (Fig. 14C); (2)
present, posterior part of each process contacting lateral process of posteriorly
successive proximal anal-fin radial (Fig. 14D). These processes are most developed
in Tomeurus gracilis. The fusion of these processes in Tomeurus gracilis forms a coossified
structure that Rosen and Bailey (1963: 36) called the secondary gonactinostal
complex. The lateral processes are present in all poeciliines examined and contact
each other in Cnesterodon decemmaculatus and Tomeurus gracilis. (1)
Dorsal and caudal fins and skeletal supports

86. Number of dorsal-fin rays: (0) 10–13; (1) fewer than nine; (2) more than 14.
Jenynsia unitaenia, Micropanchax pumilis, M. pelagicus, Platypanchax cf. modestus, Procatopus
nototaenia, Poecilia sphenops, Phallichthys amates, Valencia hispanica, and Profundulus labialis
have 10–13 dorsal-fin rays. Lamprichthys tanganicus and Hypsopanchax zebra have 14 or
more dorsal-fin rays. Anableps dowi and Valencia letourneuxi have eight through ten
dorsal-fin rays and are coded as polymorphic (0&1). (0.30:0.10)
87. Position of dorsal-fin origin: (0) at vertical that passes through or slightly anterior
to origin of anal fin; (1) at vertical that passes approximately through centre of anal-
fin base; (2) at vertical that passes posterior to anal-fin base. The dorsal-fin origin
is approximately above the middle of the anal-fin base in Oxyzygonectes dovii, Phalloceros
caudimaculatus, Phallotorynus victoriae, Gambusia affinis, Girardinus metallicus, Phallichthys
amates, Poeciliopsis latidens, and all procatopodines examined. The dorsal fin is posterior
to the anal-fin base in Anableps dowi, Alfaro cultratus, Tomeurus gracilis, and Fluviphylax
obscurum. (0.25:0.60)
88. Hypurals in adults: (0) fused, forming single continuous hypural plate (Fig. 17B);
(1) fused, forming separate symmetrical dorsal and ventral hypural elements (Fig.
17A). Costa (1998) discussed this character and states that a single hypural plate is
present in all cyprinodontoids except some anablepids. Ghedotti (1998) identified
two separate hypural plates in many anablepids, Aplocheilichthys spilauchen, and Alfaro
cultratus. Separate dorsal and ventral elements are present in Aplocheilichthys spilauchen,
some individuals of Gambusia affinis (coded 0&1), Alfaro cultratus, Poecilia sphenops,
Phallichthys amates, and all anablepids examined except some individuals of Jenynsia
multidentata (coded 0&1). (0.50)
89. Size of caudal accessory cartilage between distal neural spines of preural
vertebrae three and four: (0) slender, approximately as wide as adjacent distal neural
spines (Fig. 17A); (1) broad, wider than adjacent distal neural spines (Fig. 17B).
Costa (1996) recognized enlarged accessory caudal cartilages as a synapomorphy of
Fluviphylax, Aplocheilichthys spilauchen, and the procatopodines (=Aplocheilichthyinae
sensu Costa, 1996). This character state is not found in all procatopodines or
Aplocheilichthys spilauchen, is often polymorphic, and is also found in some poeciliines.
34 M. J. GHEDOTTI
Figure 17. Left lateral view of adult caudal skeleton in: A, Aplocheilichthys spilauchen, USNM 304139; B,
‘Micropanchax’ schioetzi, USNM 323612. Caudal-fin rays are not illustrated. Cartilage is indicated by
uniform shading. Arrows indicate character states discussed in the text. Scale bars=1 mm.
A broad caudal accessory cartilage is found in Micropanchax pumilis, ‘M.’ schioetzi,

Platypanchax cf. modestus, Fluviphylax obscurum, Phallotorynus victoriae, Girardinus metallicus,
Poecilia sphenops, Phallichthys amates, and some individuals of Micropanchax johnstoni,
Procatopus nototaenia, Cnesterodon decemmaculatus, Phalloceros caudimaculatus, and Valencia
letourneuxi (polymorphic species coded 0&1). (0.54:0.29)
Lateralis system
90. Supraorbital canal in adults: (0) closed, forming pores 2b, 3, and 4a (Clausen,
1967: fig. 19); (1) open, exposing two neuromasts (Clausen, 1967: fig. 20). Clausen
(1967) discussed the closed supraorbital systems of Lamprichthys and Plataplocheilus as
evidence of relationship between these two genera. Clausen (1967) also described a
new genus, Poropanchax, to contain a species with a closed supraorbital canal previously
placed in Aplocheilichthys. Parenti (1981) recognized an open supraorbital canal as
synapomorphic for Poeciliidae. The supraorbital canal is open in Aplocheilichthys
spilauchen,Valencia, all procatopodines examined except Micropanchax macropthalmus and
Lamprichthys tanganicus, and all poeciliines examined except Poecilia sphenops. The
supraorbital canal is variably open in Girardinus metallicus and Phallichthys amates.
These two species were coded as polymorphic (0&1). (0.43:0.64)
91. Lacrymal canal in adults: (0) closed, forming four pores; (1) open, exposing
three neuromasts. The lacrymal canal in adults is open in Gambusia affinis, Tomeurus
gracilis, Cnesterodon decemmaculatus, Phalloceros caudimaculatus, Phallotorynus victoriae, Valencia,
and all procatopodines examined except Lamprichthys tanganicus, Procatopus nototaenia,
and Hypsopanchax zebra. The lacrymal canal is variably open in adult ‘Micropanchax’
schioetzi, Phallichthys amates, and Poeciliopsis latidens, and these taxa were coded as
polymorphic (0&1). (0.44:0.67)
Colour pattern
All colour pattern states were ascertained from descriptions of colour in life in
the scientific and aquarium hobbyist literature (Scheel, 1970; Weitzman, 1982;
Fromm, 1982; Foster, 1983; White, 1983; Wildekamp, 1995) or from colour
photographs (Axelrod et al., 1993; Wischnath, 1993).
92. Dorsal iris colour in life: (0) not distinctly different than ventral iris; (1) blue or
silvery; (2) red. The iris of most poeciliids is reflective, but the differentially, more
highly reflective dorsal portion of the iris is restricted in distribution and may be
silvery, blue, or red. Silver and blue dorsal irises are not separated into distinct
character states because the blue colour is often light and difficult to distinguish
from silver. The dorsal iris in Oxyzygonectes dovii is a reflective silver (Fromm, 1982).
The dorsal irises of Micropanchax macropthalmus, M. pumilis, M. johnstoni, M. loati, ‘M’
schioetzi, and Lamprichthys tanganicus are a reflective blue or silvery blue colour (Axelrod
et al., 1993; Wischnath, 1993; Wildekamp, 1995). The dorsal irises of Aplocheilichthys
spilauchen and Procatopus nototaenia are a reflective red colour (Axelrod et al., 1993;
Wischnath, 1993; Wildekamp, 1995). Colour data or photographs showing colour
in life are not available for Pantanodon madagascariensis, Platypanchax cf. modestus,
Hypsopanchax zebra, and M. pelagicus and these taxa were coded as unknown (?). (0.50:
0.71)
93. Reflective blue dorsolateral coloration in males: (0) absent; (1) present. The
dorsolateral surface of all procatopodines examined for which data are available
(Micropanchax johnstoni, M. macropthalmus, M. pumilis, M. loati, ‘M.’ schioetzi, Lamprichthys
tanganicus, and Procatopus nototaenia), Alfaro cultratus, Priapella compressa, Girardinus metallicus,
Poecilia sphenops, and Cyprinodon variegatus are a reflective blue colour (Axelrod et al.,
1993; Wischnath, 1993; Wildekamp, 1995). (0.2:0.71)
External morphology
94. Genital laterality in adult males: (0) absent; (1) present, with both dextral and
sinistral males within each species; (2) present, with all males within a species either
dextral or sinistral. Sexual laterality in males was first described in Anableps and
Jenynsia by Garman (1895). Parenti (1981) first identified laterality in the genital
papilla of the oviparous Oxyzygonectes and recognized laterality as a synapomorphy
of the Anablepidae. Rosen and Bailey (1963) noted the presence of gonopodial
asymmetry in Poeciliopsis, Phallichthys, and Xenodexia. Ghedotti and Weitzman (1996)
discussed the presence of genital laterality in all male anablepids and female Anableps
as well as its absence in female Jenynsia. All anablepids examined exhibit sexual
laterality in the male gonopodia (anablepines) or male genital papilla (Oxyzygonectes
dovii) with both dextral and sinistral individuals in the same species. Male Phallichthys
amates and Poeciliopsis latidens have sinistral gonopodia. (0.67:0.75)
95. Adult sexual size dimorphism: (0) females larger than males; (1) males larger
than females; (2) minimal, with males and females similar in size. Parenti (1981)
recognized Aplocheilichthys spilauchen and the procatopodines as exhibiting sexual
dimorphism, with males larger than females. Males are larger than females in
Aplocheilichthys spilauchen and all procatopodines examined. Adult males and females
are approximately the same size in Cyprinodon variegatus. (0.67:0.92)
96. Branchiostegal membranes: (0) narrowly joined, left and right branchiostegal
membranes joined anterior to subopercle; (1) broadly joined, left and right branch-
iostegal membranes joined ventral to subopercle. Myers (1924b) diagnosed the genus
Micropanchax having broadly joined branchiostegal membranes. However, Clausen
36 M. J. GHEDOTTI
Figure 18. Semidiagrammatic representation of body shape in males of six poecilioid species.
(1967) disagreed and recognized this character as variable within this group. The
branchiostegal membranes are broadly joined in Micropanchax macropthalmus, M. loati,
M. pelagicus, Pantanodon madagascariensis, Fluviphylax obscurum, Cyprinodon variegatus, and
all poeciliines examined except Gambusia affinis, Alfaro cultratus, and Tomeurus gracilis.
(0.13:0.42)
97. Preorbital squamation on lacrymal: (0) absent; (1) present. Clausen (1967)
mentioned the presence of a scaled preorbital region in Lamprichthys, Plataplocheilus,
Aplocheilichthys spilauchen, and some Micropanchax. The preorbital scales are particularly
dense in Aplocheilichthys spilauchen, Lamprichthys tanganicus, and Anableps dowi. Preorbital
scales are present as either isolated scales or dense rows in all anablepids examined,
Aplocheilichthys spilauchen, Micropanchax pelagicus, ‘M’. schioetzi, Platypanchax cf. modestus,
Lamprichthys tanganicus, Hypsopanchax zebra, Procatopus nototaenia, Alfaro cultratus, and
Cyprinodon variegatus. (0.14:0.54)
98. Body shape (Fig. 18): (0) terete; (1) deep, laterally compressed. Whitehead (1962)
suggested that the deep body profiles of Pantanodon and Procatopus might indicate a
close relationship between these genera. Parenti (1981) recognized a deep body with
ventrally expanded ribs as synapomorphic of a Procatopus and Hypsopanchax clade.
‘Micropanchax’ schioetzi, Platypanchax cf. modestus, Lamprichthys tanganicus, Pantanodon
madagascariensis, Hypsopanchax zebra, Procatopus nototaenia, Phallichthys amates, Cubanichthys
pengelleyi, and Cyprinodon variegatus are laterally compressed. (0.33:0.75).
Reproductive mode
99. Viviparity: (0) absent; (1) present. Viviparity is defined as ‘‘a process in which
eggs are fertilized internally and are retained within the maternal reproductive
system for a significant period of time, during which time they develop to an
advanced state and then are released’’ (Wourms, 1981: 480). This definition avoids
the ambiguities of the more traditional use of viviparity that require the determination
that there is ‘significant’ maternal nutrient contribution to developing embryos.

