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Neotropical Ichthyology, 3(1):1-60, 2005
Copyright © 2005 Sociedade Brasileira de Ictiologia
1
PROOFS
* Laboratório de Ictiologia Sistemática, Universidade Federal do Tocantins, Campus de Porto Nacional, rua 3, Quadra 17, s/n, Caixa Postal
136, 77500-000 Porto Nacional, TO, Brazil. e-mail:lucinda@uft.edu.br
** Laboratório de Ictiologia, Pontifícia Universidade Católica do Rio Grande do Sul. Av. Ipiranga, 6681, Caixa Postal 1429, 90619-900 Porto
Alegre, RS, Brazil. e-mail: reis@pucrs.br
Systematics of the subfamily Poeciliinae Bonaparte
(Cyprinodontiformes: Poeciliidae), with an emphasis
on the tribe Cnesterodontini Hubbs
Paulo Henrique Franco Lucinda* and Roberto E. Reis**
Osteological and soft anatomical features of representatives of poeciliine genera were studied to test the monophyly of the
poeciliine tribes and to advance a hypothesis of relationships within the subfamily. The resultant hypothesis supports the
proposal of a new classification for the subfamily Poeciliinae. Diagnoses are provided for suprageneric clades. The tribe
Tomeurini is resurrected and the new tribes Brachyrhaphini and Priapichthyini as well as the supertribe Poeciliini are described.
New usages of old tribe names are proposed based on the phylogenetic framework.
Caracteres osteológicos e da anatomia mole de representantes dos gêneros de poeciliíneos foram estudados para se testar a
monofilia das tribos de Poeciliinae e para propor uma hipótese de relações dentro da subfamília. A hipótese resultante suporta
a proposição de uma nova classificação para a subfamília Poeciliinae. São fornecidas diagnoses para os clados supragenêricos.
A tribo Tomeurini é ressuscitada e as novas tribos Brachyrhaphini e Priapichthyini bem como a supertribo Poeciliini são
descritas. Novos usos para antigos nomes de tribos são propostos com base no arranjo filogenético.
Key words: Alfarini, Brachyrhaphini, Gambusiini, Girardinini, Heterandriini, Priapellini, Priapichthyini, Poeciliini, Tomeurini.
Introduction
This paper is resultant from a project that intended to
perform the taxonomic revision of the tribe Cnesterodontini,
as well as to propose a phylogenetic hypothesis of rela-
tionships among its members. Testing the monophyly of
the Cnesterodontini was the first step attempting to achieve
our aims. Thus, the original project was broadened to em-
brace a phylogenetic hypothesis of relationships and the
proposal of a provisional classification for the subfamily
Poeciliinae. The resultant hypothesis included representa-
tives of all poeciliine genera, and all described
Cnesterodontini species, as well as 24 new species revealed
by the taxonomic revisions of genera Cnesterodon Garman,
Phallotorynus Henn, and Phalloceros Eigenmann.
Intrageneric relationships of Phalloptychus Eigenmann,
Cnesterodon, Phalloceros, and Phallotorynus are provided
herein, but will be discussed in the aforementioned taxo-
nomic revisions.
Nomenclatural and Taxonomic History
Poeciliinae. The subfamily Poeciliinae is a cyprinodontiform
group widely distributed throughout the Americas. Poeciliinae
is the sister group of the Procatopodinae, a group composed
of the South-American Fluviphylax Whitley and the African
procatopodines. The clade Poeciliinae plus Procatopodinae
is the sister group of the Aplocheilichthyinae (Costa, 1996;
Ghedotti, 2000). These three subfamilies compose the family
Poeciliidae. The Poeciliinae embraces approximately two hun-
dred twenty species currently allocated in approximately
twenty-eight genera (Lucinda, 2003). Theses fishes are char-
acterized by the uniquely derived possession of a
gonopodium formed by the modified male anal-fin rays 3, 4,
and 5 (Parenti, 1981).
The Poeciliinae includes well-known aquarium fishes
such as the guppies, mosquito fishes, swordtails, platys,
and mollies. Poeciliines are well known subjects of study for
ecologists, anatomists, embryologists, and other research-
Systematics of the subfamily Poeciliinae Bonaparte
2
PROOFS
ers. Notwithstanding, this fish assemblage is only superfi-
cially studied from the perspective of systematics.
Intrageneric diversity and intergeneric relationships of the
Poeciliinae are poorly known, regardless of its huge distri-
bution, range, and notoriety. Similarly, phylogenetic hypoth-
eses for most genera are still wanting.
Taxonomic revisions and phylogenetic analyses have
provided some insight into the relationships of smaller
groups of the Poeciliinae (e.g., Rosen, 1967, 1979;
Rauchenberger, 1989; Rosa & Costa, 1993; Meyer et al.,
1994; Mojica et al., 1997; Rodriguez, 1997; Poeser, 2003) or
have dealt with higher taxa (Rosen, 1964; Parenti, 1981; Costa,
1996, 1998; Ghedotti, 2000). The only comprehensive study
is the classic revision of “Poeciliidae” by Rosen & Bailey
(1963), which did not deal with cladistic methodology. None-
theless, Rosen & Bailey (1963) is the basis for the current
internal classification of Poeciliinae. Later, Parenti &
Rauchenberger (1989) modified the classification of Rosen
& Bailey (1963) in order to accommodate it into the taxo-
nomic rank of subfamily proposed by Parenti (1981) (Table
1). Following Rosen & Bailey (1963) and Parenti &
Rauchenberger (1989), Tomeurus Eigenmann alone is the
supertribe Tomeurini. The remaining genera form the
supertribe Poeciliini, which is subdivided in the tribes
Poeciliini, Cnesterodontini, Gambusiini, Scolichthyini,
Girardinini, Heterandriini, and Xenodexini. Later, Ghedotti
(2000) proposed another classification for the Poeciliinae
(Table 2) based in his phylogenetic study of the Poecilioidea
despite the fact that only 12 genera were examined in his
cladistic analysis.
The history of the subfamily began in 1801 with the de-
scription of Poecilia vivipara Bloch & Schneider as new
genus and new species. Before establishment as a distinct
family, the history of the Poeciliidae is merged with that of
other cyprinodontiform families and with the Cyprinidae.
Table 1. Classification of Poeciliinae (Rosen & Bailey, 1963
and Parenti & Rauchenberger, 1989).
Subfamily Poeciliinae Bonaparte, 1831
Supertribe Tomeurini Eigenmann, 1912
Tomeurus
Supertribe Poeciliini Bonaparte, 1831
Tribe Poeciliini Garman, 1895
Poecilia, Xiphophorus, Alfaro, Priapella
Tribe Cnesterodontini Hubbs, 1924
Cnesterodon, Phalloceros, Phalloptychus, Phallotorynus.
Tribe Scolichthyini Rosen, 1967
Scolichthys
Tribe Gambusiini Gill, 1893
Gambusia, Belonesox, Brachyrhaphis
Tribe Girardinini Hubbs, 1924
Girardinus, Quintana, Carlhubbsia.
Tribe Heterandriini Hubbs, 1924
Heterandria, Poeciliopsis, Priapichthys, Neoheterandria,
Phallichthys
Tribe Xenodexini Hubbs, 1950
Xenodexia
Table 2. Classification of Poeciliinae (Ghedotti, 2000)
Subfamily Poeciliinae Bonaparte, 1831
Tribe Poeciliini Bonaparte, 1831
Poecilia, Xiphophorus, Phallichthys
Tribe Alfarini Hubbs, 1924
Alfaro
Tribe Cnesterodontini Hubbs, 1924
Cnesterodon, Phalloceros, Phalloptychus, Tomeurus,
Phallotorynus,
Tribe Gambusiini Gill, 1893
Gambusia, Belonesox, Brachyrhaphis
Tribe Girardinini Hubbs, 1924
Girardinus, Quintana, Carlhubbsia
Tribe Heterandriini Hubbs, 1924
Heterandria, Priapichthys, Poeciliopsis, Neoheterandria
Tribe Xenodexini Hubbs, 1950
Xenodexia
Tribe Scolichthyini Rosen, 1967
Scolichthys
Tribe Priapellini Ghedotti, 2000
Priapella
Rafinesque (1810) created the “Ordini Cyprinidi”; this
group contained a species of Mugil Linnaeus and three spe-
cies of Cyprinus Linnaeus. Later Wagner (1828) created the
family “Cyprinöidae” including the genera Lebias Goldfuss,
Poecilia Bloch & Schneider, Fundulus Lacepède, Cyprinodon
Lacepède, and Mollienesia Lesueur. The word “Cyprinöidae”
is etymologically identical to “Cyprinidi”, for they share the
same stem. Thus, Wagner failed to coin a new name. Cuvier
(1829) and Cuvier & Valenciennes (1846) followed Rafinesque
(1810) employing the name “Cyprinöides” and “Cyprins”.
Bonaparte (1831) also accepted the “Ordini Cyprinidi” of
Rafinesque (regarding it as a family), but divided it in three
groups: Cyprinini, Anableptini, and Poecilini [sic]. The
Poecilini was removed from the Cyprinidae by Bonaparte
(1840) to form a separate family Poecilidae [sic] (including
Anableptini and Poecilini). Four years before Bonaparte,
Agassiz (1834) had already separated the Cyprinodonts and
the Cyprinidae, removing Anableps Scopoli, Poecilia, Lebias,
Fundulus, Mollienesia, and Cyprinodon to a new family
“Cyprinodontes”. Swainson (1838) placed Poeciliinae as a
subfamily of Cobitidae. Gill (1857) adopted the name
Cyprinodontes and later (Gill, 1865; 1894) employed
Poeciliidae. Gill (1872) also used the name Cyprinodontidae
as a subgroup of the Haplomi, which also included the
Amblyopsidae, Esocidae, and Umbridae. Günther (1866) cre-
ated the genus Platypoecilus Günther and adopted
Cyprinodontidae as family name, dividing it in Cyprinodontidae
Limnophagae (=poeciliines) and Cyprinodontidae Carnivorae
(=remaining Cyprinodontiformes).
At the same time, new genera have been proposed:
Mollienesia, Xiphophorus Heckel and Heterandria
Agassiz. Poey (1854) added three new genera: Girardinus
Poey, Limia Poey, and Gambusia Poey. Belonesox Kner
was created in 1860. In the same year the genera
Hemixiphophorus Bleeker and Pseudoxiphophorus Bleeker
P. H. F. Lucinda & R. E. Reis
3
PROOFS
were erected. Soon after, De Filippi (1861) created Lebistes
De Filippi and Steindachner (1863) erected Poeciliodes
Steindachner.
Garman (1895) adopted the family name Cyprinodontes
and recognized eight subfamilies: Cyprinodontinae
(Cyprinodon, Tellia Gervais, Lebias, Characodon Günther,
Girardinichthys Bleeker, Neolebias Steindachner),
Jenynsiinae (Jenynsia Günther), Anablepinae (Anableps),
Haplochilinae [sic] (Orestias Valenciennes, Empetrichthys
Gilbert, Lucania Girard, Haplochilus [sic, = Aplocheilus
McClelland], Fundulus, Adinia Girard, Fundulichthys
Bleeker, Zygonectes Agassiz, Rivulus Poey, Cynolebias
Steindachner, Pterolebias, Garman, Haplochilichthys [sic, =
Aplocheilichthys Bleeker], Nothobranchius Peters,
Gambusiinae (Gambusia, Belonesox, Pseudoxiphophorus,
Heterandria) and Poeciliinae (Poecilia, Girardinus,
Platypoecilus, Mollienesia, Xiphophorus). Garman (1895)
also proposed two new genera (Glaridodon Garman and
Cnesterodon) in Poeciliinae.
Eigenmann (1903) created the genus Toxus Eigenmann.
Paragambusia Meek was proposed in 1904. Eigenmann (1907)
studied the intromittent organ in poeciliids of the La Plata
and found that it is formed by the third, fourth, and fifth anal-
fin rays. In that paper he erected the genera Acanthophacelus
Eigenmann Phalloceros and Phalloptychus. Soon after, Regan
(1908) erected Petalosoma Regan and Eigenmann (1909) cre-
ated Tomeurus.
Regan (1911) employed the name Microcyprini [=
Cyprinodontes of Garman] adding the Amblyopsidae (sub-
order Amblypsoidae) to this group. He allocated the remain-
ing species in the Poecilioidea, with one family Poeciliidae
divided in seven subfamilies: Cyprinodontinae, Fundulinae,
Orestiinae, Characodontinae, Jenynsiinae, Anablepinae, and
Poeciliinae. Regan merged the Gambusiinae and Poeciliinae
of Garman, recognizing the genera Acanthophacelus and
Petalosoma and adding Phalloceros and Phalloptychus.
Regan (1911) was the first to define the Poeciliinae by two
exclusive characters: absence of exoccipital condyles and male
anal-fin rays modified into a gonopodium.
Meek (1912) proposed the genus Alfaro Meek and treated
it as a Tomeurinae. Regan (1912) erected Petalurichthys Regan,
unneeded replacement name for Petalosoma preoccupied in
Coleoptera. Petalurichthys published November 1912 is an
objective synonym of Alfaro Meek September 12th, 1912.
Regan (1913) completed the first comprehensive revision
of the Poeciliinae proposing eight new genera: Priapichthys
Regan, Priapella Regan, Pseudopoecilia Regan, Poeciliopsis
Regan, Brachyrhaphis Regan, Leptorhaphis Regan,
Pamphorichthys Regan, and Pamphoria Regan. Langer (1913)
created Gulapinnus Langer, a junior objective synonym of
Cnesterodon. Regan (1914) proposed Heterophallus Regan.
Henn (1916) adopted the classification of Regan (1911)
and created new poeciliine genera Diphyacantha Henn,
Neoheterandria Henn, and Phallotorynus. Hubbs (1924) pro-
posed the genera Alloheterandria Hubbs, Allogambusia
Hubbs, Allopoecilia Hubbs, Darienichthys Hubbs,
Neopoecilia Hubbs, Phallichthys Hubbs, Panamichthys
Hubbs, Parapoecilia Hubbs, and Xenophallus Hubbs, and
recognized four subfamilies (Tomeurinae, Gambusiinae,
Poeciliinae, and Poeciliopsinae) and nine tribes within the
Poeciliidae (Poeciliinae sensu Parenti, 1981). Hubbs (1924)
considered the Gambusiinae as the least specialized poeciliids
whereas Tomeurus gracilis Eigenmann as the most special-
ized. Hubbs (1926) moved Alfaro from the Tomeurinae to the
Poeciliinae leaving Tomeurus as the sole genus in the
Tomeurinae. Hubbs (1926) also erected the genera Aulophallus
Hubbs, Micropoecilia Hubbs, Poecilistes Hubbs, and
Trigonophallus Hubbs. Subsequently, Hubbs proposed
Furcipenis Hubbs, Quintana Hubbs, and Allophallus Hubbs
(Hubbs, 1931; 1934; 1936, respectively).
Howell Rivero & Hubbs (1936) recognized Alfaro as dis-
tinct 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 & Gor-
don (1953).
From 1940 to 1963 eleven new genera have been proposed:
Arizonichthys Nichols, Allodontium Howell Rivero & Rivas,
Dactylophallus Howell Rivero & Rivas, Lembesseia Fowler,
Hubbsichthys Schultz, CurtipenisRivas & Myers, Xenodexia
Hubbs, Carlhubbsia Whitley, Recepoecilia Whitley,
Dicerophallus Alvarez, and Flexipenis Hubbs.
Rosen (1967) erected the poeciliid genus Scolichthys
Rosen. Rivas (1980) removed Limia from the synonym of
Poecilia and erected the subgenus Odontolimia Rivas, split-
ting Limia in two subgenera, L. (Limia) and L. (Odontolimia).
Poeser (2002) created the monotypic genus Pseudolimia
Poeser for Limia heterandria Regan.
Some taxonomic and phylogenetic studies have pro-
vided some progress into the relationships of smaller
groups of the Poeciliinae. The genera Heterandria and
Xiphophorus were reviewed by Rosen (1979), a classic
paper concerning methods of biogeographical analysis.
Rauchenberger (1989) put forward hypotheses of system-
atic and biogeographic relationships among the species of
the genus Gambusia. Rosa & Costa (1993) made a taxo-
nomic revision of the genus Cnesterodon, describing two
new species and proposing some putative synapomorphies
for Cnesterodon, in the absence of a phylogenetic analy-
sis. The phylogenetic relationships of Xiphophorus spe-
cies have been surveyed by Rosen (1979), Rauchenberger
et al. (1990), Meyer et al. (1994), Marcus & McCune (1999)
and Kallmann et al. (2004).
Mojica et al. (1997) proposed a hypothesis of relation-
ships among Brachyrhaphis species on the basis on mito-
chondrial DNA evidence. Rodriguez (1997) studied the rela-
tionships among genera of the tribe Poeciliini sensu Rosen &
Bailey, re-defining the tribe Poeciliini as comprehending the
genera Alfaro
, Priapella, Xiphophorus, Poecilia, Limia and
Pamphorichthys. Ptaceck & Breden (1998) proposed a mo-
lecular phylogeny for Poecilia, focusing on the species of
the subgenus Mollienesia. Breden et al. (1999) performed a
phylogenetic analysis for part of the species of the genus
Poecilia based on mitochondrial DNA evidence. Hamilton
Systematics of the subfamily Poeciliinae Bonaparte
4
PROOFS
(2001) proposed a phylogenetic hypothesis for Limia spe-
cies based on the mitochondrial genes sequences. Mateos et
al. (2002) proposed a historical biogeography hypothesis for
Poeciliopsis species based on sequence variation in two mi-
tochondrial genes. Poeser (2003) carried out a taxonomic re-
vision of Poecilia and proposed a phylogenetic hypothesis
for this genus.
Recent phylogenetic studies have dealt with higher taxa
but none has tackle the relationships among members of
the subfamily Poeciliinae as a whole. The only comprehen-
sive study is Rosen & Bailey (1963), but these authors
proposed a non-cladistic classification. In 1963, Rosen and
Bailey published their classic revision of poeciliid fishes.
This work separate the Poeciliidae into tree subfamilies,
Tomeurinae for Tomeurus, Xenodexiinae for Xenodexia,
and Poeciliinae for all other poeciliids including Alfaro,
which was recognized in the tribe Poeciliini. Parenti (1981)
recognized the Poeciliidae of Rosen & Bailey as a subfam-
ily (Poeciliinae) within a more inclusive family (Poeciliidae)
including the oviparous Aplocheilichthyinae and
Fluviphylacinae. Parenti & Rauchenberger (1989) modified
the classification of Rosen & Bailey (1963) to reflect the
change in taxonomic rank proposed by Parenti (1981) (Table
1). Meyer e Lydeard (1993) put forward a molecular phy-
logeny for Cyprinodontiformes including four poeciliine
genera. Costa (1996) presented new evidence concerning
the monophyly of the subfamilies of Poeciliidae, and pro-
posed a hypothesis of phylogenetic interrelationships
among them. Costa (1998) put forward a new phylogenetic
framework for the Cyprinodontiformes, differing from
Parenti’s (1981) hypothesis. Nonetheless, Parenti (1981)
and Costa (1996, 1998) did not tackle the inter- and
intrageneric relationships. Recently, Ghedotti (2000)
recognized the monophyly of the family Poeciliidae,
with three monophyletic subfamilies: (1) the
Aplocheilichthyinae containing solely Aplocheilichthys
spilauchen, (2) the Procatopodinae containing
Fluviphylax (Fluviphylacini), and the African lamp-eyed
killifishes (Procatopodini), and (3) the Poeciliinae. He also
resurrected the tribe Alfarini and proposed a new tribe,
the Priapellini for Priapella.
Cnesterodontini. The tribe Cnesterodontini as originally
erected by Hubbs (1924) was composed of genera
Phalloceros, Cnesterodon, Phallotorynus, and
Diphyacantha Henn. The Cnesterodontins were defined as
poeciliines bearing “terminal segment of ray 3 forming a
more or less specialized process” (Hubbs, 1924: 9). Hubbs
(1926) added Darienichthys to the Cnesterodontini. Later,
Rosen & Bailey (1963) recognized Diphyacantha and
Darienichthys as junior synonyms of Priapichthtys and
removed them from the Cnesterodontini, placing it in the
tribe Heterandriini. Rosen & Bailey (1963) also added
Phalloptychus to the Cnesterodontini. More recently,
Ghedotti (2000) based on his phylogenetic study of the
Poeciloidea recognized Tomeurus as a member of the tribe
Cnesterodontini and provided a diagnosis for the group
anchored in unique and unreversed synapomorphies. Thus,
as currently defined the Cnesterodontini comprises five gen-
era: Cnesterodon, Phalloceros, Phallotorynus,
Phalloptychus, and Tomeurus.
Cnesterodontines are disappointingly ill-studied from the
perspective of systematics. Except for Cnesterodon, this group
of fishes has received very little attention. The history of the
genus Cnesterodon began with Jenyns’ (1842) description of
Poecilia decemmaculata Jenyns, the first described species
currently placed in the genus. The genus Cnesterodon was
erected by Garman, with Poecilia decemmaculata
as type-
species, for it differed from the remaining genera so far as-
signed to Poeciliinae: Garman in the same paper also described
a second species for the genus: C. scalpridens Garman. A
third nominal species, C. carnegiei Haseman was described
from the rio Iguaçu drainage. Regan (1913) removed C.
scalpridens from Cnesterodon and erected the genus
Pamphoria [= Pamphorichthys] for this species. Rosa & Costa
(1993) recognized the validity of C. decemmaculatus and C.
carnegiei, and described C. brevirostratus Rosa & Costa from
the upper rio Uruguay and rio Jacuí drainages, and C.
septentrionalis Rosa & Costa from the rio Araguaia drainage.
Rosa & Costa (1993) also reported nine putative
synapomorphies for the genus. Latter, Lucinda & Garavello
(2001) described C. hypselurus Lucinda & Garavello from the
rio Paranapanema basin, and C. omorgmatos Lucinda &
Garavello, a second species from the rio Iguaçu basin.
Cnesterodon raddai Meyer & Etzel was described from the
lower portions of rio Paraná.
Studies concerning Phallotorynus species are notably
scarce, and are mostly confined to original descriptions. Henn
(1916) erected the genus Phallotorynus, based on gonopodium
structure, for his new species, P. fasciolatus Henn, from the rio
Paraíba do Sul. Later, Ihering (1930) described P. jucundus Ihering
from a tributary of rio Mogi-Guaçu in the upper rio Paraná drain-
age. Rosen & Bailey (1963) redefined the genus by osteological
characters on the basis of specimens of P. fasciolatus and speci-
mens from the neighborhood of Asunción, expanding the distri-
bution range for P. jucundus to the Paraguay drainage. Finally,
Oliveros (1983) described P. victoriae Oliveros from the lower
portions of rio Paraná basin in Argentina.
Papers concerning Phalloptychus are also extremely rare
in systematic literature, being confined to original descrip-
tions. The history of Phalloptychus began with the first de-
scribed species currently in the genus: Girardinus januarius
Hensel. A second species, G. iheringii Boulenger was de-
scribed from Rio Grande do Sul. Eigenmann (1907) created
the genus Phalloptychus for Girardinus januarius. Henn
(1916) described a third species, P. eigenmanni, from the rio
Catu at Alagoinhas, Bahia.
Phalloceros and Tomeurus are monotypic genera and dis-
appointingly ill-studied from the perspective of systematics.
Both genera were erected by Eigenmann (1907 and 1909, re-
spectively) for Girardinus caudimaculatus Hensel and
Tomeurus gracilis, respectively.
P. H. F. Lucinda & R. E. Reis
5
PROOFS
Material and Methods
Material examined is listed in the Appendix I. Museum
acronyms are from Leviton et al. (1985), and Leviton & Gibbs
(1988) except for MNHCI: Museu de História Natural do Capão
da Imbúia, Curitiba; UFPB: Universidade Federal da Paraíba,
Departamento de Sistemática e Ecologia, João Pessoa; and
ZVC-P: Sección Vertebrados, Facultad de Ciencias,
Montevideo.
Clearing and staining followed the method of Taylor &
Van Dyke (1985). Anatomical illustrations were prepared from
sketches of structures from cleared and stained specimens as
viewed through a camera lucida mounted on a dissecting ste-
reomicroscope. External characters, e.g. color pattern, were
also examined.
The current definition of Poeciliinae is Parenti’s (1981)
definition, which corresponds to the Rosen & Bailey’s (1963)
family Poeciliidae. Number and disposition of cephalic pores
follow the nomenclature of Rosen & Mendelson (1960),
Gosline (1949) and Parenti (1981). Only adult individuals have
been examined to avoid undesired ontogenetic variation.
Nomenclature of the gonopodium followed Rosen & Gordon
(1953). Anal-fin radial terminology followed Rosen & Kallman
(1959). Descriptions of gonopodium morphology are based
on fully developed gonopodia of large adult males. Anatomi-
cal nomenclature, other than gonopodial, follows Rosen &
Bailey (1963), Parenti (1981), and Rauchenberger (1989).
Proposed hypotheses of phylogenetic relationships
among studied taxa followed the phylogenetic method for-
mally put forward by Hennig (1966). The ingroup included
representatives of all poeciliine genera, and all species of
the tribe Cnesterodontini sensu Rosen & Bailey (1963). The
data matrix of 71 taxa and 144 characters (Appendix II) in-
cludes 24 new species of Cnesterodontini, whose descrip-
tions will be provided by Lucinda (in prep.) and Lucinda et
al. (in prep.). Intrageneric relationships and synapomorphy
lists for subclades of Phalloptychus, Cnesterodon,
Phalloceros, and Phallotorynus are provided herein, but
are discussed in Lucinda (in prep.) and Lucinda et al. (in
prep.). Question marks were used to indicate when a charac-
ter state could not be checked due to lacking of available
specimens. Dashes were employed for both inapplicable
coding and polymorphisms. The phylogenetic analysis
aimed to test the monophyly of the subfamily Poeciliinae as
well and its tribes (sensu Rosen & Bailey, 1963). Fundulus
heteroclitus (Linnaeus), Cyprinodon macularius Baird &
Girard, Jenynsia unitaenia Ghedotti & Weitzman,
Aplocheilichthys spilauchen (Duméril), Fluviphylax
pygmaeus (Myers & Carvalho), and Procatopus gracilis
Clausen were included as outgroup taxa. Phylogenetic analy-
ses included all 71 taxa simultaneously and were performed
with Hennig86 (Farris, 1988) coupled with Tree Gardener
(Ramos, 1997). Examination of more specimens suggested
that there were problems with homology concerning some
characters of Ghedotti’s (2000) analysis (e.g. fusion of the
dorsal-most proximal pectoral radial to the scapula) or these
characters states could not be confirmed on existing speci-
mens. Therefore these characters were excluded from the
analysis.
All transformation series were considered unordered.
Maximum parsimony analyses were undertaken using the mh*;
bb* algorithm of Hennig86. Character optimization followed
accelerated transformation model (ACCTRAN) for it is more
consistent with the concepts of homology and synapomorphy
(de Pinna, 1991). The numbers on the branches of the strict
consensus tree (Fig. 1, 2, and 3) corresponds to tree nodes
and to clade number. In the diagnoses and synapomorphy
list uniquely derived and unreversed features are indicated
by two asterisks (e.g. 47-2**); uniquely derived features are
indicated by one asterisk (e.g. 24-1*). An asterisk indicates
uniquely derived autapomorphies. Transformation series
analysis (TSA) is presented in Appendix III. Fits of individual
characters are shown in Appendix IV.
Results
Character Description and Analysis
Neurocranium
Character 0 - Mesethmoid: (0) ossified; (1) cartilaginous.
Most cyprinodontoids and other atherinomorphs possess
a bony mesethmoid (state 0). Parenti (1981) discussed carti-
laginous mesethmoid (state 1) as diagnostic of the subfamily
Aplocheilichthyinae, including Aplocheilichthys spilauchen.
However, Ghedotti (2000) verified that A. spilauchen have an
ossified mesethmoid, which has been confirmed in this study.
Costa (1998) reported a cartilaginous mesethmoid for Lebias
(Cyprinodontidae) and for most procatopodines. Ghedotti
(2000) reported a cartilaginous mesethmoid as diagnostic for
the tribe Procatopodini. Priapella is unique among
poeciliines by the possession of a cartilaginous mesethmoid.
Character 1 - Anterior margin of frontals (Ghedotti, 2000: fig.
3): (0) extending anteriorly between nasals; (1) straight or
slightly cleft medially.
The anterior margin of the frontals extends anteriorly by
between nasals (state 0) in most Poecilioidea. Among mate-
rial examined, Fluviphylax, Procatopus, Alfaro,Priapichthys,
Priapella, and Belonesox possess the anterior margin of
frontals straight or slightly cleft medially (state 1).
Character 2 - Parietals (Ghedotti, 2000: fig. 3): (0) large reach-
ing sphenotic anteriorly; (1) short restrict to epiotic region,
not reaching sphenotic anteriorly; (2) absent.
Parietals are present and large reaching sphenotic anteri-
orly (state 0) in atherinomorphs except for beloniforms and
some cyprinodontoids (Parenti, 1981; Dyer & Chernoff, 1996).
Among the studied taxa, the following possess large pari-
etals reaching sphenotic anteriorly: Jenynsia, Fundulus,
Cyprinodon, Alfaro, Brachyrhaphis, Priapichthys,
Priapella, Heterandria, Gambusia, Belonesox,
Neoheterandria, Scolichthys, Girardinus, and Xiphophorus.
Systematics of the subfamily Poeciliinae Bonaparte
6
PROOFS
According to Ghedotti (2000) parietals are present but
reduced in size (state 1) in Hypsopanchax Myers, Tomeurus,
Phalloceros, Poecilia, Girardinus, and Phallichthys.
Parenti (1981) reported the absence of parietals (state 2)
as synapomorphic for all cyprinodontines and for
Fluviphylax, Pantanodon Myers, and the procatopodines.
Parietals are also independently lacking in some groups of
poeciliines. Ghedotti (2000) recognized the lacking of pari-
etals in Aplocheilichthys, Cnesterodon, Poeciliopsis,
procatopodines (except Hypsopanchax) and in some indi-
viduals of Phallotorynus victoriae (polymorphic condition).
Additionally, this condition was found to occur also in
Pseudopoecilia, Phalloptychus, Xenodexia,
Pamphorichthys, Micropoecilia, “Poecilia” reticulata Pe-
ters and Phallotorynus.
Parietals vary considerably among poeciliines
(Rauchenberger, 1989; Figueiredo, 1997). Following our hy-
pothesis, reduction and lacking of parietals appeared inde-
pendently several times. Among poeciliines parietal reduc-
tion appeared independently four times: in Tomeurus [Clade
63]; Phallichthys + Xenophallus + Poeciliopsis +
Phalloptychus [Clade 113]; Quintana + Carlhubbsia [Clade
109]; Poecilia + Limia [Clade 93]. The loss of parietals oc-
curred twice: (1) in Phalloptychus and (2) in the Poeciliini +
Cnesterodontini [Clade 115] with reversals to [2-1] in Quintana
+ Carlhubbsia [Clade 109], Poecilia + Limia [Clade 93], and
in Phalloceros. A reversal to condition [2-0] occurs in
Xiphophorus.
Character 3 - Epiotic process (Ghedotti, 2000: fig. 3): (0) long
extending beyond first pleural rib; (1) long, longer than
exoccipital process but not reaching first pleural rib; (2) short,
shorter than exoccipital process; (3) absent.
Long epiotic process extending beyond first pleural rib
is present in Fundulus, Jenynsia, and
Aplocheilichthys and
is hypothesized as plesiomorphic (state 0). Among
poeciliines, enlarged epiotic processes are present in Alfaro,
Phallichthys, Xenophallus, Poeciliopsis, Quintana, and
Carlhubbsia.
Parenti (1981) hypothesized enlarged epiotic processes
as a synapomorphy for anablepids. However, Ghedotti (1998)
did not recognize expanded epiotic processes as uniquely
synapomorphic of the Anablepidae and recorded their pres-
ence in Anableps, Oxyzygonectes Fowler, three species of
Jenynsia, and Aplocheilichthys spilauchen. Ghedotti (2000)
also observed the presence of long epiotic processes in Fun-
dulus chrysotus Günther as well as in some poeciliines.
Girardinus and Poecilia possess an epiotic process
longer than exoccipital process but not reaching first pleural
rib (state 1). This condition is also observed in Brachyrhaphis,
Priapichthys, Neoheterandria, Scolichthys, Girardinus,
Xiphophorus, Poecilia, and Limia.
Fig. 1. Strict consensus tree of 96 most equally parsimonious trees representing a hypothesis of intergeneric relationships of
Poeciliinae. Intrageneric relationships of Cnesterodon, Phallotorynus, Phalloceros, and Phalloptychus are not depicted.
Character state transformations are based on ACCTRAN optimization. The numbers on the branches refer to the character
state transformation series listed in the Appendix II.
P. H. F. Lucinda & R. E. Reis
7
PROOFS
Fig. 2. (a) Hypothesis of relationships among the species of
Phalloptychus; (b) Hypothesis of relationships of among the
species of Cnesterodon; (c) Hypothesis of relationships of
among the species of Phallotorynus. Cladograms a, b, and c
are a continuation of that shown in Fig. 1.
Epiotic process is shorter than exoccipital process (state
2) in Heterandria, Pseudopoecilia, Xenodexia, and
“Poecilia” reticulata.
Ghedotti (2000) reported the absence of epiotic processes
in procatopodines, Tomeurus, Cnesterodon, Phallotorynus,
Cubanichthys Hubbs, Cyprinodon, Valencia Myers, and
Crenichthys Hubbs, which is corroborated by this study. The
absence of epiotic processes was also observed in Priapella,
Phalloptychus, Pamphorichthys, and Micropoecilia.
Following the hypothesis presented here state [3-1] ap-
peared once among poeciliines: in the ancestor of members
of Clade 125. Condition [3-2] appeared independently four
times in Heterandria, Pseudopoecilia, Xenodexia, and
“Poecilia” reticulata, whereas the loss of epiotic processes
occurred independently in Tomeurus, Priapella,
Phalloptychus, in the ancestor of Clade 92, and in
cnesterodontines.
Character 4 - Halves of supraoccipital process (Fig. 4a, b, c): (0)
simple; (1) bifid, outer half minute; (2) bifid, outer half larger.
In most cyprinodontiforms halves of supraoccipital pro-
cess are simple (state 0; Fig. 4a). Girardinus and
Pamphorichthys possess bifid halves of supraoccipital pro-
cess with a minute external half (state 1; Fig. 4b), which is
interpreted as independently acquired. Cnesterodontines
possess bifid halves of supraoccipital process with a large
external half (state 2; Fig. 4c). A reversal to state 0 occurs in
Phallotorynus fasciolatus.
Cephalic sensory system
Nomenclature follows Gosline (1949) and Rosen &
Mendelson (1960). We refer the reader to figures depicted in
Gosline (1949) and Rosen & Mendelson (1960) and in the
articles cited below for a detailed comprehension of charac-
ters 5 to 10.
Character 5 - Posterior supraorbital canal (pores 2b, 3, 4a): (0)
absent or opened, forming a shallow groove (Rosen &
Mendelson, 1960: fig. 3B, J-M); (1) opened, forming a sinu-
ous depression over the frontal (supraorbital bone) (Rosen,
1952: fig. 7); (2) closed (Gosline, 1949: plate II, fig. 1, 4; Rosen
& Mendelson, 1961: figs. 3A, C-D, F-I).
Posterior supraorbital canal varies among outgroup mem-
bers and among members of the ingroup. In Aplocheilichthys,
Fluviphylax, and poeciliines, except Alfaro, Brachyrhaphis,
Priapichthys, Priapella, Girardinus, Carlhubbsia, Poecilia,
and Limia, posterior supraorbital canal is lacking or is opened,
forming a shallow groove (state 0). Posterior supraorbital ca-
nal form a sinuous depression over the frontal (state 1) in
Procatopus, Alfaro, Brachyrhaphis, Priapichthys, and
Priapella. Posterior supraorbital canal is closed (state 2) in
Jenynsia, Cyprinodon, Fundulus, Girardinus, Carlhubbsia,
Poecilia, and Limia.
Following the hypothesis proposed here, a sinuous de-
pression over the frontal formed by posterior supraorbital
canal (state 1) is synapomorphic for poeciliines except
Tomeurus [clade 126] and was independently acquired by
Procatopus. State 0 is interpreted as a synapomorphic rever-
sal for Heterandriini + Gambusiini + Supertribe Poeciliini
[Clade 122]; whereas a closed posterior supraorbital canal
(state 2) appeared independently three times in Girardinus,
Carlhubbsia, and in Poecilia + Limia [clade 93].
Character 6 - Anterior section of posterior remnant of infraorbital
system (pores 4b, 5, 6a): (0) absent or opened, forming a shallow
groove (Rosen & Mendelson, 1960: fig. 3A-E, G-P); (1) opened,
pores confluent forming a major sinuous depression above and
slightly behind the orbit (Rosen, 1952: fig. 7); (2) closed (Gosline,
1949: plate II, fig. 1, 4; Rosen & Mendelson, 1960: fig. 3F).
Posterior supraorbital canal varies among members of the
outgroup and among members of the ingroup. In
Aplocheilichthys, Fluviphylax, and poeciliines, except Alfaro,
Brachyrhaphis, Priapichthys, and Priapella, anterior sec-
tion of posterior remnant of infraorbital system is lacking or is
opened, forming a shallow groove (state 0). Anterior section
of posterior remnant of infraorbital system forms a major sinu-
ous depression above and slightly behind the orbit (state 1)
in Procatopus, Alfaro, Brachyrhaphis, Priapichthys, and
Priapella. Anterior section of posterior remnant of infraor-
bital system is closed (state 2) in Jenynsia, Cyprinodon, and
Fundulus.
Systematics of the subfamily Poeciliinae Bonaparte
8
PROOFS
Following the hypothesis proposed here, a major sinuous
depression above and slightly behind the orbit formed by
anterior section of posterior remnant of infraorbital system
(state 1) is synapomorphic for poeciliines except Tomeurus
[clade 126] and was independently acquired by Procatopus.
State 0 is interpreted as a synapomorphic reversal for
Heterandriini + Gambusiini + Supertribe Poeciliini [Clade 122].
Within supertribe Poeciliini a change to state 1 occurs in
Carlhubbsia.
Character 7 - Posterior section of posterior remnant of in-
fraorbital system (canal 6b, 7): (0) closed (Gosline, 1949: plate
II, fig. 4; Parenti, 1981: fig. 14A; Ghedotti & Weitzman, 1995:
fig. 2); (1) opened into a groove (Rosen & Mendelson, 1960:
fig. 2A, B).