Tomeurus gracilis is coded as polymorphic (0&1) since Tomeurus gracilis exhibits either
viviparity, giving birth to well formed young, or oviparity, releasing fertilized eggs
relatively early in their development (Breder & Rosen, 1966). (0.67:0.92)
Phylogenetic analysis
The same three equally most parsimonious trees were found for the ingroup in
each of the three analyses based on different constrained outgroup topologies
(Parenti, 1981, outgroup topology: length=434, CI=0.39, RI=0.70; Meyer &
Lydeard, 1993, outgroup topology: length=416, CI=0.40, RI=0.70; Costa (1998)
outgroup topology: length=430, CI=0.39, RI=0.70). The phylogenetic-tree stat-
istics vary among analyses because of differences in transformation series optimization
on each of the constrained outgroup topologies. The three most parsimonious trees
have two areas of incongruence, among Heterandria formosa, Poeciliopsis latidens, and a
clade of seven other poeciliines as well as among Phallotorynus victoriae, Phalloceros
caudimaculatus, and a clade containing Cnesterodon decemmaculatus and Tomeurus gracilis.
The proposed classification in Table 4 is congruent with the relationships illustrated
in Figure 19. One of three most parsimonious trees with character state trans-
formations is illustrated in Figure 20.
Bremer decay indices (Fig. 19) show that most relationships within the Pro-
catopodinae are not as well supported as the higher taxonomic groups (e.g.,
Poeciliinae). However, a clade composed of Hypsopanchax zebra and Procatopus nototaenia
is present in all trees one, two, and three steps longer than the most parsimonious
trees. Except for a clade composed of Girardinus metallicus, Poecilia sphenops, and
Phallichthys amates all of the groups within the Poeciliinae are incongruent with
trees one step longer than the most parsimonious trees. Relationships within the
Anablepidae are well supported.
Taxonomy
This analysis requires taxonomic changes (Table 4) to ensure that taxonomic

names represent monophyletic groups. Character states diagnosing higher taxonomic
groups represented by multiple species in this study are presented in Appendix 3.
The proposed taxonomy is not congruent with currently accepted subfamilial
classifications of the Poeciliidae (Parenti, 1981; Nelson, 1994; Costa, 1996). Aplo-
cheilichthys spilauchen is the sister group to a clade composed of all other included
poeciliids. Because Aplocheilichthys spilauchen is the type species for the genus Aplo-
cheilichthys this requires its recognition as the only member of the genus Aplocheilichthys
and the subfamily Aplocheilichthyinae Myers (1928). The distinctness of this species
when compared to the other species previously placed in Aplocheilichthys was em-
phasized by Huber (1982) who also recognized a monotypic Aplocheilichthys. The
subfamilial name Procatopodinae Fowler (1916) is applied to all species previously
recognized in the subfamily Aplocheilichthyinae (sensu Costa, 1996) except Aplo-
cheilichthys spilauchen.
The generic classification within the Procatopodinae also requires alteration from
the taxonomy proposed by Parenti (1981). The removal of Aplocheilichthys spilauchen
38 M. J. GHEDOTTI
Oxyzygonectes dovii
5
3 Anableps dowi
1 Anablepidae
3 2 Jenynsia unitaenia
1 Jenynsia multidentata
Aplocheilichthys spilauchen Aplocheilichthyinae
Micropanchax macropthalmus
1
3 Micropanchax pumilis
1 1
4 Micropanchax johnstoni
1
2 2 Micropanchax loati
1 Micropanchax pelagicus
1 Procatopodinae
'Micropanchax' schioetzi
1
3 Platypanchax cf. modestus
1
2 Lamprichthys tanganicus
1
3
1 4 Hypsopanchax zebra
Alfaro cultratus
6
Priapella compressa
3 1 Gambusia affinis
2 1 Heterandria formosa
1 Girardinus metallicus
2
Poeciliinae
3 Poecilia sphenops
1 1 Phallichthys amates
1 Phalloceros caudimaculatus
2 1 Cnesterodon decemmaculatus
2 Tomeurus gracilis
Figure 19. Strict consensus of three equally most parsimonious trees. Outgroups are not illustrated.
Tree statistics are based on the analysis performed using the outgroup topology proposed by Costa
(1998). The numbers above the branches are the Bremer (1988) decay indices. The numbers below
branches indicate unique and unreversed synapomorphies, if no number is given then there are no
unique and unreversed synapomorphies supporting that node. The thick branches indicate viviparous
lineages. The cross-hatched branch indicates the facultatively viviparous Tomeurus gracilis.
from the subfamily leaves the many species previously placed within the genus
Aplocheilichthys in need of new generic designation. Micropanchax loati, the type species
for the genus Micropanchax, is part of a monophyletic group including M. macropthalmus
(type of Poropanchax), M. pumilis (type of Lacustricola), M. johnstoni (recognized as the
only member of the unnamed genus ‘Aplocheilichthys’ by Parenti, 1981), and M.
pelagicus (type of Laciris). This group is recognized as the genus Micropanchax with
three subgenera, Poropanchax, Lacustricola, and Micropanchax (including M. loati, M.
johnstoni, and M. pelagicus). This proposal is similar to that made by Huber (1982)
except that the genus Laciris is placed into synonymy with Micropanchax and a new
genus for ‘Micropanchax’ schioetzi is recognized. Wildekamp (1995) previously suggested
that ‘M.’ schioetzi and related species formed a distinct group deserving of subgeneric
recognition within Aplocheilichthys. Jean H. Huber, Museum national d’Historie
naturelle, Paris, is currently describing a new genus that will contain ‘Micropanchax’
schioetzi, M.’ nimbaensis and ‘M.’ lamberti (Huber, pers. com.). Therefore, this species
is recognized in this work as belonging to a currently unnamed genus, ‘Micropanchax’,
in anticipation of a new genus. Platypanchax is removed from the synonymy of
Hypsopanchax. Hypsopanchax zebra and H. platysternus (the type species of Hypsopanchax)
can be identified as a monophyletic group based on the synapomorphy of distinct
narrow vertical lines on the flank. The phylogenetic relationships and generic
placement of other species currently recognized in Hypsopanchax, H. catenatus, H.
jobaerti, and H. jubbi, is unclear and these species should remain within Hypsopanchax
until a more detailed taxonomic and phylogenetic study of the genus is completed.
The current taxonomy of the Poeciliinae (Table 1) also requires modification to
be consistent with the proposed phylogeny. Tomeurus gracilis is placed in the tribe
Cnesterodontini, and the tribe and supertribe Tomeurini are not recognized. The
genera Alfaro and Priapella are removed from the Poeciliini and recognized in two
separate tribes, the Alfarini and Priapellini (new tribe) respectively. The tribe Alfarini
previously was recognized by Howell and Hubbs (1936), Rosen (1952), and Rosen
and Gordon (1953). The remaining members of the Poeciliini are recognized as
forming a monophyletic group as discussed by Rodriguez (1997), with the addition
of Phallichthys. The tribe ‘Heterandrini’ is recognized as a nonmonophyletic as-
semblage based on the phylogenetic relationships of Heterandria formosa (the type),
Poeciliopsis latidens, and Phallichthys amates. Phallichthys amates is the sister taxon of
Poecilia sphenops in all most parsimonious trees. Rosen and Bailey (1963: 117) also
suggested that Phallichthys relationships were uncertain and mentioned that the
similarity between Poeciliopsis and Phallichthys could have been derived independently.
Heterandria formosa and Poeciliopsis latidens are only tentatively recognized in the tribe
Heterandrini because they are sister taxa in just one of three equally most par-
simonious trees. The other taxa currently placed in the Heterandrini should be
retained there until a more detailed phylogenetic revision of the Poeciliinae including
many more taxa currently recognized in the Heterandrini is completed.
Priapellini new tribe