Most cyprinodontiforms possess a closed posterior sec-
tion of posterior remnant of infraorbital system (state 0). Pos-
terior section of posterior remnant of infraorbital system
opened into a groove (state 1) is herein hypothesized as
apomorphic and to have been independently acquired by
Tomeurus, Gambusia + Belonesox [clade 118], Scolichthys,
Phalloptychus, Pamphorichthys, and cnesterodontines (with
a reversal in Phallotorynus).
Character 8 - Preopercular canal: (0) present and entirely
closed, except for seven pores (8, 9,10, 11,12, U, V) (Gosline,
1949: plate II, fig. 2); (1) absent or opened in a shallow groove
(Rosen & Mendelson, 1960: fig. 2A, B); (2) present and en-
tirely closed, except for four pores (8, 11, 12, V); (3) present
and partially closed, only canal between pores U - V closed;
(4) present and partially closed, only canals between pores
11-12, and 12-U closed (sometimes canal U-V also closed);
(5) present and partially closed: canal 8, 9, 10 opened in a
deep groove; pore 11 opened and elongate (sometimes
confluent with groove 8,9,10) canal between pores 12, U - V
closed (sometimes canal U-V opened); (6) restrict to closed
canal 10-11.
All studied outgroup taxa (except Fluviphylax) possess
a closed preopercular canal, except for seven pores (8, 9,10,
11,12, U, V) (state 0). In Belonesox, Scolichthys,
Pamphorichthys, and Cnesterodon preopercular canal is lack-
ing or opened in a shallow groove (state 1). Phallotorynus
species share a closed preopercular, except for four pores (8,
11, 12, V) (state 2). Gambusia presents a partially closed
preopercular canal, with canal only between pores U - V closed
(state 3). In Tomeurus, preopercular canal is present and par-
tially closed, with canals only between pores 11-12, and 12-U
closed (sometimes canal U-V also closed) (state 4). In
Phalloceros, the preopercular canal is partially closed: canal
between pores 8, 9, 10 opened in a deep groove; pore 11
opened and elongate (sometimes confluent with groove 8, 9,
10); canal between pores 12, U - V closed (state 5). Finally,
preopercular canal is restricted to closed canal 10-11 in
Fluviphylax (state 6).
Following our hypothetical history of poeciliines, state 1
appeared independently in Belonesox, Scolichthys,
Pamphorichthys, and Cnesterodon, whereas states 2, 3, 4, 5,
and 6 are interpreted as synapomorphic for Phallotorynus, Gam-
busia, Tomeurus Phalloceros, and Fluviphylax, respectively.
Fig. 3. Hypothesis of relationships of among the species of Phalloceros. Cladogram is a continuation of that shown in Fig. 1.
P. H. F. Lucinda & R. E. Reis
9
PROOFS
Fig. 4. Dorsal view of posterior crania. (a) Heterandria jonesii,
UMMZ 210999; (b) Girardinus uninotatus, AMNH 96301; (c)
Phalloceros n. sp. G, MCP 30509. DP = dermosphenotic; EO =
epiotic; F = Frontal; P = parietal; PO = pterotic; SO = supraoccipital;
SOP = supraoccipital process; SP = sphenotic. Scale bar 1mm.
Character 9 - Preorbital canal: (0) present and entirely closed,
bearing four pores (Rosen & Mendelson, 1960: fig. 2C, D); (1)
present and partially closed bearing two upper pores and a
lower deep groove; (2) absent or opened, forming a very shal-
low groove (Rosen & Mendelson, 1960: fig. 2C, D); (3) present
and entirely closed, bearing three pores.
Among studied taxa, preorbital canal is entirely closed; bear-
ing four pores (state 0) in Aplocheilichthys, Procatopus,
Jenynsia, Fundulus, Alfaro, Priapella, Girardinus,
Xenophallus, Xenodexia, Poecilia, and Limia. Preorbital canal
is partially closed bearing two upper pores and a lower deep
groove (state 1) in Brachyrhaphis, Priapichthys, Heterandria,
Phallichthys, Poeciliopsis, Quintana, Carlhubbsia,
Xiphophorus, “Poecilia”, and Micropoecilia. In Tomeurus,
gambusiines, Phalloptychus, Pamphorichthys and
cnesterodontines preorbital canal is absent or opened, forming
a very shallow groove (state 2). Cyprinodon presents preorbital
canal closed, bearing three pores. State 1 is hypothesized to has
independently appeared twice: (1) as a synapomorphy for
poeciliines except Tomeurus andAlfaro [Clade 125] (with several
subsequent reversals); (2) as a synapomorphy for Micropoecilia
+ “Poecilia” [Clade 87]. Additionally, state 2 is interpreted as
independently evolved in Fluviphylax, Tomeurus, gambusiines,
Phalloptychus, Pamphorichthys, and cnesterodontines.
Character 10 - Mandibular canal (W, X, Y, Z): (0) present and en-
tirely closed, bearing four pores (Gosline, 1949: plate II, fig. 3, 5, 6;
Ghedotti & Weitzman, 1995: fig. 3); (1) absent or opened, forming
a very shallow groove; (2) present and entirely closed, bearing
five pores; (3) present and partially closed bearing six pores; (4)
present and partially closed; canal connecting pores X and W
closed; canal connecting pores Y and Z absent; (5) present and
entirely closed, bearing three pores, pore Z absent; (6) opened,
formed by two deep grooves.
Among studied taxa, mandibular canal is entirely closed; bear-
ing four pores (state 0) in Aplocheilichthys,
Jenynsia,Cyprinodon,
and Fundulus. Mandibular canal is absent or opened, forming a
very shallow groove (state 1) in Fluviphylax,Tomeurus, and mem-
bers of the supertribe Poeciliini with the exception of Girardinus,
Xenodexia, and Poecilia. This condition is hypothesized to have
been independently acquired in these groups. Procatopus and
Priapella possess a mandibular canal entirely closed, bearing
five pores (state 2) and the presence of this feature in these taxa is
interpreted as homoplastic. Mandibular canal is partially closed
bearing six pores (state 3) in Priapichthys. Mandibular canal is
present and partially closed; canal connecting pores X and W
closed; canal connecting pores Y and Z absent (state 4) in Gam-
busia. Mandibular canal is entirely closed bearing three pores,
pore Z absent (state 5) in Girardinus. Mandibular canal is opened,
formed by two deep grooves in Xenodexia (state 6). States 3, 4, 5,
and 6 are interpreted as autapomorphic for Priapichthys,Gambu-
sia, Girardinus, and Xenodexia, respectively.
Suspensorium and mandibular arch
Character 11 - Medial surface of ascending process of pre-
maxilla (Ghedotti, 2000: fig. 3): (0) approximately straight; (1)
slightly angled laterally; (2) angled laterally at proximal end,
forming a triangle space between proximal ends of ascending
processes.
Systematics of the subfamily Poeciliinae Bonaparte
10
PROOFS
Fig. 5. Dorsal view of left premaxilla. (a) Heterandria
jonesii, UMMZ 210999; (b) Priapichthys annectens, ANSP
163139; (c) Phalloptychus iheringii, MCP 11054; (d)
Phalloceros n. sp. G, MCP 30509; (e) Girardinus
uninotatus, AMNH 96301; (f) Cnesterodon n. sp. B, MCP
19784. Scale bar 1 mm.
Ghedotti (2000) reported a laterally angled medial surface
of the ascending processes as independently acquired by
Micropanchax and the common ancestor of a clade com-
posed of Cnesterodon, Phalloceros, Phallotorynus,
Girardinus, Poecilia, Phallichthys, and Tomeurus with a re-
versal in Tomeurus. Ghedotti (2000) recognized only two char-
acter states whereas this study recognizes a third intermedi-
ate state (state 1).
Medial surface of ascending process of premaxilla is ap-
proximately straight (state 0) in all studied outgroup taxa
and Scolichthys, Neoheterandria, Pseudopoecilia,
Cnesterodon n. sp. A, C. hypselurus, C. brevirostratus, C.
septentrionalis, and Phallotorynus fasciolatus. Alfaro,
Brachyrhaphis, Priapichthys, Priapella, and Heterandria
possess a slightly laterally angled medial surface (state 1).
Medial surface of ascending process of premaxilla is later-
ally angled at proximal end, forming a triangular space be-
tween proximal ends of ascending processes (state 2) in the
remaining studied taxa.
Our phylogenetic hypothesis indicates that a slightly lat-
erally angled medial surface of ascending process of premax-
illa (state 1) is uniquely derived for a clade embracing all
poeciliines but Tomeurus [Clade 126]. State 2 appears to be
synapomorphic for Tribe Gambusiini + Supertribe Poeciliini
[Clade 121], and it shows three subsequent reversals to state
0: (1) in the common and exclusive ancestor of Scolichthys,
Neoheterandria, and Pseudopoecilia [Clade 117]; (2) in the
common and exclusive ancestor of Cnesterodon n. sp. A, C.
hypselurus, C. brevirostratus
, C. septentrionalis [Clade 103],
and in Phallotorynus fasciolatus.
Character 12 - Shape of the ascending process of premaxilla
(Fig. 5a-f): (0) Elongate, distal tip rounded; (1) elongate, distal
tip pointed; (2) short and pointed; (3) short and truncate; (4)
short, distal tip rounded; (5) minute, almost absent.
Ascending process of premaxilla is elongate with a round
distal tip (state 0; Fig. 5a) in Aplocheilichthys, Jenynsia,
Fluviphylax, Procatopus, Tomeurus, Alfaro, Brachyrhaphis,
Priapella, Heterandria, and Pseudopoecilia. Cyprinodon,
Fundulus, Priapichthys, Gambusia, Neoheterandria, and
Scolichthys possess an elongate ascending process of pre-
maxilla, with a pointed distal tip (state 1; Fig. 5b). Remaining
poeciliines except Girardinus, Quintana, Phalloceros, and
Cnesterodon n. sp. B have a short and pointed ascending
process of premaxilla (state 2; Fig. 5c). Quintana and
Phalloceros have a short and truncate ascending process of
premaxilla (state 3; Fig. 5d). Girardinus presents the ascend-
ing process of premaxilla short with distal tip round (state 4;
Fig. 5e), whereas Cnesterodon n. sp. B has a minute ascend-
ing process of premaxilla (state 5; Fig. 5f).
An elongate ascending process of premaxilla, with a
pointed distal tip (state 1; Fig. 5b) is hypothesized as
synapomorphic and independently acquired in the
Priapichthyini and the Gambusiini (with a reversal to state 0
in Pseudopoecilia). A short and pointed ascending process
of premaxilla (state 2; Fig. 5c) is interpreted as a uniquely
derived and unreversed synapomorphy for the supertribe
Poeciliini [Clade 119], with subsequent transformations to
states 3 (Quintana and Phalloceros), 4 (Girardinus), and 5
(Cnesterodon n. sp. B).
Character 13 - Contact area between premaxillae: (0) not el-
evate; (1) elevate.
The contact area between premaxillae is plain in almost all
cyprinodontoids (state 0). An elevated contact area between
premaxillae (state 1) is hypothesized as a synapomorphy for
the members of the supertribe Poeciliini, with a reversal to the
plesiomorphic condition in the Phallotorynus + Phalloceros
[Clade 106].
Character 14 - Anterior border of ventral maxilla (Fig. 6a, B):
(0) straight; (1) concave.
The anterior border of ventral maxilla is straight (state 0;
Fig. 6a) in outgroup taxa, as well as in Priapella, Gambusia,
Belonesox
, Neoheterandria, and Scolichthys and all mem-
bers of the supertribe Poeciliini with the exception of
Girardinus. Anterior border of ventral maxilla is concave (state
1; Fig. 6b) in Alfaro, Brachyrhaphis, Priapichthys,
P. H. F. Lucinda & R. E. Reis
11
PROOFS
Heterandria, Pseudopoecilia, and Girardinus and is inter-
preted as synapomorphic for Clade 126, which comprises all
poeciliines but Tomeurus. A reversal to state 0 occurs in
Priapellini and in the common and exclusive ancestor of mem-
bers of Clade 121 [Gambusiini + Supertribe Poeciliini]. Within
this clade a transformation 0 to 1 appears in Pseudopoecilia
and Girardinus.
Character 15 - Ventral surface of dentary (Costa, 1991: fig. 4G,
5E and Ghedotti, 2000: fig. 5): (0) straight or bearing a tiny
straight process; (1) bearing a curved and forward directed
process.
Most cyprinodontiforms possess the ventral surface of
dentary straight or bearing a tiny straight process (state 0).
Costa (1991) suggested the presence of a curved and forward
directed process on ventral surface of dentary (state 1) as a
putative synapomorphy for a group embracing
Pamphorichthys, Poecilia, Limia, Xiphophorus, Cnesterodon,
Phalloceros, Phallotorynus, Phalloptychus, Priapichthys,
Poeciliopsis, Priapella, Quintana, Carlhubbsia, Xenodexia,
and Phallichthys. This is partially corroborated by our results.
The current phylogenetic analysis supports this feature as a
uniquely derived and unreversed synapomorphy for the
supertribe Poeciliini [Clade 119]. In addition to the genera above
(except Priapichthys and Priapella), this group comprises
Girardinus, Xenophallus, and Micropoecilia.
Character 16 - Notch on dentary (Rosen & Bailey, 1963: fig. 21
C, F): (0) absent; (1) present.
Most poeciliids lacks a notch on dentary (state 0). A notch
on dentary is present in Phallichthys, Poeciliopsis,
Carlhubbsia, Xiphophorus, Xenodexia, Poecilia, Limia, and
“Poecilia” reticulata. This feature is interpreted as
apomorphic (state 1) and independently acquired by
Phallichthys, Poeciliopsis and by the common and exclu-
sive ancestor of members of the tribe Poeciliini, with a rever-
sal in Pamphorichthys + Micropoecilia + “Poecilia” (Clade
92). A return to state 1 occurs in “Poecilia” reticulata.
Character 17 - Anterior cleft of anguloarticular (Ghedotti, 2000:
fig. 5): (0) small, not extending beyond posterior border of
Meckel‘s cartilage; (1) large, extending beyond posterior bor-
der of Meckel´s cartilage; (2) absent.
Anterior cleft of anguloarticular is small, not extending
beyond posterior border of Meckel’s cartilage (state 0) in
Aplocheilichthys, Procatopus, Jenynsia, and in all poeciliines
except Phalloceros. In Cyprinodon and Fundulus the ante-
rior cleft of anguloarticular is large, extending beyond poste-
rior border of Meckel’s cartilage (state 1). Anterior cleft of
anguloarticular is absent (state 2) and interpreted as
synapomorphic for a clade embracing Phalloceros n. sp. S +
Phalloceros n. sp. T + Phalloceros n. sp. V + Phalloceros n.
sp. B + Phalloceros n. sp. N + Phalloceros n. sp. R +
Phalloceros n. sp. I + Phalloceros n. sp. O + Phalloceros n.
sp. P + Phalloceros n. sp. M + Phalloceros n. sp. H +
Phalloceros n. sp. Q + Phalloceros n. sp. J + Phalloceros n.
sp. L [Clade 80], with a reversal to state 0 in (Phalloceros n.
sp. P + Phalloceros n. sp. M) Clade [71].
Character 18 - Ventral process of anguloarticular (Ghedotti, 2000:
fig. 5): (0) long, extending anterior to where anguloarticular
overlaps dentary; (1) short, not extending anterior to where
anguloarticular overlaps dentary; (2) absent.
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 co-occurrence of a long retroarticular and a
long ventral process of the anguloarticular as synapomorphic
of Poecilioidea. Ghedotti (2000) treated the ventral process of
the anguloarticular and the retroarticular as separate trans-
formation series because they vary independently. Ghedotti
(2000) reported a long ventral process of the anguloarticular
in all poecilioids examined by him.
Among studied taxa, Aplocheilichthys, Jenynsia,
Procatopus and almost all poeciliines the ventral process of
anguloarticular is long, extending anterior to where
anguloarticular overlaps dentary (state 0).
This process is short, not extending anterior to where
anguloarticular overlaps dentary (state 1) in Cyprinodon,
Fundulus, Poeciliopsis, and Phalloptychus. State 1 is herein
interpreted as synapomorphic for the clade [Poeciliopsis +
Phalloptychus]. Its presence in Fundulus and Cyprinodon
is considered homoplastic.
Ventral process of anguloarticular is absent (state 2) and
interpreted as synapomorphic for the clade [Phalloceros n. sp.
S + Phalloceros n. sp. T + Phalloceros n. sp. V + Phalloceros
n. sp. B + Phalloceros n. sp. N + Phalloceros
n. sp. R +
Phalloceros n. sp. I + Phalloceros n. sp. O + Phalloceros n.
sp. P + Phalloceros n. sp. M + Phalloceros n. sp. H +
Phalloceros n. sp. Q + Phalloceros n. sp. J + Phalloceros n.
sp. L] [Clade 80], with a reversal to state 0 in the ancestor of
Phalloceros n. sp. P and Phalloceros n. sp. M [Clade 71].
Character 19 - Ventral invagination on anguloarticular (Fig.
7): (0) absent; (1) present.
Among studied taxa, Phalloceros n. sp. B, Phalloceros n.
sp. C, and Phalloceros n. sp. V possesses a ventral invagina-
tion on anguloarticular, which is interpreted as synapomorphic
for Phalloceros n. sp. C + Phalloceros n. sp. V and indepen-
dently acquired in Phalloceros n. sp. B.
Character 20 - Ascending process of parasphenoids in adults
(Ghedotti, 2000: fig. 3): (0) long, contacting pterosphenoids;
(1) short, not reaching pterosphenoids; (2) absent.
Among the studied taxa, in Aplocheilichthys, Procatopus,
Jenynsia, Fundulus, Tomeurus, Alfaro, Brachyrhaphis,
Priapella, Priapichthys, Heterandria, Belonesox,
Neoheterandria, Phallichthys, Xenophallus, Poeciliopsis,
Phalloptychus, Quintana, Carlhubbsia, Xiphophorus,
Poecilia, Limia, Pamphorichthys hollandi Henn,
Micropoecilia, “Poecilia”, Phallotorynus n. sp. A,
Phallotorynus n. sp. B, and Phalloceros, the ascending pro-
cess of parasphenoids in adults is long, contacting
Systematics of the subfamily Poeciliinae Bonaparte
12
PROOFS
pterosphenoids (state 0). In remaining taxa studied except
Xenodexia, this process is short, not reaching pterosphenoids
(state 1). In Xenodexia, ascending process of parasphenoids
is absent (state 2). Although this character contributed to the
resolution of the present topology, it presented several inde-
pendent acquisitions and reversals during the history of the
Cyprinodontiformes.
Character 21 - External teeth: (0) conical; (1) compressed.
Tooth form has been extensively employed as the basis
of classification of cyprinodontiform fishes (Günther, 1866;
Garman, 1895; Regan, 1911). The phylogenetic importance of
tooth form was minimized by other precladistic authors in
favor of reproductive characteristics (e.g. Regan, 1913; Rosen
& Bailey, 1963). Rodriguez (1997) reported compressed teeth
in outer series of premaxilla and dentary as synapomorphic
for the clade Xiphophorus, Poecilia, Pamphorichthys, and
Limia with a reversal in Pamphorichthys, which has sub-
cylindrical and pointed teeth. Ghedotti (2000) reported coni-
cal teeth for Aplocheilichthys spilauchen, all procatopodines
examined, Gambusia affinis, Alfaro cultratus, Tomeurus gra-
cilis, Valencia, and Fundulus.
Among studied taxa, procatopodines, Fundulus,
Tomeurus, Alfaro, Brachyrhaphis, Priapichthys, Priapella,
Heterandria, Gambusia, Belonesox, Pseudopoecilia,
Neoheterandria, Scolichthys, Cnesterodon brevirostratus,
and C. septentrionalis possess firmly rooted, conical, pointed
teeth (state 0). In remaining poeciliines studied, teeth are
movable, pedunculate and flattened distally (state 1). On the
basis of present hypothesis, compressed teeth is interpreted
as a synapomorphy for a clade containing the genera
Girardinus, Phallichthys, Xenophallus, Poeciliopsis,
Phalloptychus, Quintana, Carlhubbsia, Xiphophorus,
Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia,
“Poecilia”, Cnesterodon, Phallotorynus,
and Phalloceros [Supertribe Poeciliini - Clade 119], with a
reversal in Cnesterodon brevirostratus + C.
septentrionalis clade.
Hyoid arch
Character 22 - Number of branchiostegal rays (Ghedotti, 2000:
fig. 7): (0) five; (1) six.
This character has been discussed by Ghedotti (2000). 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 (state 1). In indi-
viduals with five branchiostegal rays, one of the anterior two
branchiostegal rays is absent (state 0). The possession of six
branchiostegal rays is interpreted as a synapomorphy for
poeciliines with a reversal to plesiomorphic condition in the
ancestor of members of the supertribe Poeciliini [Clade 119].
Inside this clade a change to state 1 occurred independently
in Xenophallus and Limia.
Fig. 6. Lateral view of left maxilla of (a) Alfaro huberi,
UMMZ 190567; (b) Cnesterodon n. sp. B, MCP 19784. Scale
bar 1 mm.
Fig. 7. Lateral view of inner surface of the anguloarticular of
(a) Phalloceros n. sp. F, MCP 30572; (b) Phalloceros n. sp. C,
UFPB 2214. Scale bar 1 mm.
Character 23 - First and second branchiostegal rays: (0) free
from each other; (1) united at the base.
In most atherinomorphs first and second branchiostegal
rays are free from each other (state 0). In Neoheterandria and
Scolichthys first and second branchiostegal rays are united
at the base (state 1). This condition is interpreted as
synapomorphic for Neoheterandria +Scolichthys [Clade 114].
Character 24 - Anterior process of anterior ceratohyal extend-
ing ventrally to ventral hypohyal (Ghedotti, 2000: fig. 8): (0)
present; (1) absent.
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
was reported absent in all poeciliids examined except
Aplocheilichthys spilauchen by Ghedotti (2000).
We observed this process in Aplocheilichthys, Jenynsia,
Tomeurus, and Brachyrhaphis. All remaining studied taxa
lack the anterior process of anterior ceratohyal extending
ventral to ventral hypohyal (state 1). According to present
hypothesis the absence of this process is synapomorphic for
the subfamily Poeciliinae with reversals in Tomeurus,
Brachyrhaphis, and Heterandria.
Character 25 - Interarcual cartilage: (0) present; (1) absent.
The interarcual cartilage is present (state 0) in all taxa
examined, except in Tomeurus and Priapella. The absence of
an interarcual cartilage (state 1) in these two taxa is inter-
preted as independently acquired.
Branchial arches
Character 26 - Tooth plates of third and fourth
pharingobranchials (Costa, 1991: fig. 4I, 5D and Ghedotti, 2000:
P. H. F. Lucinda & R. E. Reis
13
PROOFS
fig. 10B): (0) two separate plates with teeth irregularly distrib-
uted; (1) fused, forming an elongate structure with teeth regu-
larly distributed.
Most cyprinodontiforms possess tooth plates of third and
fourth pharingobranchials as two separate plates with teeth
irregularly distributed (state 0). Costa (1991) suggested that
tooth plates of third and fourth pharingobranchials fused, form-
ing a elongate structure with teeth regularly distributed (state
1) as a putative synapomorphy for a group embracing
Pamphorichthys, Poecilia, Limia, Xiphophorus, Cnesterodon,
Phalloceros, Phallotorynus, Phalloptychus, Priapichthys,
Poeciliopsis, Priapella, Quintana, Carlhubbsia, Xenodexia,
and Phallichthys. This is partially corroborated by our results.
The current phylogenetic analysis supports this feature as a
uniquely derived and unreversed synapomorphy for the
supertribe Poeciliini [Clade 119]. In addition to the genera above
(except Priapichthys and Priapella), this group comprises
Girardinus, Xenophallus, and Micropoecilia.
Character 27 - Teeth on fourth ceratobranchial (Ghedotti, 2000:
fig. 9): (0) present; (1) absent.
Among studied taxa Aplocheilichthys, Procatopus,
Jenynsia, Fundulus, Tomeurus, Alfaro, Brachyrhaphis,
Priapella, Priapichthys, Heterandria, Gambusia,
Belonesox, Pseudopoecilia, Neoheterandria, Cnesterodon
n. sp. A, C. hypselurus, C. brevirostratus, C. carnegiei, C.
omorgmatos, and Phalloceros n. sp. D exhibit teeth on fourth
ceratobranchial. These are lacking in the remaining studied
taxa. Global parsimony analysis demonstrate that the loss
of teeth in fourth ceratobranchial is apomorphic and oc-
curred independently in Fluviphylax, Cyprinodon,
Scolichthys, and in the ancestor of members of Clade 119,
with subsequent reversals in Phalloceros n. sp. D,
Xiphophorus and in node 103. Within Clade 103, a 0 > 1
change happened in C. septentrionalis.
Character 28 - Fifth ceratobranchial (Costa, 1991: fig. 4H; 5C):
(0) narrow bearing teeth irregularly distributed; (1) wide bear-
ing teeth regularly distributed.
Most cyprinodontiforms possess a narrow fifth
ceratobranchial, bearing irregularly distributed teeth (state
0). Costa (1991) suggested a wide fifth ceratobranchial bear-
ing regularly distributed teeth (state 1) as a putative
synapomorphy for a group embracing Pamphorichthys,
Poecilia, Limia, Xiphophorus, Cnesterodon, Phalloceros,
Phallotorynus, Phalloptychus, Priapichthys, Poeciliopsis,
Priapella, Quintana, Carlhubbsia, Xenodexia, and
Phallichthys. This is partially corroborated by our results.
The current phylogenetic analysis supports this feature as a
uniquely derived and unreversed synapomorphy for the
supertribe Poeciliini [Clade 119]. In addition to the genera
above (except Priapichthys and Priapella), this group com-
prises Girardinus, Xenophallus, and Micropoecilia.
Character 29 - Anterior margin of first hypobranchial: (0)
mostly straight; (1) concave forming distinct anterior point.
Anterior margin of first hypobranchial is mostly straight
(state 0) in Fluviphylax, Procatopus, and Fundulus. Ante-
rior margin of first hypobranchial is concave, forming distinct
anterolateral point (state 1) in Aplocheilichthys, Jenynsia,
Cyprinodon, and all poeciliines. This feature was useless for
poeciliine relationships and probably is a synapomorphy for
a more inclusive clade.
Ghedotti (2000) reported a concave anterior margin of first
hypobranchial in Oxyzygonectes dovii, Cyprinodon variegatus,
Fundulus chrysotus, and all poeciliines examined except some
individuals of Tomeurus gracilis (coded as polymorphic).
Character 30 - Third basibranchial: (0) cartilaginous; (1) ossi-
fied and toothed.
Most cyprinodontiforms possess a cartilaginous toothless
third basibranchial. Phalloceros n. sp. D, Phalloceros n. sp. G,
Phalloceros n. sp. F, Phalloceros n. sp. U, and Phalloceros n.
sp. H are unique among poeciliines by possession of a ossified
and toothed third basibranchial. This derived feature is hy-
pothesized to have appeared independently in Phalloceros n.
sp. D, Phalloceros n. sp. U, and Phalloceros n. sp. H, and in
the ancestor of Phalloceros n. sp. G and Phalloceros n. sp. F.
Pectoral fin and girdle
Character 31 - Post-temporal: (0) bifid; (1) unbranched.
Most cyprinodontiforms possess a bifid post-temporal
(state 0). In Fundulus, Scolichthys, Cnesterodon,
Phallotorynus, and Phalloceros the post-temporal is un-
branched (state 1). According to the present hypothesis of
relationships an unbranched post-temporal is assumed as
apomorphic and independently acquired by Fundulus,
Scolichthys, and by the ancestor of cnesterodontines.
Ghedotti (2000) reported an unbranched posttemporal in
Cnesterodon, Phalloceros caudimaculatus, Phallotorynus,
Valencia, Fundulus and many procatopodines.
Character 32 - Position of pectoral fins: (0) low, below midline;
(1) high, at or above midline.
The position of the pectoral fins is low, with dorsal inser-
tion below midline in most cyprinodontiforms (state 0). In
poeciliids, the position of the pectoral fins is high, with dorsal
insertion at or above midline (state 1). With the exception of
Jenynsia, Fundulus, and Cyprinodon, all studied taxa present
high pectoral fins. This feature probably is synapomorphic for
the family Poeciliidae. In fact, as pointed out by Ghedotti (2000:
25): “Parenti (1981) and Costa (1998) recognized high pectoral
fins as a synapomorphic reversal in poeciliids to the condition
in non-cyprinodontiform atherinomorphs”.
Pelvic fin and girdle
Character 33 - Number of pelvic-fin rays in males: (0) six; (1)
five; (2) four; (3) three.
Males of most atherinomorph fishes have six pelvic-fin
rays (state 0) (Parenti, 1981; Ghedotti, 2000). Ghedotti (2000)
reported less than six pelvic-fin rays as synapomorphic for
Phallotorynus, Phalloceros, Cnesterodon and Tomeurus.
Systematics of the subfamily Poeciliinae Bonaparte
14
PROOFS
Males specimens of Phalloceros and Phallotorynus spe-
cies as well as Cnesterodon n. sp. A, Cnesterodon n. sp. B,
and C. hypselurus possess five pelvic-fin rays (state 1). Males
of Cnesterodon brevirostratus, C. septentrionalis, C.
carnegiei, and C. omorgmatos, Phalloptychus januarius and
P. iheringii exhibit four pelvic-fin rays (state 2). Males of
Cnesterodon raddai Meyer & Etzel and Tomeurus possess
three pelvic-fin rays (state 3). Cnesterodon decemmaculatus
(Jenyns) was coded “-” for it is polymorphic, males having
four or five pelvic-fin rays. Phalloptychus eigenmanni Henn
was coded “?” for character state could not be checked due
to poor condition of the material studied. According to the
present phylogenetic hypothesis state 1 is interpreted as
synapomorphic for a clade containing Cnesterodon,
Phallotorynus, and Phalloceros [Clade 111]. State 2 is hy-
pothesized as independently acquired and synapomorphic
for Phalloptychus species and for the clade [Cnesterodon
brevirostratus + C. septentrionalis + C. carnegiei + C.
omorgmatos]. The presence of state 3 in C. raddai and
Tomeurus is considered homoplastic.
Character 34 - Pelvic-fin length in adult males: (0) short; sec-
ond ray not surpassing the end of anal-fin base; (1) long,
second ray surpassing the end of anal-fin base.
In most cyprinodontiform fishes the second pelvic-fin ray
does not surpass the end of anal-fin base (state 0). In
Xenophallus, Xiphophorus, Xenodexia, Poecilia, Limia,
Pamphorichthys, Micropoecilia, and “Poecilia” the second
pelvic-fin ray surpasses the end of anal-fin base (state 1). Fol-
lowing our phylogenetic study, second pelvic-fin ray surpass-
ing the end of anal-fin base is hypothesized to have been inde-
pendently acquired by Xenophallus and by the ancestor of a
clade comprising Xiphophorus, Xenodexia, Poecilia, Limia,
Pamphorichthys, Micropoecilia, and “Poecilia” [Clade 108].
Rodriguez (1997) reported this feature as synapomorphic for
Xiphophorus, Poecilia, Limia, and Pamphorichthys.
Character 35 - Position of pelvic girdle of males: (0) very pos-
terior, anterior border of basipterygium posterior to posterior
border of cleithrum; (1) posterior; posterior border of cleithrum
approximately aligned with center of basipterygium (or more
posterior); (2) not very anterior; posterior border of
basipterygium aligned with posterior border of cleithrum; (3)
very anterior; located below pectoral girdle; posterior border
of basipterygium anterior to posterior border of cleithrum; (4)
posterior border of cleithrum approximately aligned with an-
terior border of basipterygium.
Pelvic girdle of males of Aplocheilichthys, Fluviphylax,
Jenynsia, Cyprinodon, Fundulus, Brachyrhaphis, Gambu-
sia, Belonesox, Girardinus, Xiphophorus, Xenodexia,
Poecilia, Micropoecilia, and “Poecilia” is very posterior,
anterior border of basipterygium being posterior to posterior
border of cleithrum (state 0).
In Alfaro, Priapella, Priapichthys, Heterandria,
Pseudopoecilia, Phallichthys, Poeciliopsis, Phalloptychus,
Quintana, Carlhubbsia, Pamphorichthys scalpridens,
Phallotorynus, Phalloceros n. sp. B, and Phalloceros n. sp.
D pelvic girdle of males is posteriorly located, with posterior
border of cleithrum approximately aligned with center of
basipterygium (or more posterior) (state1).
In Neoheterandria, Pamphorichthys hollandi, and
Phalloceros species excepting Phalloceros n. sp. B, and
Phalloceros n. sp. D pelvic girdle is not very anteriorly lo-
cated, with posterior border of basipterygium aligned with
posterior border of cleithrum (state 2). In Tomeurus and
Cnesterodon the basipterygium is very anterior, located be-
low pectoral girdle; posterior border of basipterygium ante-
rior to posterior border of cleithrum (state 3). In Procatopus,
Scolichthys, Xenophallus, and Limia posterior border of
cleithrum is approximately aligned with anterior border of
basipterygium (state 4). Although this character contributed
to the resolution of the present topology, it presented several
independent acquisitions and reversals during the history of
the Cyprinodontiformes.
Parenti (1981), Costa (1998), and Ghedotti (2000) recog-
nized an anteriorly positioned pelvic girdle as
synapomorphic for poeciliids and also recognized the pres-
ence of an anterior pelvic girdle in cyprinodontids, and
some individuals of Jenynsia. Ghedotti (2000) also de-
scribed a pelvic girdle under the pectoral girdle in males as
synapomorphic for Tomeurus, Cnesterodon, Phalloceros,
and Phallotorynus.
Character 36 - Dorsolateral process of basipterygium in adult
males (Fig. 8): (0) absent or small; (1) large; (2) enormous.
In most cyprinodontiform fishes a dorsolateral process of
basipterygium in adult males is hardly developed or lacking
(state 0, Fig. 8a). In Tomeurus, Heterandria, Neoheterandria,
Belonesox, Poeciliopsis, and Phalloceros n. sp. U, Phalloceros
n. sp. T, and Phalloceros n. sp. S this process is large (state 1,
Fig. 8b). In Phalloptychus this process is enormous; it is as
long as the remaining basipterygium (state 2, Fig. 8c; 15). Based
on present analysis of relationships, state 1 is interpreted as
independently evolved in Tomeurus, Heterandria,
Neoheterandria, Belonesox, Poeciliopsis, and in the ancestor
of Clade 82, with a reversal in Clade 78. State 2 is hypothesized
as synapomorphic for Phalloptychus species.
Character 37 - Shape of the anterior tip of basipterygium in
adult males: (0) approximately triangular and rounded; (1)
clearly pointed; (2) sinuous; (3) clearly round and keeled.
Among studied taxa, Aplocheilichthys, Fluviphylax,
Jenynsia, Fundulus, Alfaro, Brachyrhaphis,
Neoheterandria, Scolichthys, Phallichthys, Carlhubbsia,
Micropoecilia, “Poecilia”, Phallotorynus, and
Phalloceros n. sp. B possess the anterior tip of
basipterygium approximately triangular and round in adult
males (state 0, Fig. 8d). In Cnesterodon species this struc-
ture is sinuous (state 2, Fig. 9), this condition being
synapomorphic for the genus. In Poeciliopsis and
Phalloptychus the anterior tip of basipterygium is clearly
round and keeled in adult males (state 3, Fig. 8c), which is
P. H. F. Lucinda & R. E. Reis
15
PROOFS
interpreted as synapomorphic for this clade. Remaining stud-
ied taxa exhibit a clearly pointed anterior tip of basipterygium
in adult males (state 1, Fig. 11).
Character 38 - Lateral keel of basipterygium in adult males: (0)
absent; (1) present.
Most cyprinodontiforms lack a lateral keel of
basipterygium in adult males. This derived feature appeared
to have been independently acquired by Scolichthys,
Pamphorichthys, and by the ancestor of Poeciliopsis and
Phalloptychus.
Character 39 - Narrowing of lateral surface of basipterygium
base in adult males (Fig. 9): (0) absent; (1) present.
Fig. 8. Left basipterygium. (a) Priapichthys annectens, ANSP
163139; (b) Heterandria jonesii, UMMZ 210999; (c)
Phalloptychus iheringii, MCP 11054; (d) Micropoecilia
branneri, MCP 22040. DLP = dorsolateral process. Scale bar
1 mm. A and B lateral view. C and D ventral view.
Fig. 9. Basipterygia and first pelvic-fin ray of Cnesterodon
brevirostratus, MCP 13950. Scale bar 1 mm.
Cnesterodon is unique among cyprinodontiforms by the
narrowing of lateral surface of basipterygium base in adult males
(state 1; Fig. 9). This condition is absent in remaining
atherinomorphs (state 0) and is interpreted as synapomorphic
for Cnesterodon.
Character 40 - First ray of left and right pelvic fins in adult males
(Fig. 9, 10): (0) similar to each other; (1) different from each other.
All cyprinodontiform fishes possess first ray of left and right
pelvic fins similar to each other in adult males (state 0, Fig. 9),
except for Phalloptychus species, in which first ray of left pelvic
fin is much wider and more specialized than right one (state 1;
Fig. 10). This condition is hypothesized as synapomorphic for
Phalloptychus.
Character 41 - Width of first pelvic-fin ray in adult males: (0)
approximately constant tapering gradually to tip; (1) decreas-
ing abruptly at distal portion, distal slender portion long; (2)
decreasing abruptly at distal portion, distal slender portion
short; (3) very wide, mainly right one.
Among studied taxa, members of the outgroup, Alfaro,
Brachyrhaphis, Priapella, Priapichthys, Pseudopoecilia,
Gambusia, Belonesox, Phallichthys, Quintana, Carlhubbsia,
Xenodexia, Xiphophorus, Pamphorichthys, Micropoecilia,
“Poecilia”, Phallotorynus, and Phalloceros n. sp. R possess
pelvic-fin ray gradually tapering to tip (state 0) in adult males.