Composition: Priapella Regan, 1913 containing P. bonita (Meek, 1904), P. compressa
Alvarez 1948, P. intermedia Alvarez, 1952, P. olmecae Meyer and Spinosa Peréz, 1990.
Diagnosis: Same as for the genus as diagnosed and described by Rodriguez (1997).
DISCUSSION
The proposed phylogeny (Fig. 21A) supports monophyly of the Anablepidae and
the Poeciliidae (sensu Parenti, 1981). This contradicts the hypothesis proposed by
Meyer and Lydeard (1993) based on X-src DNA sequence data that the Poeciliinae
is the sister group of the Anablepidae. However, my results agree with the conclusions
of both Parenti (1981) and Meyer and Lydeard (1993) that viviparity has arisen
twice within the superfamily. Clausen (1959) discussed the remarkable degree of
similarity between some members of the Procatopodine genus Procatopus and the
Poeciliine genus Xiphophorus. Members of both genera exhibit sexual dichromatism,
40 M. J. GHEDOTTI
3
Oxyzygonectes dovii
2 5
4 Anableps dowi
7 Anablepidae
6 Jenynsia unitaenia
8
Jenynsia multidentata
10
1 Aplocheilichthys spilauchen Aplocheilichthyinae
13
16
Micropanchax macropthalmus
15 18
12 Micropanchax pumilis
17 20
9 Micropanchax johnstoni
19 22
21 Micropanchax loati
14 23
Micropanchax pelagicus
Procatopodinae
26
'Micropanchax' schioetzi
25 28
27 Platypanchax cf. modestus
11 29
24 Lamprichthys tanganicus
31
30 33
32 Hypsopanchax zebra
34 Procatopus nototaenia
36
Alfaro cultratus
35 38
Priapella compressa
37 40
Gambusia affinis
42
39 Heterandria formosa
44
41 47
46 Girardinus metallicus
43 49 Poeciliinae
48 Poecilia sphenops
50
45 Phallichthys amates
53
52 Phallotorynus victoriae
54
51 Phalloceros caudimaculatus
56
55 Cnesterodon decemmaculatus
57
Tomeurus gracilis
Figure 20. One of three equally most parsimonious trees. This topology is isomorphic with the majority-
rule consensus tree. Outgroups are not illustrated. Character state transformations are based on the
analysis performed using the outgroup topology proposed by Costa (1998) and ACCTRAN optimization.
The numbers above the branches refer to the character state transformations listed: (1) 2-1, 4-2, 10-
1, 22-1, 23-1, 88-1, 97-1; (2) 4-2, 9-1, 10-1, 11-3, 12-1, 24-1, 27-2, 55-0, 59-1, 64-0, 88-1, 94-1; (3)
2-1, 5-0, 30-1, 32-0, 35-1, 36-1, 87-1, 92-1; (4) 33-1, 67-2, 71-1, 72-2, 80-1, 99-1; (5) 16-1, 32-0, 36-
1, 55-2, 58-1, 62-2, 63-1, 66-1, 87-2; (6) 22-0, 30-1, 70-1, 75-1, 78-1; (7) 24-0, 31-0, 86-0; (8) 4-0, 71-
0; (9) 2-1, 3-2, 29-0, 42-1, 51-1, 53-1, 54-1, 57-1, 66-1, 90-1, 93-1, 95-1; (10) 4-2, 16-1, 88-1, 10-1,
47-1, 92-2; (11) 5-0, 25-1, 31-0, 34-1, 49-1, 58-1, 87-1; (12) 6-1, 8-1, 18-0, 26-1, 28-1, 40-1, 91-1, 97-
0; (13) 2-0, 22-2, 26-2, 49-0, 54-0, 69-2, 87-2, 89-1, 93-0, 96-1; (14) 1-1, 16-1, 47-1, 92-1; (15) 24-2,
30-1, 32-2, 52-1; (16) 7-1, 59-1, 90-0, 96-1; (17) 18-1, 31-1, 40-0; (18) 6-0, 14-1, 25-0, 28-0, 69-0, 86-
0, 89-1; (19) 19-1; (20) 17-1, 30-0, 32-1; (21) 27-1, 49-0, 96-1; (22) 54-0; (23) 13-1, 69-0, 86-0, 97-1;
(24) 27-1, 69-0, 97-1, 98-1; (25) 14-1, 89-1; (26) 18-1, 19-1, 28-0; (27) 6-0, 25-0, 52-1, 54-0, 59-1, 86-
0; (28) 8-0, 17-1, 27-0, 33-1, 37-1, 39-1; (29) 7-1, 31-1, 32-2, 48-1, 86-2, 89-0, 90-0, 91-0; (30) 8-0, 9-
2, 10-1, 15-1, 16-0, 33-1, 48-1, 56-1, 92-2; (31) 16-2, 20-1, 22-2, 26-2, 38-1, 41-1, 42-0, 43-1, 54-0,
62-1, 66-0, 96-1, 97-0; (32) 28-0, 30-1, 39-1, 63-2, 86-0, 91-0; (33) 2-0, 3-1, 31-1,37-1, 56-0, 86-2;
(34) 25-0, 49-0; (35) 3-0, 27-1, 30-1, 32-0, 35-2, 42-2, 43-1, 44-1, 45-1, 61-1, 63-1, 66-0, 67-1, 70-1,
71-1, 83-1, 84-1, 85-1, 95-0, 99-1; (36) 4-0, 37-1, 73-1, 79-1, 87-2, 88-1; (37) 46-1, 50-2, 74-1, 77-1,
82-1; (38) 30-0, 54-0, 55-0, 86-0, 98-1; (39) 2-0, 26-2, 27-0, 28-1, 81-2, 91-1, 93-0, 97-0; (40) 24-1,
males with dorsal or ventral caudal-fin extensions, and males that perform similar
mating displays. The caudal-fin extensions are almost certainly independently derived
in these two genera because they are absent in other poeciliid taxa. However, the
sister group relationship between the Procatopodinae and the Poeciliinae suggests
that the similar sexual dichromatism and mating displays may be the result of
common ancestry because both of these characteristics are widespread within the
Procatopodinae and the Poeciliinae and are absent in the Anablepidae. Further
behavioral studies need to be performed on a wide variety of poecilioid taxa to be
able to make a robust hypothesis of behavioral homology that when interpreted
within a phylogenetic context would be able support or reject this hypothesis.
The Procatopodinae is composed of two large clades that can be roughly
characterized by overall body form. One clade contains approximately seven genera
and is composed of laterally compressed, deep-bodied species, and the other clade
contains the primitively terete procatopodines in the genus Micropanchax (Fig. 21B).
Within each of these clades one species independently evolved pelagic habits in the
African Great Lakes (Fig. 21B). Lamprichthys tanganicus is a deep-bodied species living
in off shore areas of Lake Tanganyika, and Micropanchax pelagicus is a terete species
living in off shore areas of Lake Edward. The more pelagic behaviour of most
procatopodines, schooling and often swimming in open water, as compared to other
cyprinodontiforms (Clausen, 1967; Foster, 1967; White, 1983) may help to explain
why two pelagic species have evolved in this group when none have evolved in the
African aplocheilids.
The phylogeny of the Poeciliinae is somewhat consistent with the statement by
Hubbs (1924) that Gambusia is among the least specialized in the group and Tomeurus
is the most specialized (Fig. 21C). Based on this analysis, Alfaro is the most basal
genus in the Poeciliinae confirming the ‘primitiveness’ of this group suggested
previously by Hubbs (1931) and Howell and Hubbs (1936). More recently, Tomeurus
was thought to be the basal member of the Poeciliinae exhibiting an au-
tapomorphically unique, highly specialized gonopodium (Rosen & Bailey, 1963;
Parenti, 1981; Meyer & Lydeard, 1993). Rosen and Bailey (1963) and Meyer and
Lydeard (1993) suggested that facultative viviparity in Tomeurus is a retained ancestral
condition that represents an intermediate stage in the evolution of obligate viviparity.
However, the results of this study support a sister group relationship between
Tomeurus gracilis and Cnesterodon as well as the placement of this clade in a derived
position within the subfamily (Fig. 21C). Facultative viviparity in Tomeurus is thus
most parsimoniously interpreted as a novel condition unique to this species rather
than a retained ancestral condition.
There is significant contradictory evidence for the placement of Tomeurus gracilis
in this analysis as shown by Bremer decay index values of one for all nodes between
Tomeurus gracilis and the base of the poeciliine clade among Alfaro cultratus, Priapella
50-0, 76-1, 82-2, 83-0; (41) 3-2, 4-0, 11-0, 18-0, 21-1, 32-1, 41-1, 68-1, 96-1; (42) 3-2, 52-1; (43) 26-
0, 28-0, 65-1; (44) 15-1, 49-0, 62-1, 94-2; (45) 3-1, 4-1, 13-1, 38-1, 50-1, 81-0, 89-1; (46) 4-2, 20-1,
27-1, 30-2, 90-0, 91-0, 93-1; (47) 12-2, 72-2, 83-2; (48) 18-1, 32-0, 50-0, 81-1, 86-0, 87-0, 88-1; (49)
11-4, 44-2, 73-1, 25-0, 68-0, 82-2; (50) 4-0, 27-0, 28-1, 50-2, 67-1, 94-2, 96-0; (51) 26-2, 28-1, 52-1,
60-1, 65-2, 72-1; (52) 83-2; (53) 12-2; (54) 4-0; (55) 8-1, 18-1, 42-0, 45-0, 46-0, 56-1, 79-1, 85-2, 87-
0, 89-0; (56) 3-2, 76-1; (57) 11-1, 13-0, 21-0, 38-0, 40-1, 41-0, 44-2, 49-0, 52-0, 61-0, 62-1, 74-0, 87-
2, 96-0, 99-0.
42 M. J. GHEDOTTI
Aplocheilichthyinae
Fluviphylax obscurum – Fluviphylacini
Oxyzygonectinae Micropanchax macropthalmus
ANABLEPIDAE Micropanchax pumilis
Anablepinae
Micropanchax johnstoni
Aplocheilichthyinae
Procatopodini
Micropanchax loati
Procatopodinae POECILIIDAE Micropanchax pelagicus
Poeciliinae 'Micropanchax' schioetzi
Platypanchax cf. modestus
Lamprichthys tanganicus
Hypsopanchax zebra
A B Poeciliinae
Aplocheilichthyinae
Procatopodinae
Alfaro cultratus Alfarini
Priapella compressa Priapellini
Gambusia affinis Gambusini
Heterandria formosa
Heterandrini
Girardinus metallicus Girardini
Poecilia sphenops
Poeciliini
Phallichthys amates
Phalloceros caudimaculatus
Cnesterodontini
Cnesterodon decemaculatus
C Tomeurus gracilis
Figure 21. A, phylogenetic relationships of the subfamilies within the superfamily Poecilioidea. The
thick branches represent viviparous lineages. B, phylogenetic relationships of the Poeciliidae with
emphasis on the relationships within the Procatopodinae. The thick branches represent pelagic species
that live in the African Great Lakes. C, phylogenetic relationships of the Poeciliidae with emphasis on
the relationships within the Poeciliinae. The dash represents the inferred evolution of a more anterior
position of the pelvic girdle (and gonopodium) in males than in females (T.S. 63-1). The thick branches
represent lineages with fewer than six pelvic-fin rays (T.S. 60-1), a far anterior position of the pelvic
girdle (and gonopodium) (T.S. 63-2) in males and paired cirri on anal-fin ray three in males (T.S.
72-1).
compressa, and Gambusia affinis (Fig. 19). Most of this evidence for a basal position of
Tomeurus is based on the morphology of the oral and pharyngeal jaws (T.S. 11, 13,
21, 38, 41) and most of the evidence for placement in the Cnesterodontini is based
on the morphology of the gonopodium and gonopodial suspensorium (T.S. 45, 65,
72, 79, 85). The principle of parsimony supports the latter hypothesis and the
homoplastic reversals in jaw structure possibly may be explained as a developmental
by-product of reduced size.
The position of Tomeurus gracilis as a derived poeciliine closely related to Cnesterodon
suggests that the poeciliine gonopodium arose as a more simple structure similar in
position to the female anal fin that became anteriorly placed and more complex
during the evolutionary diversification of the group. Based on the phylogeny, it is
likely that gonopodium has become more anterior than the female anal fin in two
stages (T.S. 65). Somewhat more anteriorly positioned gonopodia are found in all
poeciliines except Alfaro, Priapella, and the tribe Gambusini, and the most anterior
gonopodia are found in the tribe Cnesterodontini (Fig. 21C). Similarly the gono-
podium can be inferred to have increased in length (T.S. 68) and developed some
Procatopodini
Aplocheil-
Fluviphylacini
ichthyinae
Poeciliinae
Anablepidae
Figure 22. Approximate distributions of poecilioid taxa and phylogenetic relationships illustrating trans-
Atlantic relationships.
terminal specializations (T.S. 72 & 74) after its origin in the ancestor of the
Poeciliinae.
Few detailed biogeographic conclusions can be drawn from this study because of
the representative nature of this analysis and the lack of information concerning the
relationships of genera and species not included. The sister group relationship
between Cnesterodon decemmaculatus (distributed in southern South America) and
Tomeurus gracilis (distributed in northern South America) is similar to the largely anti-
Amazonian pattern seen in Jenynsia (distributed in southern South America) and
Anableps (distributed in northern South America). This pattern is largely consistent
with the biogeographic pattern proposed by Costa (1995) for Neotropical fishes.
However, the absence of other cnesterodontin species (especially other species of
Cnesterodon) from this analysis makes this biogeographic interpretation only tentative.
The best supported biogeographic conclusions that can be drawn from this study
are based on the distribution of higher taxonomic groups as discussed by Parenti
(1981). The New World and African poecilioids do not segregate into two geo-
graphically separate clades. Based on the principles of vicariance biogeography
(Nelson & Rosen, 1981; Wiley, 1981) all hypothesized ancestors (including the
ancestor of the Procatopodinae) shown in Figure 22 are assumed to be distributed
in both Africa and the New World. Therefore, the phylogeny suggests an origin of
the Poecilioidea before the separation of South America and Africa.
ACKNOWLEDGEMENTS
I would like to express my appreciation to the Department of Ecology and

Evolutionary Biology, the Natural History Museum of the University of Kansas,
and the Regis University Department of Biology for support during this study. I
44 M. J. GHEDOTTI
also received support for travel to examine specimens at the Natural History Museum
in London from an Ernst Mayr Grant from the Harvard Museum of Comparative
Zoology and a Panorama Society Grant from the University of Kansas Natural
History Museum. I thank the curators and collections managers at institutions that
provided loans of specimens. I would also like to thank J.S. Ashe, W.W. Dimmick,
C. Fielitz, C. Haufler, L.R. Parenti, R.O. Prum, C.R. Robins, K.A. Shaw, K.L.
Tang, E. O. Wiley, and two anonymous reviewers for comment on various forms
of this manuscript.
REFERENCES
Agassiz JLR. 1853. Recent researches of Professor Agassiz. American Journal of Science 16: 134–136.
Ahl E. 1924. Neue afrikanische Zahnkarpfen aus dem zoologischen Museum Berlin. Zoologischer Anzeiger
61(3–4): 135–136.
Ahl E. 1928. Beiträge zur Systematik der africanischen Zahnkarpfen. Zoologischer Anzeiger 79(3–4):
115–116.
Alvarez J. 1948. Contribución al concocimiento del género Priapella y descripción de una nueva
especie (Pisces, Poeciliidae). Revista de Sociedad Mexicana de Historia Natural 9: 331–340.
Alvarez J. 1952. Cuatro especies nuevas de peces dulceacuicolas del suroeste de Mexico. Ciencia 11:
281–289.
Axelrod HR, Burgess WE, Pronek N, Walls JG. 1993. Dr. Axelrod’s atlas of freshwater aquarium fishes,
Seventh edition revised and expanded. Neptune City: T. F. H. Publications Inc.
Bleeker P. 1863. Mémoire sur les poissons de la Côte de Guinée. Naturkundige Verhandelingen van de
Hollandsche Maatschap van Wetenschappen, Haarlem 2(18): 116–117.
Bloch ME, Schneider JOG. 1801. Systema Ichthyologiae. Vols. 1–2. Berlin.
Bonaparte CL. 1831. Saggio di una distribuzione metodica degli animali vertebrati.
Boulenger GA. 1898. Report on the collection of fishes made by Mr. J.E.S. Moore in Lake Tanganyika
during his expedition, 1895–1896. Transactions of the Zoological Society, London 15(1): 1–30.
Boulenger GA. 1901. Diagnoses of new fishes, discovered by Mr. W.L.S. Loat in the Nile. Annals
and Magazine of Natural History 7(8): 446.
Boulenger GA. 1904. Descriptions of new West African freshwater fishes. Annals and Magazine of
Natural History 14(7): 16–20.
Boulenger GA. 1906. Fourth contribution to the ichthyology of Lake Tanganyika, Report on the
collection of fishes made by Dr. Cunningham, during the third Tanganyika expedition, 1904–1905.
Transactions of the Zoological Society, London 17(6): 554.
Breder CM Jr, Rosen DE. 1966. Modes of reproduction in fishes. Garden City, New York: The Natural
History Press.
Bremer K. 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction.
Evolution 42: 795–803.
Chambers J. 1987. The cyprinodontiform gonopodium, with an atlas of the gonopodia of the fishes
of the genus Limia. Journal of Fish Biology 30: 389–418.
Clark E, Aronson LR. 1951. Sexual behavior in the guppy, Lebistes reticulatus (Peters). Zoologica 36:
49–65.
Clausen HS. 1959. Description of two subgenera and six new species of Procatopus Boul., a little-
known West African genus of cyprinodont fishes. Videnskabelige Meddeleser fra Dansk Naturhistorisk
Forening I København. 121: 261–291.
Clausen HS. 1967. Tropical Old World cyprinodonts. Copenhagen: Akademisk Forlag.
Costa WJEM. 1995. Pearl killifishes, the Cynolebiatinae, systematics and biogeography of the neotropical annual
fish subfamily (Cyprinodontiformes: Rivulidae). Neptune City, New Jersey: T. F. H. Publications, Inc.
Costa WJEM. 1996. Relationships, monophyly and three new species of the neotropical miniature
poeciliid genus Fluviphylax (Cyprinodontiformes: Cyprinodontoidei). Ichthyological Exploration of Fresh-
waters 7(2): 111–130.
Costa WJEM. 1998. Phylogeny and classification of the Cyprinodontiformes (Euteleostei: Ather-
inomorpha): A reappraisal. In: Malabarba LR, Reis RE, Vari RP, Lucena ZMS, Lucena CAS,
eds. Phylogeny and classification of Neotropical fishes. Porto Alegre, Brazil: Edipuers.
Dawes J. 1991. Livebearing fishes: A guide to their aquarium care, biology, and classification. London: Blandford.
Dingerkus G, Uhler LD. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for
demonstration of cartilage. Stain Techniques 52: 229–232.
Duméril AHA. 1861. Reptiles et poissons de l’Afrique occidentale. Tome 10. Etude précedée de
considérations générals sur leur distribution géographique. Archives du Muséum d’Histoire Naturelle 10:
258.
Dyer BS, Chernoff B. 1996. Phylogenetic relationships among atheriniform fishes (Teleostei:
Atherinomorpha). Zoological Journal of the Linnean Society 117: 1–69.
Eigenmann CH. 1907. The poeciliid fishes of Rio Grande do Sul and the La Plata Basin. Proceedings
of the United States National Museum 32: 425–433.
Eigenmann CH. 1909. Reports on the expedition to British Guiana of the Indiana University and
the Carnegie Museum, 1908. Some new genera and species of fishes from British Guiana. Annals
of the Carnegie Museum 6(1): 4–54.
Foster NR. 1967. Trends in the evolution of reproductive behavior in killifishes. Studies in Tropical
Oceanography 5: 549–566.
Foster NR. 1983. The Tanganyikan lampeye, Lamprichthys tanganicus (Boulenger). Journal of the American
Killifish Association 16(5): 165–171.
Fowler HW. 1916. Notes on fishes of the orders Haplomi and Microcyprini. Proceedings of the Academy
of Natural Sciences, Philadelphia 68: 415–439.
Fromm D. 1982. The white eye, Oxyzygonectes dovii (Günther). Journal of the American Killifish Association
15: 201–210.
Garman S. 1895. The cyprinodonts. Memoirs of the Museum of Comparative Zoology 19: 1–170.
Ghedotti MJ. 1998. Phylogeny and classification of the Anablepidae (Teleostei: Cyprinodontiformes).
In: Malabarba LR, Reis RE, Vari RP, Lucena ZMS, Lucena CAS, eds. Phylogeny and classification
of Neotropical fishes. Porto Alegre, Brazil: Edipuers.
Ghedotti MJ, Weitzman SH. 1996. A new species of Jenynsia (Cyprinodontiformes: Anablepidae)
from Brazil with comments on the composition and taxonomy of the genus. Occasional Papers of the
Natural History Museum of the University of Kansas 179: 1–25.
Greenwood PH, Rosen DE, Weitzman SH, Myers GS. 1966. Studies of teleostean fishes, with
a provisional classification of living forms. Bulletin of the American Museum of Natural History 131:
345–354, 393–403.
Günther A. 1866. Catalog of the fishes of the British Museum—VI. London: Taylor & Francis.
Günther A. 1893. Second report on the reptiles, batrachians, and fishes, transmitted by Mr. H.H.
Johnston C.B. from British Central Africa. Proceedings of the Zoological Society, London 627–628.
Heckel J. 1848. Eine neue Gattung von Poecilien mit rochenartigem Anklammerungs-Organe.
Sitzungsber Akad Wiss Wien, Math Nat Cl 1: 289–303.
Henn AW. 1916. On various South American poeciliid fishes. Annals of the Carnegie Museum 10: 93–142.
Howell Rivero L, Hubbs CL. 1936. Studies of the fishes of the order Cyprinodontes. XV. The
characters and relationships of Furcipenis huberi and Alfaro cultratus. Occasional Papers of the Museum of
Zoology, University of Michigan 339: 1–11.
Hubbs CL. 1924. Studies of the fishes of the order Cyprinodontes. Miscellaneous Publications of the
Museum of Zoology, University of Michigan 13: 1–31.
Hubbs CL. 1926. Studies of the fishes of the order Cyprinodontes VI. Miscellaneous Publications of the
Museum of Zoology, University of Michigan 16: 1–86.
Hubbs CL. 1931. Studies of the fishes of the order Cyprinodontes. IX. A new and primitive genus
of Poeciliidae from Central America. Occasional Papers of the Museum of Zoology, University of Michigan
230: 1–3.
Hubbs CL. 1934. Studies of the fishes of the order Cyprinodontes. XIII. Quintana atrizona, a new
poeciliid. Occasional Papers of the Museum of Zoology, University of Michigan 301: 1–8.
Hubbs CL. 1950. Studies of cyprinodont fishes. XX. A new subfamily from Guatemala, with ctenoid
scales and a unilateral pectoral clasper. Miscellaneous Publications of the Museum of Zoology, University of
Michigan 78: 1–28.
Huber JH. 1982. (cover date 1981). A review of the cyprinodont fauna of the coastal plain in Rio Muni, Gabon,
Congo, Cabinda, and Zaire, with taxonomic shifts in Aphyosemion, Epiplatys and West African Procatopodins.
London: British Killifish Association.
Jordan DS. 1923. A classification of fishes including families and genera as far as known. Stanford
University Pubublications, University Series Biological Sciences 3(2): 77–243.
Kamel A-H. 1947. On the shifting of the trunk features into anterior part of the original tail region
during the later development of Gambusia affinis. Bull Facul Sci (Cairo) 26: 265–268.
46 M. J. GHEDOTTI
Kner R. 1860. Über Belonesox beliezanus, nov. gen. et spec. aus der Familie der Cyprinodonten.
Sitzungsber Akad Wiss Wein, Math Nat Cl 40(10): 419–422.
Lazara KJ. 1983. A taxonomic and nomenclatural overview of the subfamily Aplocheilichthyinae.
Journal of the American Killifish Association 16(5): 146–158.
Leviton AE, Gibbs RH Jr, Heal E, Dawson CE. 1985. Standards in herpetology and ichthyology:
Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.
Copeia 1985: 802–832.
Meek SE. 1904. The fresh-water fishes of Mexico north of the Isthmus of Tehuantepec. Publications
of the Field Columbian Museum, Zoology Series 5: 1–252.
Meek SE. 1912. New species of fishes from Costa Rica. Publications of the Field Columbian Museum,
Zoology Series 10(7): 69–75.
Meffe GK, Snelson FF Jr, eds. 1989. Ecology and Evolution of Livebearing Fishes (Poeciliidae). Englewood
Cliffs, New Jersey: Prentice Hall Inc.
Meinken H. 1932. Über einige neue Zahnkerpfen aus dem tropischen Westafrika. Blätter für Aquarien-
und Terrarienkunde 43(4): 53–54.
Meyer A, Lydeard C. 1993. The evolution of copulatory organs, internal fertilization, placentas,
and viviparity in killifishes (Cyprinodontiformes), as inferred from a DNA phylogeny of the tyrosine
kinase gene X-src. Proceedings of the Royal Society, London Series B 254: 153–162.
Meyer A, Morrissey JM, Schartl M. 1994. Recurrent origin of a sexually selected trait in Xiphophorus
fishes inferred from molecular phylogeny. Nature 386: 539–542.
Meyer MK, Espinosa Peréz H. 1990. Priapella olmecae sp.n., a new species from veracruz (Mexico).
Zool Abhandlungen Staatliches Museum für Tierkunde Dresden 45: 121–126.
Miller RR. 1979. Ecology, habits, and relationships of the Middle American cuatro ojos, Anableps
dowi (Pisces: Anablepidae). Copeia 1979: 82–91.
Mojica CL, Meyer A, Barlow GW. 1997. Phylogenetic relationships of species of the Genus
Brachyraphis (Poeciliidae) inferred from partial mitochondrial DNA sequences. Copeia 1997: 298–305.
Myers GS. 1924a. A new poeciliid fish of the genus Micropanchax from Ubangi. American Museum
Novitates 122: 1–3.
Myers GS. 1924b. New genera of African poeciliid fishes. Copeia 129: 42–43.
Myers GS. 1928. The systematic position of the phallostethid fishes, with diagnosis of a new genus
from Siam. American Museum Novitates 295: 1–12.
Myers GS. 1931. The primary groups of oviparous cyprinodont fishes, Order Cyprinodontes
(Microcyprini). Stanford University Publications 6: 1–14.
Myers GS. 1938. Studies on the genera of cyprinodont fishes. XIV. Aplocheilichthys and its relatives
in Africa. Copeia 1938: 136–143.
Myers GS. 1955. Notes on the classification and names of cyprinodont fishes, XIV. Tropical Fish
Magazine 1955 (March): 7.
Nelson G, Rosen DE. 1981. Vicariance biogeography: a critique. New York: Columbia University Press.
Nelson JS. 1994. Fishes of the world, 3rd Ed. New York: John Wiley & Sons.
Nichols JT, Griscom L. 1917. Freshwater fishes of the Congo basin obtained by the American
Museum Congo expedition. Bulletin of the American Museum of Natural History 37(25): 653–756.
Parenti LR. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei,
Atherinomorpha). Bulletin of the American Museum of Natural History 168: 335–557.
Parenti LR, Rauchenberger M. 1989. Systematic overview of the poeciliines. In: Meffe GK,
Snelson FF Jr, eds. Ecology and Evolution of Livebearing Fishes (Poeciliidae). Englewood Cliffs, New Jersey:
Prentice Hall Inc., 3–13.
Parker A. 1997. Combining molecular and morphological data in fish systematics: Examples from
the Cyprinodontiformes. In: Kocher TD, Stepien CA, eds. Molecular systematics of fishes. New York.
Academic Press, 163–188.