In Neoheterandria, Heterandria, Girardinus, Xenophallus,
Poeciliopsis
, Poecilia, Limia, Phalloceros, (exceptPhalloceros
n. sp. R) width of first pelvic-fin ray in adult males decreases
abruptly at distal portion, and distal slender portion is long (state
1, Fig. 11). In adult males of Tomeurus and Cnesterodon, the first
pelvic-fin ray decreases abruptly at distal portion, and distal
slender portion is short (state 2, Fig. 9), which is synapomorphic
for the genus and independently acquired in Tomeurus. In
Phalloptychus the first pelvic-fin ray in adult males is very wide,
especially the right (state 3, Fig. 10). This condition is inter-
preted as synapomorphic for Phalloptychus.
Character 42 - Second pelvic-fin ray in adult males: (0)
branched; (1) unbranched.
Second pelvic-fin ray is branched (state 0) in adult males
of cyprinodontiforms, with the exception of Tomeurus and
Cnesterodon species, which possess unbranched second
pelvic-fin ray (state 1). Following the present hypothesis,
Tomeurus and Cnesterodon independently acquired an un-
branched second pelvic-fin ray in adult males. A reversal oc-
curs in C. septentrionalis.
Character 43 - Lateral projection near the bifurcation of sec-
ond right pelvic-fin ray in adult males (Fig. 10a): (0) absent;
(1) present.
Phalloptychus is unique among cyprinodontiform fishes
by the possession of a lateral projection near the bifurca-
tion of second right pelvic-fin ray in adult males (Fig. 10a).
This condition is interpreted as synapomorphic for
Phalloptychus.
Systematics of the subfamily Poeciliinae Bonaparte
16
PROOFS
Fig. 12. First right pelvic-fin ray in adult male of Phalloptychus
iheringii, MCP 11054. C = callosity. Scale bar 1 mm.
Character 44 - Number of pelvic-fin rays in females: (0) six or
seven; (1) five; (2) three.
Females of most cyprinodontiforms possess six or seven
pelvic-fin rays (state 0). Ghedotti (2000) reported less than six
pelvic-fin rays as synapomorphic for Phallotorynus,
Phalloceros, Cnesterodon and Tomeurus.
Females of Phalloptychus, Cnesterodon, Phallotorynus,
and Phalloceros possess five anal-fin rays (state 1). Females
ofTomeurus exhibit three pelvic-fin rays (state 2). Tracking the
present historical hypothesis for poeciliines, state 1 is pro-
posed as synapomorphic for Phalloptychus januarius and P.
iheringii. This derived feature is independently acquired by
and also synapomorphic for a clade embracing Cnesterodon,
Phallotorynus, and Phalloceros [Clade 111].
Character 45 - Callosity at the distal portion of first right pel-
vic fin in adult males (Fig. 12): (0) absent; (1) present.
Phalloptychus is unique among cyprinodontiform fishes
by the possession of a callosity at the distal portion of right
pelvic fin in adult males. This condition is interpreted as
synapomorphic for Phalloptychus (state 1; Fig. 12).
Axial skeleton
Character 46 - Ligastyle (Fig. 13): (0) absent; (1) with one axis;
(2) triangular; (3) tripartite (three axis).
Ghedotti (2000) reported the presence of an ossified
ligastyle in Alfaro, Priapella, Gambusia, Heterandria,
Poeciliopsis, Girardinus, Phallichthys, Phallotorynus,
Phalloceros, and Cnesterodon.
A ligastyle is absent in Aplocheilichthys, Fluviphylax,
Procatopus, Jenynsia, Fundulus, Tomeurus, Pseudopoecilia,
Scolichthys, Poecilia, Limia, Pamphorichthys,
Micropoecilia
, “Poecilia”, and Cnesterodon (state 0).
Cyprinodon, Alfaro, Brachyrhaphis, Gambusia,
Priapichthys, Poeciliopsis, Xenophallus, Xiphophorus,
Xenodexia, Carlhubbsia, Phallotorynus, and Phalloceros
possess a rod-like monoaxial ligastyle (state 1, Fig. 13a). The
ligastyle is triangular (state 2, Fig. 13b) in Priapella,
Heterandria, Belonesox, Girardinus, Phallichthys, and
Quintana; whereas Phalloptychus and Neoheterandria pos-
sess a tripartite ligastyle (state 3, Fig. 13c). Although this
character contributed to the resolution of the present topol-
ogy, it presented several independent acquisitions and re-
versals during the history of the Cyprinodontiformes.
Character 47 - Haemal arch and spine of vertebrae 13-17 in
adult males: (0) typical and similar to the remaining; (1) ab-
sent; (2) modified in gonapophyses; (3) modified in rudimen-
tary gonapophyses.
Haemal arch and spine of vertebrae 13-17 in adult males
are typical and similar to the remaining vertebrae in the
outgroup and Alfaro (state 0). These are absent (state 1) in
Tomeurus and are modified in gonapophyses (state 2) in all
remaining poeciliines. In Cnesterodon, gonapophyses are
rudimentary (state 3). Tracking the evolutive history of this
character reveals that: (1) plesiomorphic condition is only
kept by Alfaro; (2) Tomeurus lacked the haemal arch and spine
Fig. 10. First and second pelvic-fin rays in adult male of
Phalloptychus iheringii, MCP 11054. (a) Second right pelvic-fin
ray; (b) first right pelvic-fin ray; (c) second left pelvic-fin ray; (d)
first left pelvic-fin ray. LP = lateral projection. Scale bar 1 mm.
Fig. 11. Basipterygia and first pelvic-fin ray of Phalloceros
n. sp. G, MCP 30509. Scale bar 1 mm.
P. H. F. Lucinda & R. E. Reis
17
PROOFS
of vertebrae 13-17 in adult males; (3) haemal arch and spine of
vertebrae 13-17 in adult males were modified in gonapophyses
in the ancestor of all remaining poeciliines; (4) Cnesterodon
shows a hypotrophy of gonapophyses to a vestigial stage.
Rosen & Bailey (1963) described the structure and distri-
bution of gonapophyses in many poeciliines. Ghedotti (2000)
reported the presence of gonapophyses in poeciliines, ex-
cept in Tomeurus and Cnesterodon.
Character 48 - Number of well-developed gonapophyses: (0)
zero (absent); (1) three; (2) two, rarely one; (3) four.
Among studied taxa, members of the outgroup, Tomeurus,
Alfaro, and Cnesterodon species lack well-developed
gonapophyses (state 0). Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” possess two (rarely one) well-
developed gonapophyses (state 2). Xenodexia exhibit four
well-developed gonapophyses (state 3). The remaining stud-
ied taxa present three well-developed gonapophyses (state
1). Results of the present phylogenetic analysis support the
following interpretation of character evolution: (1) state 1
appeared once in the history of poeciliines in the ancestor of
members of Clade 125; (2) this state is posteriorly modified to
state 2 in the ancestor of members of Clade 99; (3) state 3 is
interpreted as autapomorphic for Xenodexia; and (4) a rever-
sal to state 0 occurred in Cnesterodon.
Character 49 - Position of functional gonapophyses: (0) ab-
sent; (1) on vertebrae 14, 15, 16; (2) located between verte-
brae 13 to 15, but never on vertebra 16 (13-14-15; 14-15; 13-
14; 13); (3) on vertebrae15, 16, and 17; (4) on vertebrae 13-16
or 14-17.
Functional gonapophyses are absent in members of the
outgroup and in Tomeurus, Alfaro, and Cnesterodon (state
0). Functional gonapophyses are located on vertebrae 14, 15,
16 (state 1). Functional gonapophyses are located on verte-
brae13 to 15, but never on vertebra 16 (13-14-15; 14-15; 13-14;
13) (state 2). Functional gonapophyses are located on verte-
brae 15, 16, and 17 (state 3). Functional gonapophyses are
located on vertebrae 13-16 or 14-17 (state 4). Our results sup-
port the assumption that state 1 is synapomorphic for a clade
comprising all poeciliines but Tomeurus and Alfaro [Clade
125], with a reversal to state 0 in Cnesterodon [Clade 0107].
State 2 is interpreted as a synapomorphy for the clade
[Poecilia + Limia + Pamphorichthys + Micropoecilia +
“Poecilia”] [Clade 99]. State 3 and 4 are considered as
autapomorphic for Phalloceros n. sp. R and for Xenodexia,
respectively.
Character 50 – First gonapophysis reduced to a support for
the adjacent gonapophysis located in vertebra 12 or 13
(Rodriguez, 1997: fig. 4B): (0) absent; (1) present.
Rodriguez (1997) proposed that Pamphorichthys is unique
among cyprinodontiform fishes by the possession of a re-
duced gonapophysis to a support for the adjacent
gonapophysis located in vertebra 12 or 13. This condition is
interpreted as synapomorphic for Pamphorichthys.
Fig. 13. Ligastyle of (a) Carlhubbsia kidderi
, UMMZ 184619;
(b) Heterandria jonesii, UMMZ 210999; (c) Phalloptychus
iheringii, MCP 11054. Scale bar 1 mm.
Character 51 - Hollister’s foramen on first or second
gonapophysis (Rodriguez, 1997: fig. 4B): (0) absent; (1)
present.
Rodriguez (1997) reported the presence of Hollister’s
foramen on first or second gonapophysis as
synapomorphic for Pamphorichthys, Poecilia, and Limia.
However, we did not observe this feature in Limia. We
have observed this feature in Poecilia, Pamphorichthys,
Micropoecilia, and “Poecilia”. Hollister’s foramen on first
or second gonapophysis is herein interpreted as
synapomorphic for a clade containing Poecilia,
Pamphorichthys, Limia, Micropoecilia, and “Poecilia”,
with a reversal in Limia.
Character 52 - Gonapophysis of vertebra 14: (0) slightly
curved; (1) very curved; (2) straight; (3) rudimentary; (4)
bearing an abrupt break forming an acute angle at subdistal
portion.
Gonapophysis of vertebra 14 is slightly curved (state 0,
Rosen & Bailey, 1963: fig. 46) in Brachyrhaphis,Priapichthys,
Priapella, Heterandria, Gambusia, Belonesox,
Pseudopoecilia, Neoheterandria, Scolichthys, Girardinus,
Phallichthys, Xenophallus, Quintana, Carlhubbsia,
Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia”. In Poeciliopsis and
Phalloptychus gonapophysis of vertebra 14 is strongly
curved (state 1, Rosen & Bailey, 1963: fig. 26C; 56A-D).
Gonapophysis of vertebra 14 is straight (state 2, Rosen &
Bailey, 1963: fig. 24A) in Xiphophorus and Phalloceros n. sp.
J. Gonapophysis of vertebra 14 is rudimentary (state 3, Rosen
& Bailey, 1963: fig. 26D) in Cnesterodon. Phalloceros spe-
cies except Phalloceros n. sp. J. possess gonapophysis of
vertebra 14 bearing an abrupt break forming an acute angle at
Systematics of the subfamily Poeciliinae Bonaparte
18
PROOFS
subdistal portion (state 4, Rosen & Bailey, 1963: fig. 29B). The
present phylogenetic analysis indicate: (1) state 1 as
synapomorphic for Poeciliopsis + Phalloptychus [Clade 105];
(2) state 2 independently acquired in Xiphophorus and
Phalloceros n. sp. J; (3) state 3 synapomorphic for Cnesterodon;
and (4) state 4 as synapomorphic for Phalloceros species, with
subsequent change to state 2 in Phalloceros n. sp. J.
Character 53 - Curvature of first gonapophysis relative to
vertebral column: (0) 46-75 degrees; (1) 16-45 degrees; (2) 5-
15 degrees; (3) zero degree (approximately parallel); (4) ap-
proximately 90 degrees (approximately perpendicular).
In Brachyrhaphis, Priapichthys, Phallichthys, Quintana,
and Xenodexia first gonapophysis is angled 46-75 degrees
relative to vertebral column (state 0). In Priapella,
Heterandria, Belonesox, Neoheterandria, Scolichthys,
Girardinus, Xenophallus, Carlhubbsia, Poecilia, Limia,
Phallotorynus, Phalloceros (except Phalloceros n. sp. C),
the first gonapophysis is angled 16-45 degrees relatively to
vertebral column (state 1). A curvature of 5-15 degrees (state
2) is present in Gambusia, Pseudopoecilia, Poeciliopsis,
Micropoecilia sp., “Poecilia”, and Phalloceros n. sp. C. In
Phalloptychus, Pamphorichthys, Micropoecilia branneri
(Eigenmann), first gonapophysis is parallel to vertebral col-
umn (state 3). In Xiphophorus, first gonapophysis is approxi-
mately perpendicular to vertebral column (state 4). Outgroup
taxa, Tomeurus, Alfaro, and Cnesterodon were coded “-” for
they lack gonapophysis 1. Although this character contrib-
uted to the resolution of the present topology, it presented
several independent acquisitions and reversals during the
history of the Cyprinodontiformes.
The curvature of gonapophyses (in general) was employed
by Rodriguez (1997), however, this author codified only two
character states: gonapophyses forming an acute angle relative
to vertebral column and perpendicular to vertebral column.
Character 54 - Curvature of second gonapophysis relative to
vertebral column: (0) 15-45 degrees; (1) 45-70 degrees; (2)
zero degrees (approximately parallel); (3) zero-15 degrees; (4)
approximately 90 degrees (approximately perpendicular).
In Priapella, Heterandria
, Belonesox, Pseudopoecilia,
Neoheterandria, Scolichthys, Girardinus, Phallichthys,
Xenophallus, Poeciliopsis, Poecilia, Phallotorynus, and
Phalloceros (except Phalloceros n. sp. B) second
gonapophysis is angled 15-45 degrees relative to vertebral
column (state 0). In Priapichthys, Quintana, Carlhubbsia,
and Xenodexia the second gonapophysis is angled 45-70
degrees relatively to vertebral column (state 1). In
Phalloptychus, Pamphorichthys, Micropoecilia branneri,
second gonapophysis is approximately parallel to vertebral
column (state 2). A curvature of zero-15 degrees (state 3) is
present in Gambusia, Micropoecilia branneri, and
“Poecilia”. In Brachyrhaphis and Xiphophorus second
gonapophysis is approximately perpendicular to vertebral
column (state 4). Outgroup taxa, Tomeurus, Alfaro, and
Cnesterodon were coded “-” for they lack gonapophysis 2.
Although this character contributed to the resolution of the
present topology, it presented several independent acquisi-
tions and reversals during the history of the
Cyprinodontiformes.
Character 55 - Curvature of third gonapophysis relative to
vertebral column: (0) 90 degrees (approximately perpendicu-
lar); (1) 35-70 degrees; (2) 10-32 degrees; (3) zero-10 degrees;
(4) zero (parallel).
In Brachyrhaphis and Xiphophorus, third gonapophysis
is approximately perpendicular to vertebral column (state 0).
In Belonesox, Priapella, Priapichthys, Quintana,
Carlhubbsia, and Xenodexia third gonapophysis is angled
35-70 degrees relatively to vertebral column (state 1). A cur-
vature of 10-32 degrees (state 2) is present in Heterandria,
Pseudopoecilia, Scolichthys, Girardinus, Xenophallus,
Poeciliopsis, Phallotorynus, and Phalloceros. In
Phalloptychus third gonapophysis is angled zero-10 degrees
relatively to vertebral column (state 3). In Gambusia third
gonapophysis is approximately parallel to vertebral column
(state 4). Outgroup taxa, Tomeurus, Alfaro, Poecilia, Limia,
Pamphorichthys, Micropoecilia, “Poecilia”, and
Cnesterodon were coded “-” for they lack third gonapophysis.
The present phylogenetic analysis indicates that state 1
appeared at the ancestor of Clade 124. State 2 is supposed to
have been acquired by the ancestor of members of Clade 122,
with reversals to state 1 in Belonesox, Phallichthys, and in
Clade 112. A reversal to state 0 occurs in Xiphophorus. State
3 is interpreted as synapomorphic for Phalloptychus.
Character 56 - Distal portion of ribs (6, 7, and 8) in adult males
(Fig. 14): (0) not expanded; (1) expanded.
Adult males of cyprinodontiform fishes (with the exception
of Cnesterodon species) exhibit the distal portions of pleural
ribs 6, 7, and 8 not expanded (state 0). Rosa & Costa (1993)
recognized the presence of winglike expansions on distal por-
tions of male pleural ribs, as synapomorphic for Cnesterodon
species. Cnesterodon is unique among cyprinodontiforms by
the expansion of distal portion of pleural ribs 6, 7, and 8 in adult
males (state 1). This condition is herein also interpreted as
synapomorphic for Cnesterodon species.
Character 57 - Length of pleural rib 7 in adult males (Fig. 14):
(0) shorter than pleural rib 8; (1) longer than pleural rib 8.
In most cyprinodontiforms the distal tip of pleural rib 7 in
adult males does not surpass that of pleural rib 8 (state 0).
Cnesterodon is unique among cyprinodontiforms by the
length of pleural rib 7 in adult males, which surpasses that of
pleural rib 8. This condition is interpreted as synapomorphic
for Cnesterodon (state 1). The possession of this state by
Phalloceros n. sp. I is assumed as homoplastic.
Character 58 - Pleural rib 9 in adult males (Fig. 15): (0) normally
developed; (1) well-developed.
Most cyprinodontiform fishes possess the pleural rib
normally developed, i.e. similar to the remaining
P. H. F. Lucinda & R. E. Reis
19
PROOFS
Fig. 15. Basipterygium and pleural ribs 6-9 in adult male of
Phalloptychus iheringii, MCP 11054. Scale bar 1 mm.
Character 59 - Curvature of pleural ribs in adult males: (0)
approximately parallel and slightly arched forward, their dis-
tal tip not converging to the same point; (1) pleural ribs 7, 8,
and 9 are curved forward not converging to the same point
towards pelvic girdle; (2) pleural ribs 6, 7, and 8 are curved
forward converging to the same point towards pelvic girdle.
Pleural ribs 6-9 of males of most cyprinodontiform fishes
are approximately parallel and slightly arched forward, their
distal tip not converging to the same point (state 0). In
Priapella, Heterandria, Pseudopoecilia, Neoheterandria,
Scolichthys, Girardinus, Phallichthys, Xenophallus,
Poeciliopsis, Phalloptychus, Quintana, and Carlhubbsia
pleural ribs 7, 8, and 9 are curved forward not converging to
the same point towards pelvic girdle (state 1). In Cnesterodon
species, excepting C. raddai, Phallotorynus and Phalloceros
pleural ribs 6, 7, and 8 are curved forward converging to the
same point towards pelvic girdle (state 2).
The curvature of pleural ribs in adult males was described
and discussed by Rosen & Bailey (1963) and Rosa & Costa
(1993). Ghedotti (2000: 24) discussed the rib condition in adult
male. However, the rib condition of Tomeurus as described
by Ghedotti (2000) himself is far autapomorphically peculiar
and different from states 1 and 2 of the present character,
therefore Tomeurus was coded 0 for this character.
Results of the present phylogenetic analysis support the
following interpretation of character evolution: (1) state 1
appeared once in the history of poeciliines in the ancestor of
members of Clade 123, with reversals in the Gambusia +
Belonesox clade [Clade 118] and in the clade composed of
Xiphophorus, Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” [Clade 108]; (2) state 2 is
synapomorphic for a clade comprising Cnesterodon,
Phallotorynus, and Phalloceros [Clade 111], with a reversal
to state 0 in Cnesterodon raddai.
Character 60 - Pleural ribs association with haemal arches in
males: (0) absent; (1) present.