Patterson C. 1992. Supernumerary median fin-rays in teleostean fishes. Zoological Journal of the Linnean
Society 106: 147–161.
Poey F. 1854. Los guajacones, pecesillos de agua dulca. In: Poey F. Memorias sobre Historia Natural de
la Isla de Cuba, Volume 1. Havana, 347–392.
Poll M. 1952. Notes sur les Cyprinodontidae du Musée de Congo Belge. 2e partie: les Aplocheilichthyini
et les Lamprichthyini. Revue de Zoologie et de Botanique Africaines 45(3–4): 300–301.
Poll M. 1971. Un genre nouveau et une espece nouvelle de Cyprinodontidae congolaise. Revue de
Zoologie et de Botanique Africaines 83(3–4): 303–308.
Regan CT. 1911. The osteology and classification of teleostean fishes of the order Microcyprini.
Annals and Magazine of Natural History 8(7): 320–327.
Regan CT. 1913. A revision of the cyprinodont fishes of the subfamily Poeciliinae. Annals and Magazine
of Natural History 11: 977–1018.
Roberts TR. 1970. Description, osteology, and relationships of the Amazonian cyprinodont fish
Fluviphylax pygmaeus (Myers and Carvalho). Breviora 347: 1–28.
Rodriguez CMl. 1997. Phylogenetic analysis of the Tribe Poeciliini (Cyprinodontiformes: Poeciliidae).
Copeia 1997: 663–679.
Rosa RS, Costa WJEM. 1993. Systematic revision of the Genus Cnesterodon (Cyprinodontiformes:
Poeciliidae) with the description of two new species from Brazil. Copeia 1993: 696–708.
Rosa-Molinar E, Hendricks SE, Rodriguez-Sierra JF, Fritzsch B. 1994. Development of the
anal fin appendicular support in the western mosquitofish, Gambusia affinis affinis (Baird and Girard,
1854): A reinvestigation and reinterpretation. Acta Anatomica 151: 20–35.
Rosen DE. 1952. A revision of the fishes of the subfamily Alfarinae in the family Poeciliidae. Zoologica
37: 151–156.
Rosen DE. 1964. The relationships and taxonomic position of the halfbeaks, killifishes, silversides,
and their relatives. Bulletin of the American Museum of Natural History 127: 217–268.
Rosen DE. 1965. Oryzias madagascariensis Arnoult redescribed and assigned to the east African fish
genus Pantanodon (Atheriniformes, Cyprinodontoidei). American Museum Novitates 2240: 1–10.
Rosen DE. 1967. New poeciliid fishes from Guatemala, with comments on the origins of some South
and Central American forms. American Museum Novitates 2303: 1–15.
Rosen DE. 1979. Fishes from the uplands and intermontane basins of Guatemala: Revisionary studies
and comparative geography. Bulletin of the American Museum of Natural History 162: 269–375.
Rosen DE, Bailey RM. 1963. The poeciliid fishes (Cyprinodontiformes). Their structure, zoo-
geography and systematics. Bulletin of the American Museum of Natural History 126: 1–176.
Rosen DE, Gordon M. 1953. Functional anatomy and evolution of male genitalia in poeciliid fishes.
Zoologica 38: 1–47.
Scheel JJ. 1970. The genus Procatopus. Tropical Fish Hobbyist 18(168): 4–15.
Sethi RP. 1960. Osteology and phylogeny of oviparous cyprinodont fishes (order Cyprinodontiformes).
Unpublished D.Phil. Thesis, University of Florida.
Sorenson MD. 1996. TreeRot. Unpublished Computer Program. University of Michigan.
Stiassny MLJ. 1993. What are grey mullets? Bulletin of Marine Science 52(1): 197–219.
Swofford DL. 1993. PAUP: phylogenetic analysis using parsimony version 3.1.1. Unpublished Computer
Program. Illinois Natural History Survey.
Tigano C, Parenti LR. 1988. Homology of the median ethmoid ossifications of Aphanius fasciatus
and other atherinomorph fishes. Copeia 1988: 866–870.
Turner CL. 1941. Morphogenesis of the gonopodium in Gambusia affinis affinis. Journal of Morphology
69: 161–185.
Turner CL. 1950. The skeletal structure of the gonopodium and gonopodial suspensorium of Anableps
anableps. Journal of Morphology 86: 329–366.
Weitzman SH. 1974. Osteology and evolutionary relationships of the Sternoptychidae with a new
classification of stomiatoid families. Bulletin of the American Museum of Natural History 153: 327–478.
Weitzman SH. 1982. Fluviphylax pygmaeus, a miniature killifish from tropical South American
freshwaters. Journal of the American Killifish Association 15: 190–199.
Weitzman SH, Vari RP. 1988. Miniaturization in South American freshwater fishes: an overview
and discussion. Proceedings of the Biological Society of Washington 101: 444–465.
White S. 1983. The genus Aplocheilichthys Bleeker: A hobbyist’s introduction. Journal of the American
Killifish Association 16: 176–184.
Whitehead PJP. 1962. The Pantanodontinae, edentulous toothcarps from east Africa. Bulletin of the
British Museum of Natural History, Zoology 9(3): 105–137.
Whitley GP. 1951. New fish names and records. Proceedings Royal Zoological Society, New South Wales
1949–1950: 61–68.
Whitley GP. 1965. Some fish genera scrutinized. Proceedings Royal Zoological Society, New South Wales
1964–1965: 25–26.
Wildekamp RH. 1995. A world of killies, Atlas of the oviparous cyprinodontiform fishes of the world, Volume 2.
Mishawaka, Indiana: American Killifish Association.
Wiley EO. 1981. Phylogenetics, the theory and practice of phylogenetic systematics. New York: John Wiley &
Sons.
Wischnath L. 1993. Atlas of livebearers of the world. Neptune City: T. F. H. Publications Inc.
Worthington EB. 1932. Scientific results of the Cambridge expedition to the East African lakes
1930–1931. 2. Fishes other than Cichlidae. Journal of the Linnean Society, London 38(258): 121–134.
Wourms JP. 1981. Viviparity: the maternal-fetal relationship in fishes. American Zoologist 21: 473–515.
48 M. J. GHEDOTTI
APPENDIX 1: MATERIAL EXAMINED
The total number of specimens examined is indicated after the catalog number. The number of
cleared and stained specimens examined follows this number in parentheses.
Alfaro cultratus KU 11122, 25 (3); Costa Rica: Heredia: rı́o Santa Clara.
Anableps anableps MCZ 31107, 8 (2); Surinam. UMMZ 108901, 3; Guyana, Beatrice River. UMMZ
191789, 4 (2); aquarium specimens.
Anableps dowi ANSP 91678, 8 (2); Honduras, rı́o Nova Mulca. CAS 148298, 9 (3); México, Oaxaca,
rı́o Tequisistlan. FMNH 12114, 4; El Salvador, San Marcos, rı́o Lempa. KU 18689, 14 (3); Nicaragua,
Chinandega, rı́o Estero Real. UMMZ 190543, 12 of 68 (2); Guatemala, Santa Rosa & Jutiapa, rı́o
Los Esclavos. UMMZ 191725, 12 of 15 (3 skel.); México, Oaxaca, rı́o Tehuantepec.
Aplocheilichthys spilauchen CAS 163435, 5 of 36; Ghana, Avedzake Creek. CAS 163440, 5 of 83 (2);
Ghana, mouth of Volta River. USNM 195961, 5; Nigeria. USNM 246177, 4, Togo, Zebe. USNM
304139, 50 (3); Cameroon, rı́o del Rey.
Cnesterodon decemmaculatus KU 22411, 6 (2); Uruguay, Montevideo, Pajas Blancas. KU 22415, 9;
Uruguay, Montevideo, Santiago Vasquez.
Crenichthys baileyi KU 11862, 111 (3); USA, Nevada, Preston Springs. KU 14031, 43 (4); USA,
Nevada, irrigation ditch in Preston.
Cubanichthys pengelleyi UMMZ 166660, 2 of 6 (2); Jamaica, 6 mi. SE of Black River. UMMZ 213381,
5 of 22; Jamaica, St. Elizabeth, 1–2 mi. NE of Middle Quarters.
Cyprinodon variegatus KU 17040 55 (3); USA, Texas, Compano Bay. KU 17117 49 (4); USA, Georgia,
Catherine’s Island. KU 19634 36 (2); USA, Florida, Lake Eustis.
Fluviphylax obscurum CAS 36639, 5 of 7 (2); Brazil, Amazonas, rio Taruma.
Fluviphylax simplex MCZ 46713, 12 of 40 (3), MCZ 49958, 2 of 4 (2); Brazil, Pará, lago Parauacuı́.
Fundulus chrysotus KU 18165, 59 (6); USA, Florida, Hogeye Sink.
Gambusia affinis KU 16320, 162 (4); USA, Texas, San Gabriel River. KU 22033, 30 (4); USA,
Nebraska, Schuyer City Park.
Girardinus metallicus UMMZ 103333, 5 of 30; Cuba, Pinar del Rio, rı́o TacoTaco. UMMZ 108685,
4 of 21 (4); Cuba, Santa Clara, arroyo Los Gansos. UMMZ 158697, 7 of 29 (2), Cuba, Santa Clara,
cienega de Zapata.
Hylopanchax sylvestris CAS 163380, 10 of 36; Congo, Orientale, Stanleyville.
Hypsopanchax platysternus AMNH 6078, 2 paratypes; Congo, Orientale, Stanleyville.
Hypsopanchax zebra USNM 191521, 10 of 22 (4); French Equatorial Africa.
Jenynsia lineata BMNH 1843.2.8,43, 3 syntypes, BMNH 1917.7.14, 20–23, 4 syntypes; Uruguay,
Maldonado. AMNH 73235, 2; Uruguay, Florida, arroyo Molles. FMNH 71514, 105427–105428, 19;
Uruguay, Rocha y Minas, rı́o Cebollati, Passo Averias. FMNH 71515, 105429, 50 (4); Uruguay,
Minas, arroyo Polanco.
Jenynsia multidentata BMNH 1917.7.14.24, holotype; Uruguay, Montevideo. CAS 47062, 4 (4); Brazil,
Rio Grande do Sul. MZUSP uncat., 7, USNM 300246, 8; Uruguay, Maldonado, mouth of arroyo
Maldonado at Punta del Este. KU 22416, 49 (4); Uruguay, Montevideo, Santiago Vasquez at the
mouth of rı́o Santa Lucia. KU 19190, 60 (6); Argentina, Mendoza, Dique Benegas. KU 23872, 23;
Argentina, Buenos Aires, arroyo Pescado. KU 23873, 17; Argentina, Buenos Aires, rı́o de La Plata at
Punta. NMC uncat., 200 (23); Brazil, Rio Grande do Sul, Bosque, rio Grande.
Jenynsia unitaenia MZUSP 47615, holotype, KU 23866, 15 paratypes, MZUSP 47616–47620,
66 paratypes, MCP 17419–17421, MCP 17425, 54 paratypes, USNM 300232, 50 (5), USNM
300243–300244, USNM 300250, USNM 300297–300298, 63 paratypes; Brazil, Santa Catarina,
municı́pio de Praia Grande, rio Faxinalzinho at Mãe dos Homens. KU 23865, 10 paratypes, MCP
17422–17424, 91 paratypes, MZUSP 47621–47624, 124 paratypes, USNM 300238, USNM 300241,
USNM 300245, USNM 300299, 161 paratypes (4); Brazil, Santa Catarina, rio Jordão in Jordão Alto.
USNM 300731, 2; Brazil, Santa Catarina, rio Pique, Morro Cortado, on road between Meleiro and
Ararangua. USNM 326070, 5 paratypes; Brazil, Santa Catarina, municı́pio de Praia Grande, rio
Canoas approx. 8 km from Praia Grande in the direction of Mãe dos Homens. USNM 326104, 3
paratypes; Brazil, Rio Grande do Sul, municı́pio de Terra de Areira, rio Tres Pinhêiros on BR 101.
Lamprichthys tanganicus AMNH 38420, 1 (1); Lake Tanganyika. AMNH 38421, 1 (1); Lake Tanganyika.
UMMZ 199863, 8 of 18 (2); Zambia, Northern, Lake Tanganyika.
Micropanchax johnstoni ANSP 83282, 10 of 323 (4); Zimbabwe, Kaufe River. UMMZ 200190, 20 of
62 (4); Zambia, Northern, Chambeshi River. USNM 246173, 10 of 25 (4); Botswana, Xugana.
Micropanchax loati USNM 94205, 10 of 36 (4); Egypt, Gezira aquarium.
Micropanchax macropthalmus USNM 323614, 10 of 43 (4); Benin, Iquidi River. USNM 323621, 2 of
12 (2); Togo, Togble-Kope.
Micropanchax myersi ANSP 75219, 2; Zaire, Leopoldville.
Micropanchax pelagicus FMNH 62956, 10 of 18 (2); Zaire, Nord-Kivu, Semliki River.
Micropanchax pumilis USNM 84158, 10 of 49 (4); Lake Tanganyika.
‘Micropanchax’ schioetzi USNM 260051, 14; Ghana, Ankobra River. USNM 323612, 28 (5); Ghana,
Adansu River.
Oxyzygonectes dovii AMNH 17657, 8 (2), AMNH 37733, 12 of 25 (2); Costa Rica, Golfito. ANSP
140669, 6; Costa Rica, North Side of Golfito, 8°50′N 83°6′W. KU 18174, 13 (5); Panama, Veraguas,
rı́o San Pablo.
Pantanodon madagascariensis AMNH 20526, 2 (1) paratypes; Madagascar, Mahambo.
Phalloceros caudimaculatus UMMZ 11 of 310 (11), Paraguay, Paraguari, small stream trib. to arroyo
Yby. UMMZ 206426, 5 of 29; Paraguay, Canendiyu, 1.3 km W of Salto del Guaira.
Phallotorynus victoriae UMMZ 14 of 94 (4), Paraguay, San Pedro, rı́o Aguaray.
Platypanchax cf. modestus MCZ 100634, 10 of 43 (4); Uganda, Toro, Mpanga River.
Poecilia sphenops KU 11118, 65 (4); Costa Rica, Heredia, rı́o Santa Clara. KU 22711, 31 (4);
Nicaragua, laguna de Apoyo.
Poeciliopsis latidens KU 4416, 30; Mexico, Sonora, rı́o Cachujaqui. KU 5465, 158 (6); Mexico, Sonora,
rı́o Alamos.
Procatopus nototaenia CAS 147714, 5 of 7; Cameroon, Lokunje River. CAS 155515, 5 (2); Cameroon,
Lobi River system. USNM 92967, 2 cotypes; Cameroon, tributary to Lobi River.
Profundulus labialis KU 18163, 88 (7); Guatemala, rı́o Carchada.
Profundulus punctatus KU 18177, 67 (4); Guatemala, rı́o Camaya.
Tomeurus gracilis AMNH 28619, 10 (5); aquarium specimens. AMNH 48639, 4 (4); Suriname,
Corantijn River. UMMZ 167944, 15 of 44 (4); Guyana, aquarium specimens.
Valencia hispanica AMNH 38401, 3 of 5 (2); Spain.
Valencia letourneuxi UMMZ 213901, 8; Greece, Epirus, Louros River. UMMZ 213902, 13 of 24 (3);
Greece, Epirus, Acheron Basin.
50 M. J. GHEDOTTI
APPENDIX 2: DATA MATRIX
Character state data matrix for 29 poecilioid taxa and seven outgroup taxa. See text for discussion
of transformation series. All transformation series are unordered. Taxa with polymorphic character
states for a given transformation series are indicated with a letter (A=0&1; B=0&3, C=1&2).
1 1111111112 2222222223 3333333334
1234567890 1234567890 1234567890 1234567890
Oxyzygonectes dovii 0102000011 B100000100 0111002011 1000110000