There is no consensus among authors concerning this
character. Basal atherinomorph fishes do not exhibit pleural
ribs associated with haemal arches (state 0). Parenti (1981)
recognized pleural ribs on haemal arches as synapomorphic
for Poeciliidae. However, Costa (1998) did not notice the de-
rived feature in the poeciliids examined by him: Alfaro,
Aplocheilichthys Brachyrhaphis, Cnesterodon, Fluviphylax,
Hylopanchax, Limia, Pamphorichthys, Phalloceros
,
Phalloptychus, Poecilia, Procatopus, Tomeurus, and
Xiphophorus.
On the other hand, Ghedotti (2000) reported pleural ribs
on haemal arches in Aplocheilichthys, Micropanchax Myers,
Procatopus, Fluviphylax, Cubanichthys, Cyprinodon, and
all poeciliines examined (except Tomeurus and Poeciliopsis):
Alfaro, Priapella, Gambusia, Heterandria, Girardinus,
Poecilia, Phallichthys, Phallotorynus, Phalloceros, and
Cnesterodon.
We opted to examine this character in males and females
separately, i.e. we split this character in two, because the
atherinomorphs (state 0). Phalloptychus is unique among
cyprinodontiform fishes by having pleural rib 9 well-devel-
oped in adult males, i.e. longer than remaining pleural ribs,
curved forward and expanded at distal tip (state 1). This con-
dition is interpreted as synapomorphic for Phalloptychus.
Fig. 14. Pleural ribs 5-9 in adult male of Cnesterodon
brevirostratus, MCP 13950. Scale bar 1 mm.
Systematics of the subfamily Poeciliinae Bonaparte
20
PROOFS
presence of haemal arches associated with pleural ribs may
vary between sexes. Pleural ribs associated with haemal arches
were found in males of Alfaro, Gambusia, Belonesox,
Priapella, Phalloptychus, Scolichthys, Poecilia, “Poecilia”,
Xiphophorus, Limia, Pamphorichthys, Micropoecilia,
Carlhubbsia, Xenodexia, Cnesterodon (except C.
septentrionalis), Phalloceros, and Phallotorynus. We inter-
preted this character as independently acquired in Alfaro,
Priapella, in Gambusia + Belonesox clade, Scolichthys,
Phalloptychus, in the ancestor of members of Clade 115, with
reversals in C. septentrionalis and Quintana.
Character 61 - Pleural ribs association with haemal arches in
females: (0) absent; (1) present.
Pleural ribs associated with haemal arches were found in
females of Alfaro, Priapella, Gambusia, Phalloptychus,
Quintana, Poecilia, “Poecilia”, Xiphophorus, Limia,
Pamphorichthys, Micropoecilia, Carlhubbsia, and Xenodexia,
Cnesterodon decemmaculatus, C. hypselurus, Cnesterodon n.
sp. A, C. omorgmatos,
C. brevirostratus, Phallotorynus (ex-
cept Phallotorynus n. sp. A), and Phalloceros.
Although this character contributed to the resolution of
the present topology, it presented several independent ac-
quisitions and reversals during the history of the
Cyprinodontiformes.
Dorsal Fin
Character 62 - Position of the first proximal radial of dorsal fin
in adult males: Located between neural arches of vertebrae:
(0) 12 and 13; (1) 13 and 14; (2) 8 and 9; (3) 23 and 24 or 24 and
25; (4) 10 and 11; (5) 15 and 16 or 16 and 17; (6) 11 and 12; (7)
14 and 15; (8) 7 and 8.
Ghedotti (2000) studied the position of dorsal-fin origin
relative to the origin of anal fin. We opted to examine this
character by means of the position of the first proximal radial
of dorsal fin in adult males relative to neural arches of verte-
brae. We also examined males and females separately, i.e.we
split this character in two, because the position of dorsal-
and anal-fin origins may vary between sexes.
First proximal radial of dorsal fin in adult males is located be-
tween neural spines of vertebrae 12 and 13 (state 0) in Fundulus,
Priapichthys,Priapella,Poeciliopsis,Phalloptychus,“Poecilia”,
Cnesterodon decemmaculatus, and Cnesterodon n. sp. B.
In Gambusia, Pseudopoecilia, Scolichthys, Girardinus,
Micropoecilia, Cnesterodon n. sp. A, C. brevirostratus, C.
carnegiei, C. omorgmatos, Phallotorynus, and most
Phalloceros species (except Phalloceros n. sp. U, Phalloceros
n. sp. R, Phalloceros n. sp. N, Phalloceros n. sp. O, Phalloceros
n. sp. M, Phalloceros n. sp. H, and Phalloceros n. sp. J) the
first proximal radial of dorsal fin in adult males is located be-
tween neural spines of vertebrae 13 and 14 (state 1).
Neoheterandria, Carlhubbsia, and Xiphophorus exhibit
this structure located between neural spines of vertebrae 8
and 9 (state 2). In Tomeurus the first proximal radial of dorsal
fin in adult males is located between neural spines of verte-
brae 23 and 24 or 24 and 25 (state 3). Jenynsia, Phallichthys,
Xenophallus, Limia, and Pamphorichthys present this ele-
ment located between neural spines of vertebrae 10 and 11
(state 4)
In Fluviphylax, this radial is located between neural spines
of vertebrae 15 and 16 or 16 and 17 (state 5). Brachyrhaphis,
Heterandria, Quintana, Xenodexia, Poecilia, and
Cnesterodon septentrionalis possess the first proximal ra-
dial of dorsal fin in adult males located between neural spines
of vertebrae 11 and 12 (state 6). In Phalloceros n. sp. U,
Phalloceros n. sp. R, and Phalloceros n. sp. O, this structure
is located between neural spines of vertebrae 14 and 15 (state
7) and in Cyprinodon it is located between neural spines of
vertebrae 7 and 8 (state 8).
Although this character contributed to the resolution of
the present topology, it presented several independent ac-
quisitions and reversals (24 steps) during the history of the
Cyprinodontiformes.
Character 63 - Position of the first proximal radial of dorsal fin
in adult females: Located between neural arches of vertebrae:
(0) 12 and 13; (1) 13 and 14; (2) 10 and 11; (3) 11 and 12; (4) 14
and 15; (5) 15 and 16; (6) 23 and 24 or 24 and 25; (7) 8 and 9.
First proximal radial of dorsal fin in adult females located
between neural spines of vertebrae 12 and 13 (state 0) in
Aplocheilichthys, Fundulus, Phalloptychus, Poeciliopsis, C.
brevirostratus, C. carnegiei, and C. omorgmatos. In
Belonesox, Pseudopoecilia, Scolichthys, Girardinus,
Micropoecilia, Cnesterodon n. sp. A, Cnesterodon n. sp. B.,
Phallotorynus jucundus, P. victoriae, Phallotorynus n. sp.
B, and Phalloceros species (except Phalloceros n. sp. P,
Phalloceros n. sp. M, Phalloceros n. sp. J) the first proximal
radial of dorsal fin in adult females located between neural
spines of vertebrae 13 and 14 (state 1).
Cyprinodon
, Phallichthys, Xenophallus, Xenodexia,
Limia, and Pamphorichthys exhibit this structure located
between neural spines of vertebrae 10 and 11 (state 2). In
Brachyrhaphis, Quintana, “Poecilia”, and C.
septentrionalis the first proximal radial of dorsal fin in adult
females is located between neural spines of vertebrae 11 and
12 (state 3). Alfaro, Gambusia, Neoheterandria,
Phallotorynus n. sp. A, Phalloceros n. sp. P, Phalloceros n.
sp. M, and Phalloceros n. sp. J present this element located
between neural spines of vertebrae 14 and 15 (state 4). In
Fluviphylax and Phallotorynus fasciolatus this radial is lo-
cated between neural spines of vertebrae 15 and 16 (state 5).
Tomeurus possess the first proximal radial of dorsal fin in
adult females located between neural spines of vertebrae 23
and 24 or 24 and 25 (state 6). In Carlhubbsia and Xiphophorus
this structure is located between neural spines of vertebrae 8
and 9 (state 7). Although this character contributed to the
resolution of the present topology, it presented several inde-
pendent acquisitions and reversals during the history of the
Cyprinodontiformes.
Character 64 - Number of dorsal-fin rays (males and females):
(0) ten or more; (1) nine; (2) eight; (3) seven; (4) six.
P. H. F. Lucinda & R. E. Reis
21
PROOFS
Among the studied taxa, Procatopus, Fundulus,
Cyprinodon, Alfaro, Priapella, Priapichthys, Heterandria,
Pseudopoecilia, Phallichthys, Carlhubbsia, Xiphophorus,
Xenodexia, and Phallotorynus jucundus possess 10 or more
dorsal-fin rays (state 0). Jenynsia, Brachyrhaphis,
Scolichthys, Girardinus, Xenophallus, Phalloptychus,
Quintana, and Phalloceros n. sp. S have nine dorsal-fin rays
(state 1). Eight dorsal-fin rays (state 2) are present in
Aplocheilichthys, Gambusia, Poeciliopsis, Poecilia,
Micropoecilia branneri, Cnesterodon, Phallotorynus (ex-
cept jucundus), and Phalloceros (except Phalloceros n. sp.
S). Finally, Pamphorichthys possess seven dorsal-fin rays
(state 3), and Tomeurus has six (state 4). Although this char-
acter contributed to the resolution of the present topology, it
presented several independent acquisitions and reversals
during the history of the Cyprinodontiformes. This character
was discussed at some length by Ghedotti (2000).
Anal Fin
Character 65 - Number of anal-fin rays in females: (0) twelve
or more; (1) ten; (2) eleven; (3) nine; (4) eight.
Among the studied taxa Aplocheilichthys, Procatopus,
Cyprinodon, and Alfaro possess 12 or more anal-fin rays
(state 0). Fundulus, Jenynsia, Priapichthys, Brachyrhaphis,
Scolichthys, Belonesox, Neoheterandria
, Xenophallus,
Poeciliopsis, Phalloptychus, Xiphophorus, Limia,
“Poecilia”, Cnesterodon, Phallotorynus (except P.
fasciolatus and P. victoriae), Phalloceros n. sp. D,
Phalloceros n. sp. P, Phalloceros n. sp. M, and Phalloceros
n. sp. Q have 10 anal-fin rays (state 1). Eleven anal-fin rays
(state 2), is found in Tomeurus, Priapella, Heterandria,
Girardinus, Gambusia, Phallichthys, Pseudopoecilia,
Quintana, Carlhubbsia, Xenodexia, Phallotorynus
fasciolatus, P. victoriae, and most Phalloceros species.
Poecilia, Micropoecilia, and Pamphorichthys have nine
anal-fin rays (state 3), and finally Fluviphylax possess eight
anal-fin rays (state 4). Although this character contributed to
the resolution of the present topology, it presented several
independent acquisitions and reversals during the history of
the Cyprinodontiformes. This character was discussed at
some length by Ghedotti (2000).
Character 66 - Disposition of anal-fin proximal radials of fe-
males: (0) parallel; (1) divergent, i.e., first radial slightly in-
clined forwards; (2) convergent.
Most cyprinodontiform fishes possess anal-fin proximal
radials of females in parallel disposition (state 0). Quintana,
Carlhubbsia, Phallotorynus victoriae, Phallotorynus
jucundus, and Phallotorynus n. sp. A exhibit a divergent dis-
position of anal-fin proximal radials of females (state 1). Anal-
fin proximal radials of females converge to the same point,
i.e., present a convergent disposition in Xenophallus,
Poeciliopsis, and Phalloptychus (state 2). The results of the
present phylogenetic hypothesis indicate that state 1 was
independently acquired by (1) the ancestor of Quintana and
Carlhubbsia; and (2) by the ancestor of Phallotorynus
victoriae, Phallotorynus jucundus, Phallotorynus n. sp. A,
and Phallotorynus n. sp. B, with a reversal in Phallotorynus
n. sp. A. State 2 is interpreted as uniquely derived and
unreversed synapomorphy for a clade comprising
Xenophallus + Poeciliopsis + Phalloptychus [Clade 110].
Character 67 - Second, third, and fourth gonactinosts fused
into a complex gonactinost: (0) absent; (1) present.
Second, third, and fourth gonactinosts are fused into a
gonactinost complex in all poeciliines except Xenodexia. This
feature is therefore hypothesized as a synapomorphy for
Poeciliinae with a reversal in Xenodexia.
This character has long been used as diagnostic for
poeciliines (e.g. Rosen & Gordon, 1953; Rosen & Bailey, 1963).
Character 68 - Inclination of gonactinost complex relative to
body longitudinal axis: (0) very inclined backwards, forming
a less than 45 degrees angle with body longitudinal axis; (1)
gonactinost complex approximately perpendicular to body
longitudinal axis; (2) inclined forward, forming a more than 90
degrees angle with body longitudinal axis; (3) little inclined
backwards, forming a angle between 45 and 90 degrees with
body longitudinal axis.
Ghedotti (2000) employed this character, however, recog-
nizing two states “(0) anteriorly inclined or vertical and (1)
posteriorly inclined“. This author reported a posteriorly in-
clined gonactinost complex as synapomorphic for
Cnesterodon and Tomeurus and independently acquired in
Alfaro.
In Tomeurus and Cnesterodon the gonactinost complex
is very inclined backwards to an angle smaller than 45
º
rela-
tive to the body longitudinal axis (state 0). On the basis of the
present hypothesis of relationships, the condition in
Cnesterodon is interpreted as a synapomorphic reversal and
as plesiomorphic in Tomeurus.
Gonactinost complex is approximately perpendicular to
body longitudinal axis (state 1) in Alfaro, Scolichthys,
Poecilia, Pamphorichthys, Micropoecilia, “Poecilia”,
Phalloceros, and Phallotorynus. This condition is herein
hypothesized as independent acquisitions in Alfaro,
Scolichthys, Poecilia, in Pamphorichthys + Micropoecilia
+ “Poecilia” Clade [92], and in Phallotorynus + Phalloceros
Clade [106].
In Brachyrhaphis, Priapella, Priapichthys, Heterandria,
Gambusia, Belonesox, Neoheterandria, Pseudopoecilia,
Girardinus, Phallichthys, Xenophallus, Poeciliopsis,
Quintana, Carlhubbsia, Xiphophorus, Xenodexia, and Limia
the gonactinost complex is inclined forward, forming an angle
wider than 90º relative to the body longitudinal axis (state 2).
Our results support the supposition that this state is a
uniquely derived synapomorphy (with subsequent reversals)
acquired by the ancestor of members of Clade 125.
In Phalloptychus the gonactinost complex is a little in-
clined backwards, forming an angle of 45
º
and 90
º
with body
longitudinal axis (state 3); this is hypothesized as a uniquely
derived and unreversed feature of Phalloptychus species.
Systematics of the subfamily Poeciliinae Bonaparte
22
PROOFS
Character 69 - Basal process on first gonactinost: (0) absent;
(1) small; (2) large.
Brachyrhaphis, Priapella, Belonesox, Pseudopoecilia,
Scolichthys, Girardinus, Carlhubbsia, Xiphophorus,
Phallotorynus, and Phalloceros species (except Phalloceros
n. sp. A) possess a small basal process on first gonactinost
(state 1, Rosen & Bailey, 1963: fig. 29A). This derived feature
presented several independent acquisitions and reversals dur-
ing the history of the Cyprinodontiformes. In Cnesterodon
species this process is enlarged (state 2, Rosen & Bailey, 1963:
fig. 30) and is synapomorphic for the species of Cnesterodon.
Remaining studied taxa lack such process (state 0).
Character 70 - Anterior border of second gonactinost: (0)
straight; (1) bearing a convex expansion.
The anterior border of second proximal radial of anal fin in
adult males of most atherinomorphs fishes is straight (state
0). However, in Priapichthys, Heterandria, Pseudopoecilia,
Neoheterandria, Girardinus, Phallichthys, Xenophallus,
Poeciliopsis, Carlhubbsia, Pamphorichthys hollandi,
Micropoecilia, “Poecilia”, and Phallotorynus the anterior
border of second gonactinost exhibits a convex expansion in
adult males (state 1, Rosen & Bailey, 1963: fig. 29A). Although
this character contributed to the resolution of the present
topology, it presented several independent acquisitions and
reversals during the history of the Cyprinodontiformes.
Character 71 - Distal portion of second and third gonactinosts
(Fig. 16): (0) separate; (1) fused; (2) coalescent only at distal
tip, forming an oblong aperture.
In most cyprinodontiform fishes the distal portion of sec-
ond and third anal-fin proximal radials in adult males is sepa-
rate (state 0, Fig. 16a). In Alfaro, Priapichthys, Heterandria,
Belonesox, Neoheterandria, Girardinus, Phallichthys,
Xenophallus, Poeciliopsis, Phalloptychus, Limia,
Pamphorichthys, Cnesterodon n. sp. A, Phalloceros n. sp.
A, and Phalloceros n. sp. H the distal portion of second and
third gonactinost is fused (state 1, Fig. 16b, e). This feature
presented several independent acquisitions and reversals
during the history of the Cyprinodontiformes.
In Phallotorynus
n. sp. A and Phallotorynus n. sp. B
these are coalescent only at distal tip, forming an oblong
aperture (state 2, Fig. 16c), with is interpreted as a
synapomorphy for these two species.
Character 72 - Fusion of second and third anal-fin gonactinosts
(Fig. 16a): (0) absent; (1) complete; (2) partial.
In outgroup taxa, Xenodexia, and Micropoecilia sp. sec-
ond and third gonactinosts are free in adult males (state 0,
Fig. 16a). These elements are completed fused (state 1, Fig.
16b) in males of Tomeurus, Alfaro, Priapichthys, Gambusia,
Belonesox, Neoheterandria, Phallichthys, Xenophallus,
Poeciliopsis, Phalloptychus, Limia, Pamphorichthys
hollandi, Phalloceros n. sp. A, and Phalloceros n. sp. H.
Remaining taxa studied exhibit second and third gonactinosts
partially fused (state 2, Fig. 16c, e). Although this character
contributed to the resolution of the present topology, it pre-
sented several independent acquisitions and reversals dur-
ing the history of the Cyprinodontiformes.
Character 73 - Shape of gonactinostal complex
(Rauchenberger, 1989: fig. 22 C, D): (0) plate-like; (1) fused
into a column.
In most poeciliines the gonactinostal complex is expanded
in a laminar plate in the anterior-posterior plane; gonactinosts
are spread out in this plate-like spokes in a fan (state 0).
Belonesox and Gambusia possess second, third, and fourth
gonactinosts fused into a column (state 1). This is interpreted
as a synapomorphic for a clade composed by both genera.
This feature has been proposed by Rauchenberger (1989).
Character 74 - Distal portion of third and fourth gonactinosts:
(0) separate; (1) completely fused; (2) separate, except by tip
of gonactinost 3, which is arched backward towards
gonactinost 4; (3) completely fused, except by a small notch.
Members of the outgroup, Brachyrhaphis, Priapella,
Pseudopoecilia, Neoheterandria, Scolichthys, Girardinus,
Phallichthys, Quintana, Carlhubbsia, Xiphophorus,
Xenodexia, Limia, Pamphorichthys scalpridens,
Micropoecilia sp, Cnesterodon n. sp. A, Cnesterodon n. sp.
B, P. jucundus,
P. victoriae, P. fasciolatus, and almost all
species of Phalloceros exhibit the distal portion of third and
fourth gonactinosts separate in adult males (state 0, Fig. 16a).