Anableps dowi 0002100011 BA??010100 0111002010 1010010000
Jenynsia unitaenia 0002100011 3100000100 0010002011 0110000000
Jenynsia multidentata 0000100011 3100000100 0A11002011 1110000000
Aplocheilichthys spilauchen 0122100001 1000010100 0A10000000 1100000000
Fluviphylax obscurum 0021?10100 1000000000 0210120100 0101000001
Micropanchax macropthalmus 1121?11100 1000010000 0112110101 0201000001
Micropanchax pumilis 1121000100 1001010100 0112010001 1201000000
Micropanchax johnstoni 1121?10100 1000011110 0112110100 1101000000
Micropanchax loati 11210A0100 1000010110 0112111101 1201000000
Micropanchax pelagicus 1121?10100 1010010110 0112111101 1201000000
‘Micropanchax’ schioetzi 1121?10100 1001010110 0110111000 0101000001
Platypanchax cf. modestus 1121000000 1001011000 0110010100 0111001011
Lamprichthys tanganicus 1121001100 1001010000 0110011100 1201000001
Pantanodon madagascariensis 1121?10021 1000120001 0210121100 0111???101
Hypsopanchax zebra 1011?10021 1000100000 0110111001 1111001011
Procatopus nototaenia 1121?10021 1000100000 01100A1001 0111000011
Alfaro cultratus 0100000000 1000000100 0110101001 0001101000
Priapella compressa 0101000000 1000000100 0110101000 0001100000
Gambusia affinis 0001000000 1000000100 0111120101 0001100000
Heterandria formosa 0020000A00 0000000000 1110120102 0101100000
Poeciliopsis latidens 0020000000 0000100000 1110100001 0101100000
Girardinus metallicus 0012000000 0210000001 1110101002 0101100100
Poecilia sphenops 0012000000 4010000101 1110001002 0001100100
Phallichthys amates 0010000000 0010000101 1110100102 0001100100
Phallotorhynus victoriae 00C1000000 0210000000 1110120101 0101100100
Phalloceros caudimaculatus 0010000000 0010000000 1110120101 0101100100
Cnesterodon decemmaculatus 0021000100 0010000100 1110120101 0101100100
Tomeurus gracilis 0011000100 1000000100 0110120101 0101A00001
Cubanichthys pengelleyi 0001000100 BA01010100 0000022001 1000000000
Cyprinodon variegatus 1021000100 3201010000 0000112010 1010100100
Valencia hispanica 0001100000 1000000100 0000020010 1000010000
Valencia letourneuxi 0001100000 1000000100 0000000010 1100000000
Fundulus chrysotus 0002100000 1000020100 0000000010 0100100000
Crenichthys baileyi 0001000010 2000020000 0000000000 0110000010
Profundulus labialis 0000000000 0000000000 0000000000 0000000000
APPENDIX 2—continued
4444444445 5555555556 6666666667 7777777778