In Alfaro, Priapichthys, Gambusia, Belonesox,
Xenophallus, Poeciliopsis, Phalloptychus, Poecilia,
Pamphorichthys hollandi, Phallotorynus n. sp. A,
Phallotorynus n. sp. B, Phalloceros n. sp. D, Phalloceros n.
sp. F, Phalloceros n. sp. H, Phalloceros n. sp. Q, and
Phalloceros n. sp. J the distal portions of third and fourth
gonactinosts are completely fused (state 1, Fig. 16b, c).
Tomeurus, “Poecilia”, Micropoecilia branneri,
Cnesterodon decemmaculatus, C. hypselurus, C.
brevirostratus, C. septentrionalis, C. omorgmatos, and C.
carnegiei possess the distal portions of third and fourth
gonactinosts separate, except by the tip of gonactinost 3,
which is arched backward towards gonactinost 4 (state 2,
Fig. 16d). Distal portions of third and fourth gonactinosts are
completely fused, except by a small notch (state 3, Fig. 16e) in
Heterandria.
Although this character contributed to the resolution of
the present topology, it presented several independent ac-
quisitions and reversals during the history of the poeciliines.
Ghedotti (2000) recognized 3 states of this character. We
add a fourth state (75-3). This author reported third and fourth
proximal anal-fin radials completely fused in Gambusia affinis,
Poeciliopsis latidens, and Heterandria formosa and partially
fused, forming an oblong opening in Phallichthys amates
and Poecilia sphenops.
Character 75 - Lateral flanges on ventral portion of fourth
gonactinost (Ghedotti, 2000: fig. 16): (0) absent; (1) present
and continuous, without dorsal cleft; (2) present and cleft
dorsally forming separate dorsally directed processes.
P. H. F. Lucinda & R. E. Reis
23
PROOFS
Fig. 16. Gonactinostal complexes. (a) Xenodexia ctenolepis, AMNH 32137; (b) Phalloptychus iheringii, MCP 11054; (c)
Phallotorynus n. sp. A, NRM 42823; (d) Cnesterodon brevirostratus, MCP 13950; (e) Heterandria jonesii UMMZ 2109990.
GNA = gonactinost; GNAC = gonactinost complex; R = ray. Scale bar 1 mm.
This character was employed by Ghedotti (2000). Most
cyprinodontiforms lack lateral flanges on ventral portion of
fourth anal-fin radial in adult males (state 0). Lateral flanges
on ventral portion of fourth gonactinost are continuous, with-
out dorsal cleft (state 1) in Gambusia and Belonesox and are
hypothesized as a synapomorphy uniting both. In Girardinus,
Phallotorynus, and Phalloceros lateral flanges are present
and cleaved dorsally forming separate dorsally directed pro-
cesses (state 2). The topology of our strict consensus tree
supports the hypothesis that state 2 appeared independently
in Girardinus and in the ancestor of Phallotorynus and
Phalloceros.
Character 76 - Posteroventral projection of ventral flange of
fourth gonactinost (Fig. 17): (0) absent; (1) large; (2) small.
Phallotorynus species are unique among
cyprinodontiforms by having a posteroventral projection of
ventral flange of fourth gonactinost (Fig. 17). The absence of
this structure in remaining cyprinodontiform taxa is consid-
ered plesiomorphic (state 0). In Phallotorynus fasciolatus
and P. victoriae this projection is large (state 1, Fig. 17a). In
Phallotorynus jucundus, Phallotorynus n. sp. A, and
Phallotorynus n. sp. B the projection is small (state 2; Fig.
17b). The presence of a posteroventral projection of ventral
flange of fourth gonactinost is interpreted as synapomorphic
for Phallotorynus with a derived size reduction in
Phallotorynus jucundus, Phallotorynus n. sp. A, and
Phallotorynus n. sp. B.
Character 77 - Fusion of posterior middle radials of anal-fin
(5
th
to last) in adult males to respective proximal radials: (0)
absent; (1) present.
Posterior median radials of anal-fin (5
th
to last one) in adult
males are independent and separate in most atherinomorphs
(state 0). Procatopus, Tomeurus, Alfaro,Xenodexia, Poecilia,
Limia, Pamphorichthys, Micropoecilia, “Poecilia”, and
Cnesterodon are unique among cyprinodontiforms by the
fusion of posterior median radials of anal-fin (5
th
to last one)
Systematics of the subfamily Poeciliinae Bonaparte
24
PROOFS
in adult males to respective proximal radials (gonactinosts of
Poeciliinae) (state 1). This derived feature is interpreted as
acquired by the ancestor of poeciliines, with a synapomorphic
reversal in clade 125 (which comprehends all poeciliines ex-
cept Tomeurus and Alfaro). Within Clade 125 state 1 is inde-
pendently acquired by Cnesterodon and by the ancestor of
Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” [Clade 104].
Character 78 - Anterior process on base of fifth middle anal-
fin radial in adult males (Fig. 18): (0) absent; (1) pointed and
upward directed; (2) hardly developed, and rounded.
The outgroup taxa, Tomeurus, Xenophallus, Poeciliopsis,
Pseudopoecilia, Phalloptychus, Carlhubbsia, Xenodexia,
Limia, Pamphorichthys, Micropoecilia, “Poecilia”, and
Phalloceros lack an anterior process on base of fifth middle
anal-fin radial in adult males (state 0, Fig. 18a). This process is
hardly developed, and round (state 2, Fig. 18c) in Cnesterodon,
Brachyrhaphis, and Belonesox. This process is pointed and
upward directed (state 1, Fig. 18b) in remaining taxa studied.
Although this character contributed to the resolution of the
present topology, it presented several independent acquisi-
tions and reversals during the history of the
Cyprinodontiformes.
Character 79 - Lateral process on base of fifth and sixth middle
anal-fin radial in adult males: (0) absent; (1) large; (2) very
large; (3) asymmetrical; (4) minute.
A lateral process on base of fifth and sixth middle anal-fin
radial in adult males is absent in the outgroup taxa and in
Tomeurus (state 0, Rosen & Bailey, 1963: fig. 15). In
Brachyrhaphis, Priapichthys, Priapella, Heterandria,
Pseudopoecilia, Scolichthys, Neoheterandria, Carlhubbsia,
Quintana, Xiphophorus, Limia, Micropoecilia, Phalloceros,
and Phallotorynus (except Phallotorynus n. sp. B) the lateral
process is large (state 1, Rosen & Bailey, 1963: fig. 29A, B)
and in Cnesterodon is even more enlarged (state 2, Rosen &
Bailey, 1963: fig. 30). In Phallichthys,
Xenophallus,
Poeciliopsis, and Phalloptychus this process is asymmetri-
cal (state 3, Fig. 19; Rosen & Bailey, 1963: fig. 56). In remain-
ing taxa studied the lateral process on base of fifth and sixth
middle anal-fin radial in adult males is minute (state 4, Rosen
& Bailey, 1963: fig. 23).
Character 80 - Asymmetry of middle anal-fin radials 5, 6, and
7 in adult males (Fig. 19) (0) absent; (1) present.
In most cyprinodontiforms middle anal-fin radials 5, 6,
and 7 are symmetrical in adult males, with right and left lateral
projections similar in shape and size (state 0). Middle anal-fin
radials 5, 6, and 7 are asymmetrical in adult males, with right
lateral projection more compressed and much larger than left
one in Phallichthys, Xenophallus, Poeciliopsis,
Phalloptychus, and Xenodexia (state 1, Fig. 19). This feature
is hypothesized as independently acquired by Xenodexia
and in the ancestor of a clade containing Phallichthys,
Xenophallus, Poeciliopsis, and Phalloptychus [Clade 113].
Fig. 19. Posterior view of sixth mesonost of Phalloptychus
iheringii, MCP 11054. LLP = left lateral process; RLP = right
lateral process; Scale bar 1 mm.
Character 81 - Gonactinost 5: (0) free; (1) fused to complex
gonactinost.
Gonactinost 5 is free in almost all poeciliines (state 0). In
Limia, “Poecilia” reticulata, and Micropoecilia gonactinost
5 is fused to gonactinostal complex (state 1). This condition
is interpreted as derived and independently acquired by the
aforementioned genera.
Character 82 - Ventral portion of proximal anal-fin radials 6 to
10 in adult males (Ghedotti, 2000: fig. 14): (0) laterally com-
pressed with anterior and posterior flanges; (1) not laterally
compressed without anterior and posterior flanges.
Most cyprinodontiforms possess the ventral portion of
proximal anal-fin radials 6-10 in adult males laterally com-
pressed, with anterior and posterior flanges (state 0). In
poeciliines the ventral portion of gonactinosts 6-10 is not
Fig. 17. Posterior view of gonactinostal complex and fifth
gonactinost of (a) Phallotorynus victoriae, NRM 42907; (b)
Phallotorynusn. sp. A, NRM 42823. GNAC = gonactinost com-
plex; GNA5 = gonactinost 5; PVP = posteroventral projection of
ventral flange. Scale bar 1 mm.
Fig. 18. Fifth mesonost of (a) Phalloptychus iheringii, MCP 11054;
(b) Cnesterodon brevirostratus, MCP 13950; (c), Phallotorynus
jucundus, UFPB 2161. AP= anterior process; LP = lateral process.
Scale bar 1 mm.
P. H. F. Lucinda & R. E. Reis
25
PROOFS
laterally compressed, without anterior and posterior flanges
(state 1), which is proposed as synapomorphic for the sub-
family Poeciliinae. This character has been discussed by
Ghedotti (2000).
Character 83 - Eighth anal-fin gonactinost (Rosen & Bailey, 1963:
fig. 53): (0) straight; (1) bearing wing-like lateral projections.
Xenophallus and Priapichthys are unique among
poeciliines by eighth gonactinost bearing wing-like lateral
projections. This feature is interpreted as apomorphic (state
1) and independently acquired by these two taxa, whereas
the absence of this structure in remaining atherinomorph
fishes is herein interpreted as plesiomorphic (state 0).
Character 84 - Ninth anal-fin gonactinost (Rosen & Bailey,
1963: fig. 53): (0) straight; (1) bearing wing-like lateral projec-
tions.
Rosen & Bailey (1963) reported the presence of wing-like
projections in Brachyrhaphis, Priapichthys, and
Neoheterandria (including Xenophallus) and employed this
character as diagnostic (in combination with other charac-
ters) for the genera above. Meyer & Etzel (1996) reported the
presence of “plate-like outgrowths” in the ninth gonactinost
of Neoheterandria tridentiger (Garman) and in the eighth
gonactinost of Xenophallus umbratilis (Meek). These au-
thors stated that the presence of wing-like lateral projections
as a “primitive character in poeciliid fishes” (Meyer & Etzel,
1996: 3), arguing that this condition is very common among
poeciliines: “because plate-like outgrowths of the
gonactinostal system are found in several taxa” (Meyer &
Etzel, 1996: 3). However, this statement is made in the ab-
sence of a cladistic analysis, which prevents the evaluation
of character polarity. Besides, the fact that a character state is
very common does not guarantee it is the plesiomorphic state.
Brachyrhaphis, Priapichthys, Neoheterandria, and
Xenophallus, are unique among poeciliines by the posses-
sion of wing-like lateral projections on the ninth gonactinost.
The presence of this structure in each of four taxa above is
interpreted as apomorphic (state 1) and homoplastic (inde-
pendently derived). On the other hand, the absence of these
wing-like lateral projections in remaining atherinomorphs is
viewed as the plesiomorphic condition (state 0).
Character 85 - Number of anal-fin rays on males: (0) ten; (1)
twelve; (2) eleven; (3) eight; (4) nine; and (5) thirteen or
more.
Ghedotti (2000) studied this character, however without
discriminating males and females. Fundulus, Scolichthys,
Xenophallus, Poeciliopsis, Quintana, Xiphophorus,
Poecilia, Limia, “Poecilia”, Micropoecilia branneri,
Phalloceros n. sp. C, Phalloceros n. sp. D, Phalloceros n.
sp. A, Phalloceros n. sp. E, Phalloceros n. sp. U, Phalloceros
n. sp. T, Phalloceros n. sp. S, Phalloceros n. sp. I, and
Phalloceros n. sp. J exhibit 10 anal-fin rays on males (state 0).
Males specimens of Cyprinodon and Alfaro possess twelve
anal-fin rays (state 1), whereas males specimens of Priapella,
Priapichthys, Heterandria, Gambusia, Belonesox,
Girardinus, Phallichthys, Carlhubbsia, and Xenodexia pos-
sess eleven anal-fin rays (state 2). Fluviphylax, Tomeurus,
Pamphorichthys hollandi, Phalloptychus iheringii,
Phallotorynus fasciolatus, and Phallotorynus victoriae
present eight anal-fin rays on males (state 3). Nine anal-fin
rays (state 4) is found in males of Pseudopoecilia,
Neoheterandria, Phalloptychus januarius, Pamphorichthys
scalpridens, Micropoecilia sp., Cnesterodon, P. jucundus,
Phallotorynus n. sp. A, Phallotorynus n. sp. B, Phalloceros
n. sp. G, Phalloceros n. sp. F, Phalloceros n. sp. B,
Phalloceros n. sp. V, Phalloceros n. sp. R, Phalloceros n. sp.
N, Phalloceros n. sp. O, Phalloceros n. sp. M, Phalloceros
n. sp. H, Phalloceros n. sp. Q, and Phalloceros n. sp. L. Male
individuals of Aplocheilichthys and Procatopus have 13 or
more anal-fin rays (state 5). Although this character contrib-
uted to the resolution of the present topology, it presented
several independent acquisitions and reversals during the
history of the poeciliines.
Character 86 - Anal-fin rays 3, 4, and 5 in adult males (R3, R4,
R5): (0) similar to remaining anal-fin rays; (1) modified into a
gonopodium.
The presence of a gonopodium formed by modified anal-
fin rays 3, 4, and 5 in adult males has long been used as
diagnostic for poeciliines (e.g. Garman, 1895; Eigenmann, 1907;
Regan, 1911, 1913; Hubbs, 1924; Parenti, 1981; Ghedotti, 2000).
Anal-fin rays 3, 4, and 5 in adult males are normally devel-
oped, i.e. are similar to remaining anal-fin rays in most
atherinomorphs (state 0). Poeciliines are unique by the pos-
session of a copulatory structure (gonopodium) formed by
modified anal-fin rays 3, 4, and 5 in adult males (state 1). This
condition is hypothesized as a uniquely derived and
unreversed synapomorphy for the subfamily.
Character 87 - Symmetry of anal fin in adult males: (0) present;
(1) absent.
Most cyprinodontiform fishes exhibit a symmetrical anal
fin in adult males (state 0). The anal fin in adult males speci-
mens is asymmetrical in Phallichthys, Xenophallus,
Poeciliopsis, and Phalloptychus,
Quintana, Carlhubbsia,
and Xenodexia (state 1). Rosen & Bailey (1959) discussed
the gonopodium asymmetry of Phalloptychus, Poeciliopsis,
Phallichthys, Xenophallus, Carlhubbsia, Quintana, and
Girardinus. They believed that characters related to
gonopodium folding were highly adaptive and may had
evolved independently more than once within the subfamily.
This feature is herein hypothesized as evolving three times
in poeciliine history: (1) in the common and exclusive ancestor
of Phallichthys, Xenophallus, Poeciliopsis, and Phalloptychus
[Clade 113]; (2) in the common and exclusive ancestor of
Quintana + Carlhubbsia [Clade 109]; (3) in Xenodexia.
Character 88 - Palp (membranous appendix) in subdistal seg-
ments of R3 (Rosen & Bailey, 1963: fig. 25; Rodriguez, 1997:
fig. 5 C-G): (0) absent; (1) present.
Systematics of the subfamily Poeciliinae Bonaparte
26
PROOFS
Most cyprinodontiforms do not exhibit a palp in subdistal
segments of R3 (state 0), whereas this structure is present
(state 1) in some poeciliids.
Rosen & Bailey (1963) used the presence of a palp in
Alfaro and Poecilia as evidence that Alfaro should be
placed in the tribe Poeciliini. Rodriguez (1997) suggested
that the presence of a palp was independently derived in
Alfaro and in the ancestor of a clade including Poecilia,
Pamphorichthys, and Limia. Ghedotti (2000) reported the
presence of a palp in Alfaro and Poecilia. Among the taxa
studied, a palp was observed in Alfaro, Poecilia, Limia,
Pamphorichthys, Micropoecilia, and “Poecilia”. Our re-
sults support the hypothesis that a palp in subdistal seg-
ments of R3 was independently derived in Alfaro and the
common and exclusive ancestor of a clade including
Poecilia, Limia, Pamphorichthys, Micropoecilia, and
“Poecilia”.
Character 89 - V-shaped ventral projection at distal por-
tion of R3 (Rosen & Bailey, 1963: fig. 31A): (0) absent; (1)
present.
Phallotorynus species are unique among
cyprinodontiforms by the possession of a V-shaped ventral
projection at distal portion of R3. This condition is hypoth-
esized as apomorphic (state 1) and exclusively shared by
Phallotorynus species.
Character 90 - Pedicel in R3 united to R4 (Rosen & Bailey,
1963: fig. 31A, B, C, D, F, G): (0) absent; (1) present.
Most cyprinodontiform fishes lack a pedicel in R3 united to
R4 (state 0). Phallotorynus, Phalloceros, and Cnesterodon
species are unique among cyprinodontiforms by the posses-
sion of a pedicel in R3 united to R4 (state 1). This condition is
hypothesized as a synapomorphy for the tribe Cnesterodontini.
Character 91 - Pedicel at tip of R3 (Rosen & Bailey, 1963: fig.
31A, B, C, D, F, G): (0) absent; (1) present.
The presence of a pedicle at tip of R3 long been reported
by different authors (Hubbs, 1924; Rosen & Bailey, 1963).
Phallotorynus, Phalloceros and Cnesterodon species are
unique among cyprinodontiforms by the possession of a
pedicel at tip of R3 (state 1). This condition is hypothesized
as a synapomorphy for the tribe Cnesterodontini.
Character 92 - Membranous appendix at tip of R3 (Rosen &
Bailey, 1963: fig. 31 A, B, C, D, F, G): (0) absent; (1) present.
The presence of a membranous appendix at tip of R3 long
been reported by different authors (Hubbs, 1924; Rosen &
Bailey, 1963; Ghedotti, 2000). Phallotorynus, Phalloceros,
and Cnesterodon species are unique among
cyprinodontiforms by the possession of a membranous ap-
pendix at tip of R3 (state 1). This condition is hypothesized as
a synapomorphy for the tribe Cnesterodontini.
Character 93 - Paired appendix at tip of R3 (Rosen & Bailey,
1963: fig. 31 C, D): (0) absent; (1) present.
Paired appendix at tip of R3 has long been reported as
diagnostic for Phalloceros (e.g. Eigenmann, 1907; Rosen &
Bailey, 1963) Phalloceros
species are unique among
cyprinodontiforms by the possession of a paired appendix at
tip of R3 (state 1). This condition is hypothesized as
synapomorphic for the genus Phalloceros.
Character 94 - Hooks on paired gonopodial appendix (at least
the right one) (Figs. 20-22): (0) absent; (1) large and sickle-
like; (2) not sickle-like.