1234567890 1234567890 1234567890 1234567890
Oxyzygonectes dovii 0000000000 0000000010 0000000010 0000000000

Anableps dowi 0000000000 0000200110 0210012010 1000002001
Jenynsia unitaenia 0000000000 0000000010 0000002011 1000102101
Jenynsia multidentata 0000000000 0000000010 000A002011 0000102101
Aplocheilichthys spilauchen 0100001000 1011101000 0001010010 0000000000
Fluviphylax obscurum 0100000000 101010A100 0001010020 0000000000
Micropanchax macropthalmus 0100001010 1111101110 0001010010 0000000000
Micropanchax pumilis 0100001010 1111101100 0001010000 0000000000
Micropanchax johnstoni 0100001010 1111101100 0001010010 0000000000
Micropanchax loati 0100001000 1110101100 0001010010 0000000000
Micropanchax pelagicus 0100001000 1111101100 0001010000 0000000000
‘Micropanchax’ schioetzi 0100001010 1011101100 0001010000 0000000000
Platypanchax cf. modestus 0100001010 1110101110 0001010000 0000000000
Lamprichthys tanganicus 0100001110 1110101110 0001010000 0000000000
Pantanodon madagascariensis 1010001110 1010111100 0101000000 A000000000
Hypsopanchax zebra 0100001110 1011101100 0021010000 0000000000
Procatopus nototaenia 0100001100 1011111100 0021010000 0000000000
Alfaro cultratus 0211100010 1011101100 1011001011 1010000010
Priapella compressa 0211110012 1010001100 1011001011 1001001000
Gambusia affinis 0211110010 10111A1100 1011001011 1001011000
Heterandria formosa 1211110012 1111101100 1011001111 1001001000
Poeciliopsis latidens 1211110002 10111A1100 1111101111 1001001000
Girardinus metallicus 1211110011 1011101100 1011101111 1201001000
Poecilia sphenops 1212110010 1011101100 1011101011 1011001000
Phallichthys amates 1211110012 1011101100 1111101111 1001001000
Phallotorhynus victoriae 1211110011 1111101101 1011201111 1101001000
Phalloceros caudimaculatus 1211110011 1111101101 1011201111 1101001000
Cnesterodon decemmaculatus 1011000011 1111111101 1011201111 1101011010
Tomeurus gracilis 0012000001 10??111101 0111201111 1100001010
Cubanichthys pengelleyi 0110001010 0100001000 0001000010 0000000000
Cyprinodon variegatus 0010000010 0000100002 0021000010 0000000000
Valencia hispanica 0000000000 0110100010 00000100A0 0000000000
Valencia letourneuxi 0000000000 0110100010 0000010010 0000000000
Fundulus chrysotus 0000000000 0100100000 0000000010 0000000000
Crenichthys baileyi 1300000000 001000001? ??????00A0 0000000000
Profundulus labialis 0000000000 0000000000 0000000000 0000000000
52 M. J. GHEDOTTI
APPENDIX 2—continued
8888888889 999999999
1234567890 123456789
Oxyzygonectes dovii 0000011100 010100100

Anableps dowi 00000A2100 000100101
Jenynsia unitaenia 0000000100 000100101
Jenynsia multidentata 0000010A00 000100101
Aplocheilichthys spilauchen 0000010101 021010100
Fluviphylax obscurum 0000012011 100011000
Micropanchax macropthalmus 0000011000 111011000
Micropanchax pumilis 0000001011 111010000
Micropanchax johnstoni 00000110A1 111010000
Micropanchax loati 0000011001 111011000
Micropanchax pelagicus 0000001001 1??011100
‘Micropanchax’ schioetzi 0000011011 A11010110
Platypanchax cf. modestus 0000001011 1??010110
Lamprichthys tanganicus 0000021000 011010110
Pantanodon madagascariensis 0000011001 1?1011010
Hypsopanchax zebra 0000021001 0??010110
Procatopus nototaenia 00000010A1 021010110
Alfaro cultratus 0011112101 001000101
Priapella compressa 0111101001 001000111
Gambusia affinis 2201111A01 100000001
Heterandria formosa 2111111001 100001001
Poeciliopsis latidens 2111111001 A00201001
Girardinus metallicus 012111101A 001001001
Poecilia sphenops 1211100110 001001001
Phallichthys amates 111110011A A01200001
Phallotorhynus victoriae 0121111011 100001001
Phalloceros caudimaculatus 01211110A1 100001001
Cnesterodon decemmaculatus 01112100A1 100001001
Tomeurus gracilis 0111212001 10000000A
Cubanichthys pengelleyi 0000010000 000000010
Cyprinodon variegatus 0000010000 001021110
Valencia hispanica 0000000001 10?000000
Valencia letourneuxi 00000A00A1 10?000000
Fundulus chrysotus 0000010000 000000000
Crenichthys baileyi 0000010000 000000000
Profundulus labialis 0000000000 000000000
APPENDIX 3: DIAGNOSTIC SYNAPOMORPHIES
Character states diagnosing higher taxonomic groups represented by multiple species in this study.
Character states that were unique and unreversed in this analysis are indicated with an asterisk (∗).
Putative reversals are noted
Poecilioidea 1. Elongate ventral process of anguloarticular (T.S. 23-1)∗
Anablepidae 1. Individually variable dextral and sinistral laterality in males (T.S. 94-
1)∗
2. Broad prootic bridge (T.S. 9-1)
3. Tricuspid outer row of teeth (T.S. 11-3)
4. Tricuspid inner rows of teeth (T.S. 12-1)
5. Broad ventral postcleithrum (T.S. 59-1)
Anablepinae 1. Tubular gonopodium formed from four or more rays (T.S. 67-2)∗
2. Sixth anal-fin ray unbranched in adult males (T.S. 77-1)∗
3. First proximal anal radial fused to second in males (T.S. 80-1)∗
4. Vomer contacts lateral ethmoids (T.S. 5-1)
5. Broad basihyal (T.S. 33-1)
6. Viviparity (T.S. 99-1)
Jenynsia 1. Fifth anal-fin ray short (T.S. 75-1)∗
2. Seventh middle anal radial enlarged (T.S. 78-1)∗
3. Second anal-fin ray short (T.S. 70-1)
Poeciliidae 1. High dorsal insertion of pectoral fins (T.S. 51-1)∗
2. Supracleithrum lacks flange (T.S. 53-1)
3. Posteroventral coracoid rounded (T.S. 57-1, 0 in Fluviphylax obscurum)
4. Anterior position of pelvic girdle (T.S. 64-1)
Poeciliinae 1. Gonopodium formed from anal-fin rays 3–5 (T.S. 67-1)∗
2. Proximal anal radials 7–9 in males lack anterior and posterior flanges
(T.S. 84-1)∗
3. Anterior margin of first hypobranchial concave (T.S. 35-1)
4. Exoccipital condyles absent (T.S. 43-1)
5. Ligastyle in adult males (T.S. 44-1)
6. Gonapophyses present (T.S. 45-1, 0 in Cnesterodon and Tomeurus)
7. Ventral flange on male pelvic bone (T.S. 61-1, in Tomereus)
8. Posterior process on pelvic bone absent (T.S. 63-1)
9. Second anal-fin ray short (T.S. 70-1)
10. Third anal-fin ray unbranched (T.S. 71-1)
11. Lateral processes on ventral proximal radials in males (T.S. 85-1)
12. Viviparity (T.S. 99-1)
Cnesterodontini 1. Less than six pelvic-fin rays (T.S. 60-1)∗
2. Male pelvic girdle far anterior, under pectoral girdle (T.S. 65-2)∗
3. Paired bony cirri on the third anal-fin ray in males (T.S. 72-1)∗
4. Dorsoposterior region of lacrymal very narrow (T.S. 26-2)
Procatopodinae 1. Vomer unossified (T.S. 6-1, 0 in Platypanchax sp., Lamprichthys tanganicus,
Micropanchax pumilis, and some M. loati)
2. Interarcual cartilage absert (T.S. 40-1, 0 in Micropanchax johnstoni M.
pelagicus, M. loati and M. pelagicus)
3. Adult males larger than adult females (T.S 95-1)
Procatopodini 1. Mesethmoid unossified (T.S. 1-1)
2. Dorsoposterior region of lacrymal broad (T.S. 26-1, 0 in some Procatopus
nototaenia, 2 in Pantanodon madagascariensis)
3. First hemal arch expanded (T.S. 47-1)
Genus Micropanchax 1. Narrow elongate retroarticular (T.S. 24-2)∗
Subgenus Micropanchax 1. Dentary deep near symphysis with ventral expansion (T.S. 19-1)

Phylogenetic Analysis and Taxonomy of The Poeciloid Fishes

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Phylogenetic Analysis and Taxonomy of The Poeciloid Fishes

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Zoological Journal of the Linnean Society (2000), 130: 1–53.

doi:10.1006/zjls.1999.0213, available online at http://www.idealibrary.com on

Phylogenetic analysis and taxonomy of

Natural History Museum, University of Kansas, Lawrence, KS 66045-2454, U.S.A.

Received July 1998; accepted for publication July 1999

ADDITIONAL KEY WORDS:—Poecilioidea – morphology – phylogeny – Poeciliidae –

The superfamily Poecilioidea is a diverse group of cyprinodontiform fishes com-

TAXONOMIC HISTORY OF THE POECILIOIDEA

The Family Anablepidae currently contains three genera of fishes distributed in

Subfamily Poeciliinae Garman, 1895

Aplocheilichthyinae and Procatopodinae

The subfamilies Aplocheilichthyinae and Procatopodinae contain more than 100

T 2. Classification of the taxa included in the subfamily Aplocheilichthyinae as proposed by

pelagicus, a pelagic species endemic to Lake Edward. Huber (1982) recognized

However, I recognize both family group names as separate, subfamilial-rank taxa.

Specimens examined in this study are listed in Appendix 1. Standard institutional

mesethmoid to diagnose the suborder Aploceiloidei. However, Tigano and Parenti

parietals as synapomorphic for all cyprinodontids except Cubanichthys and potentially

Jaws and suspensorium

Meckel's cartilage retroarticular

present (Fig. 5C). Parenti (1981) recognized a robust dentary as a synapomorphy

ventral anterior posterior

urohyal anterior ceratohyal

34. Anterior process of anterior ceratohyal extending ventral to ventral hypohyal:

bb1 35-0 35-1

not recognized as a separate transformation series. In Crenichthys baileyi, the fifth

Vertebrae and ribs

1st vertebral pleural rib

Pectoral fin and girdle

Pelvic fin and girdle

(1981) recognized an anteriorly positioned pelvic girdle as synapomorphic for

Anal fin and skeletal supports

(1960) suggested that an increased number of anal-fin rays in Procatopus, Hypsopanchax,

Dorsal and caudal fins and skeletal supports

A broad caudal accessory cartilage is found in Micropanchax pumilis, ‘M.’ schioetzi,

that there is ‘significant’ maternal nutrient contribution to developing embryos.

This analysis requires taxonomic changes (Table 4) to ensure that taxonomic

Priapellini new tribe

I would like to express my appreciation to the Department of Ecology and

APPENDIX 1: MATERIAL EXAMINED

APPENDIX 2: DATA MATRIX

Oxyzygonectes dovii 0102000011 B100000100 0111002011 1000110000

4444444445 5555555556 6666666667 7777777778

Oxyzygonectes dovii 0000000000 0000000010 0000000010 0000000000

Oxyzygonectes dovii 0000011100 010100100

APPENDIX 3: DIAGNOSTIC SYNAPOMORPHIES

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