Among Phalloceros species, Phalloceros n. sp. B,
Phalloceros n. sp. D, Phalloceros n. sp. A, Phalloceros n.
sp. C, and Phalloceros caudimaculatus lack hooks on paired
gonopodial appendix (state 0; Fig. 20). Hooks are large and
sickle-like (state 1; Fig. 21) in Phallocerosn. sp. E, Phalloceros
n. sp. G, and Phalloceros n. sp. F. Hooks are not sickle-like in
remaining species of Phalloceros (state 2; Fig. 22; Rosen &
Bailey, 1963: fig. 31 D). State 1 is herein interpreted as
synapomorphic for a clade composed of Phalloceros n. sp. E,
Phalloceros n. sp. G, and Phalloceros n. sp. F [Clade 81].
State 2 is hypothesized as synapomorphic for the clade
[Phalloceros n. sp. U + Phalloceros n. sp. T + Phalloceros n.
sp. S + Phalloceros n. sp. V + Phalloceros n. sp. B +
Phalloceros n. sp. N + Phalloceros n. sp. R + Phalloceros n.
sp. I + Phalloceros n. sp. O + Phalloceros n. sp. P +
Phalloceros n. sp. M + Phalloceros n. sp. H + Phalloceros n.
sp. Q + Phalloceros n. sp. J + Phalloceros n. sp. L] [Clade 82].
A reversal to state 0 occurred in Phalloceros n. sp. B.
Character 95 - Similarity of the halves of gonopodial paired
appendix: (0) similar to each other; (1) different from each
other; right half wider than left one.
In most species of Phalloceros the halves of gonopodial
paired appendix are similar to each other (state 0; Fig. 20, 21;
Rosen & Bailey, 1963: fig. 31 D). In Phalloceros n. sp. O,
Phalloceros n. sp. P, Phalloceros n. sp. H, Phalloceros n. sp.
Q, Phalloceros n. sp. J, and Phalloceros n. sp. L halves of
gonopodial paired appendix are different from each other;
right half is wider than left one (state 1; Fig. 22), which is
interpreted as synapomorphic for the ancestor of a clade con-
taining these species plus Phalloceros n. sp. M, with a rever-
sal in the latter.
Character 96 - Shape of the halves of gonopodial paired ap-
pendix: (0) not sickle-like, with a medium corner; (1) sickle-
like, lacking a medium corner; (2) straight and perpendicular
to R3.
Phalloceros caudimaculatus, Phalloceros n. sp. B,
Phalloceros n. sp. D, Phalloceros n. sp. A, Phalloceros n. sp.
C, Phalloceros n. sp. U, Phalloceros n. sp. T, Phalloceros n. sp.
S, Phalloceros n. sp. V, Phalloceros n. sp. N, Phalloceros n. sp.
R, Phalloceros n. sp. I, and Phalloceros n. sp. B possess halves
of gonopodial paired appendix not sickle-like, with a medium
corner (state 0; Rosen & Bailey, 1963: fig. 31 C, D). Halves are
sickle-like, lacking a medium corner (state 1; Figs. 21, 22) in
Phalloceros n. sp. E, Phalloceros n. sp. G, Phalloceros n. sp. F,
P. H. F. Lucinda & R. E. Reis
27
PROOFS
Phalloceros n. sp. O, Phalloceros n. sp. P, Phalloceros n. sp.
M, Phalloceros n. sp. H, Phalloceros n. sp. Q, and Phalloceros
n. sp. L. In Phalloceros n. sp. J halves of gonopodial paired
appendix are straight and perpendicular to R3 (Fig. 20).
State 1 is herein interpreted as synapomorphic and inde-
pendently acquired for a clade composed of Phalloceros n.
sp. E + Phalloceros n. sp. G + Phalloceros n. sp. F [Clade 81]
and for the clade [Phalloceros n. sp. O + Phalloceros n. sp. P
+ Phalloceros n. sp. M + Phalloceros n. sp. H + Phalloceros
n. sp. Q + Phalloceros n. sp. J + Phalloceros n. sp. L] [Clade
74]. State 2 is hypothesized as autapomorphic for Phalloceros
n. sp. J.
Character 97 - Hooks large directed downward and located in
the corner of gonopodial appendix: (0) absent; (1) present.
In most Phalloceros species hooks are small,
anterodorsally directed and located nearer the base of the
gonopodial appendix (Fig. 21). Phalloceros n. sp. I,
Phalloceros n. sp. R, and Phalloceros n. sp. N, are unique
among poeciliines by the possession of large hooks, down-
ward directed and located in the corner of the gonopodial
appendix (Fig. 23). This feature is hypothesized as
synapomorphic for these three species.
Character 98 - Presence of hooks on gonopodial paired ap-
pendix Character 99 - Presence of hooks on gonopodial paired
appendix: (0) absent; (1) present in both halves; (2) present
only on left half.
Among Phalloceros species, Phalloceros n. sp. B,
Phalloceros n. sp. D, Phalloceros n. sp. A, Phalloceros n.
sp. C, and Phalloceros caudimaculatus lack hooks on paired
gonopodial appendix (state 0; Fig. 20). Hooks are present in
both halves of gonopodial appendix (state 1; Figs. 21, 24) in
Phalloceros n. sp. E, Phalloceros n. sp. G, Phalloceros n.
sp. F, Phalloceros n. sp. U, Phalloceros n. sp. T, Phalloceros
n. sp. S, Phalloceros n. sp. V, Phalloceros n. sp. I,
Phalloceros n. sp. N, and Phalloceros n. sp. R. Hooks are
present only on left half (state 2; Fig. 22) in remaining spe-
cies of Phalloceros.
State 1 is herein interpreted as synapomorphic for the
clade [Phalloceros n. sp. F + Phalloceros n. sp. G +
Phalloceros n. sp. E + Phalloceros n. sp. U + Phalloceros n.
sp. T + Phalloceros n. sp. S + Phalloceros n. sp. V +
Phalloceros n. sp. B + Phalloceros n. sp. N + Phalloceros n.
sp. R + Phalloceros n. sp. I + Phalloceros n. sp. O +
Phalloceros n. sp. P + Phalloceros n. sp. M + Phalloceros n.
sp. H + Phalloceros n. sp. Q + Phalloceros n. sp. J +
Phalloceros n. sp. L] [Clade 83]. A reversal to state 0 ap-
peared in Phalloceros n. sp. B. State 2 is hypothesized as
synapomorphic for the clade [Phalloceros n. sp. O +
Phalloceros n. sp. P + Phalloceros n. sp. M + Phalloceros n.
sp. H + Phalloceros n. sp. Q + Phalloceros n. sp. J +
Phalloceros n. sp. L] [Clade 74].
Character 99 - Minute paired terminal hook on R3 (Rosen &
Bailey, 1959: fig. 7): (0) absent; (1) present.
Fig. 23. Gonopodium tip of Phalloceros n. sp. R, holotype,
MZUSP 79669. H = hook. Scale bar 1 mm.
Fig. 20. Gonopodium tip of Phalloceros n. sp. J, MCP 29270.
(a) ventrolateral view; (b) lateral view. Scale bar 1 mm.
Fig. 21. Gonopodium tip of Phalloceros n. sp. F, MCP 30572. (a)
ventrolateral view; (b) lateral view. H = hook. Scale bar 1 mm.
Fig. 22. Gonopodium tip of Phalloceros n. sp. L, MCP 20579. (a)
ventrolateral view; (b) lateral view. H = hook. Scale bar 1 mm.
Systematics of the subfamily Poeciliinae Bonaparte
28
PROOFS
Rosen & Bailey (1959) suggested that Girardinus and
Quintana were closely related and share a homologous minute
paired terminal hook on R3. However our results support that
Girardinus and Quintana are not closely related and that the
possession of a minute paired terminal hook on R3 was inde-
pendently acquired by both taxa. Thus this derived feature is
regarded as non-homologous in Girardinus and Quintana.
Character 100 - Trowel-like appendix at tip of R3 (Rosen &
Bailey, 1963: fig. 31 A, B): (0) absent; (1) present.
Rosen & Bailey (1963) reported a trowel-like appendix at
tip of R3 as synapomorphic for Phallotorynus species.
Phallotorynus is unique among cyprinodontiforms by the
possession of a trowel-like appendix at tip of R3. This con-
dition (state 1) is hypothesized as a synapomorphy for the
genus, while the absence of such structure in remaining
cyprinodontiform fishes is interpreted as plesiomorphic
(state 0).
Character 101 - Unpaired appendix at tip of R3 (Rosen & Bailey,
1963: fig. 31 F, G): (0) absent; (1) present.
Unpaired appendix at tip of R3 was suggested as a putative
synapomorphy for Cnesterodon species by Rosen & Bailey
(1963) and Rose & Costa (1993). Cnesterodon species are unique
among cyprinodontiforms by the possession of an unpaired
appendix at tip of R3. The possession of this structure (state 1)
is interpreted as synapomorphic for Cnesterodon.
Character 102 - Large membrane on pedicel of gonopodium:
(0) absent; (1) present:
Phallotorynus species are unique among cyprinodontiforms
by the possession of a large membrane on pedicel of the
gonopodium. This condition is herein hypothesized as a
synapomorphy for the genus Phallotorynus (state 1).
Character 103 - Constriction of unpaired appendix of
gonopodium: (0) absent; (1) present.
Cnesterodon decemmaculatus Cnesterodon n. sp. B, and
Cnesterodon raddai lack a constriction on unpaired appen-
dix of the gonopodium (state 0; Rosa & Costa, 1993: fig. 10);
whereas Cnesterodon n. sp. A, C. hypselurus, C.
septentrionalis, C. carnegiei, and C. omorgmatos possess
this feature (state 1; Rosa & Costa, 1993: fig. 12, 13); which is
interpreted as a synapomorphy for a clade containing these
species plus C. brevirostratus [Clade 103]. Cnesterodon
brevirostratus is polymorphic and was coded “-”.
Character 104 – Shape of the trowel-like appendix at tip of R3:
(0) absent; (1) narrow and elongate; (2) wide and short.
Phallotorynus species are unique among
cyprinodontiforms by the possession of a trowel-like appen-
dix at tip of R3. In the light of the present evidence, the ab-
sence of this appendix in cyprinodontiform fishes other than
Phallotorynus is interpreted as plesiomorphic (state 0). In
Phallotorynus victoriae, P. jucundus, and Phallotorynus n.
sp. B the trowel-like appendix is narrow and elongate (state 1;
Figs. 24, 27). In Phallotorynus n. sp. A the appendix is wide
and short (state 2; Fig. 25). The presence of a trowel-like
appendix at tip of R3 is interpreted as synapomorphic for
Phallotorynus with a derived widening and reduction in length
in
Phallotorynus n. sp. A. Phallotorynus fasciolatus was
coded “?” for character state could not be checked due to
poor condition of the material studied.
Character 105 - Lateral processes on the trowel-like appendix:
(0) small; (1) large.
Phallotorynus victoriae, Phallotorynus n. sp. A, and
Phallotorynus n. sp. B possess small lateral processes on
the trowel-like appendix (state 0; Figs. 25, 27). Phallotorynus
jucundus presents large lateral processes (state 1; Fig. 24),
and this feature is interpreted as autapomorphic for P.
jucundus. Phallotorynus fasciolatus was coded “?” for char-
acter state could not be checked due to poor condition of the
material studied.
Character 106 - Profile of lateral border of left and right halves
of trowel-like appendix: (0) straight; (1) concave.
Phallotorynus victoriae, Phallotorynus jucundus, and
Phallotorynus n. sp. B possess the profile of lateral border of
left and right halves of trowel-like appendix straight (state 0;
Figs. 24, 27). Phallotorynus n. sp. A presents a concave pro-
file (state 1; Fig. 25), and this feature is interpreted as
autapomorphic for Phallotorynus n. sp. A. Phallotorynus
fasciolatus was coded “?” for character state could not be
checked due to poor condition of the material studied.
Character 107 - Level of separation/union of left and right
halves of the trowel-like appendix: (0) halves separate by a
gap along two thirds of its extension; (1) halves united along
its whole extension; (2) halves separate along approximately
five sixths of its whole extension.
In Phallotorynus victoriae and Phallotorynus n. sp. B the
halves of trowel-like appendix are separate by a gap along two
thirds of its extension (state 0; Fig. 26). In Phallotorynus
jucundus halves are united along its full extension (state 1; Fig.
24). Phallotorynus n. sp. A presents halves separate along ap-
proximately five sixths of its whole extension (state 2; Fig. 25).
Phallotorynus species are unique among cyprinodontiforms by
the possession of a trowel-like appendix at tip of R3. States 1 and
2 are interpreted as autapomorphic for Phallotorynus jucundus
and Phallotorynus n. sp. A, respectively.
Character 108 - Spines on subdistal segments of R3: (0) ab-
sent; (1) square and antrorse or directed downwards; (2) ret-
rorse.
Among studied taxa, Tomeurus, Alfaro, Brachyrhaphis,
Priapichthys, Priapella, Heterandria, Belonesox, and
Xenodexia exhibit squared, antrorse or downward directed
spines on subdistal segments of R3 (state 1, Rosen & Bailey,
1963: fig. 25 A, C, H). In Xiphophorus, “Poecilia”, and
Micropoecilia these spines are retrorse (state 2, Rosen &
Bailey, 1963: fig. 25 B, D). In remaining studied taxa these
P. H. F. Lucinda & R. E. Reis
29
PROOFS
structures are absent (state 0, Rosen & Bailey, 1963: fig. 25 F).
Our results support state 1 as a synapomorphy for Poeciliinae
with a reversal to state 0 in the base of clade 119. Within this
clade a change 0 > 2 occurs at node 108; a reversal to state 1
appeared in Xenodexia, and two independent reversals took
place in both Limia and Pamphorichthys.
Character 109 - Lateral wings on segments of R3: (0) absent;
(1) symmetrical; (2) asymmetrical.
Carlhubbsia, Xiphophorus, Xenodexia, Poecilia, Limia,
Pamphorichthys, Micropoecilia, and “Poecilia” are unique
among poeciliines by the possession of lateral wings on seg-
ments of R3. Wings are asymmetrical in Carlhubbsia (state
2; Rosen & Bailey, 1959: fig. 3D, 10). Xiphophorus,Xenodexia,
Poecilia, Limia, Pamphorichthys, Micropoecilia, and
“Poecilia” possess symmetrical lateral wings on segments
of R3 (state 1, Rosen & Bailey, 1963: fig. 25B). According to
the present hypothesis of relationships State 2 is hypoth-
esized as autapomorphic for Carlhubbsia, whereas state 2 is
synapomorphic for a clade composed of Xiphophorus,
Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” [Clade 108].
Character 110 - Membranous tip anterior to R4 and R5: (0)
absent; (1) curved downwards; (2) straight.
Most cyprinodontiforms lack a membranous tip anterior
to R4 and R5 (state 0). Heterandria, Limia, Cnesterodon spe-
cies except C. brevirostratus, Phallotorynus and Phalloceros
are unique among poeciliines by the presence of a membra-
nous tip anterior to R4 and R5. This membranous tip is curved
downwards (state 1, Rosen & Bailey, 1963: fig. 31 C, D, F, G) in
Heterandria, Limia, Cnesterodon species except C.
brevirostratus, and in Phalloceros. In Phallotorynus it is
straight (state 2, Rosen & Bailey, 1963: fig. 31 A). According
to the present hypothesis state 1 has appeared independently
in Heterandria, Limia, and in the ancestor of Cnesterodon,
Phallotorynus and Phalloceros Clade 111], with a reversal in
C. decemmaculatus. State 2 is interpreted as synapomorphic
for Phallotorynus.
Character 111 - Size of membranous tip anterior to R4 and R5:
(0) absent; (1) small; (2) reduced; (3) large.
Most cyprinodontiform fishes lack a membranous tip an-
terior to R4 and R5 (state 0, Rosen & Bailey, 1963: fig. 39).
Heterandria, Limia, Cnesterodon except C. decemmaculatus,
Phallotorynus jucundus, and P. victoriae possess a small
membranous tip anterior to R4 and R5 (state 1, Rosen & Bailey,
1963: fig. 31 F, G). This membrane is very reduced in
Phallotorynus n. sp. A, and Phallotorynus n. sp. B (state 2,
Rosen & Bailey, 1963: fig. 31 A) and enlarged in Phalloceros
species (state 3, Rosen & Bailey, 1963: fig. 31 D).
The present phylogenetic study support the hypothesis
that state 1 was independently acquired by Heterandria,
Limia, and by the ancestor of Cnesterodon, Phallotorynus,
and Phalloceros, with a reversal to state 0 in C.
decemmaculatus. State 2 is interpreted as synapomorphic for
Phallotorynus n. sp. A, and Phallotorynus n. sp. B, whereas
state 3 is interpreted as synapomorphic for Phalloceros spe-
cies with a reversal in Phalloceros n. sp. D.
Fig. 24. Gonopodium appendix of Phallotorynus jucundus,
MCP 25415, (a) lateral view; (b) dorsal view. LP = lateral
process; LH = left half; RH = right half. Scale bar 1 mm.
Fig. 25. Gonopodium appendix of Phallotorynus n. sp. A,
NRM 42823, (a) lateral view; (b) dorsal view. LH = left half;
RH = right half.Scale bar 1 mm.
Fig. 26. Gonopodium appendix of Phallotorynus victoriae,
NRM 42907, (a) lateral view; (b) dorsal view. LP = lateral pro-
cess; LH = left half; RH = right half. Scale bar 1 mm.
Systematics of the subfamily Poeciliinae Bonaparte
30
PROOFS
Character 112 - R5a, R5p, R4p, R4a directed upwards
(Rauchenberger, 1989: fig. 23): (0) absent; (1) present.
Rauchenberger (1989) reported R5a, R5p, R4p, R4a directed
upwards as a synapomorphy uniting Brachyrhaphis, Gam-
busia, and Belonesox. However, our phylogenetic framework
support this derived feature as independently acquired by
Brachyrhaphis and by the ancestor of Gambusia and
Belonesox.
Character 113 - Ventral projection of R4a towards R3
(Rauchenberger, 1989: fig. 20; 38): (0) absent; (1) present.
Rauchenberger (1989) reported a ventral projection of R4a
towards R3 as an autapomorphy for Gambusia. However,
following the present phylogeny, this derived feature is also
present and is hypothesized as synapomorphic for Scolichthys
+ Neoheterandria [Clade 114]. The presence of this charac-
ter state in Gambusia is interpreted as homoplastic.
Character 114 - Number of series of subdistal retrorse spines
on R4p: (0) zero; (1) one; (2) two.
Members of the outgroup, Tomeurus, Alfaro, Xenophallus,
and Xenodexia lack subdistal retrorse spines on R4p (state
0). Almost all remaining studied taxa possess at least one
series of subdistal retrorse spines on R4p. Xiphophorus and
Pamphorichthys scalpridens exhibit two series. State 1 is
herein proposed as synapomorphic for a clade comprising all
poeciliines except Alfaro and Tomeurus [Clade 125]. Two sub-
sequent independent reversals took place: (1) in Xenophallus
and (2) in Xenodexia. The presence of two series of subdistal
retrorse spines on R4p in Xiphophorus and Pamphorichthys
scalpridens is interpreted as homoplastic.
Character 115 - Depth of distal segments of R4p posterior to
serrae (Rodriguez, 1997; Fig. 5f, g): (0) wider than deep; (1)
deeper than wide.
In most poeciliines distal segments of R4p posterior to
serrae are wider than deep (state 0). Rodriguez (1997) pro-
posed distal segments of R4p posterior to serrae deeper than
wide (state 1) as a synapomorphy uniting Limia and
Pamphorichthys. This was confirmed herein. Additionally,
this feature was also observed to be present in Xenodexia.
Our results support the assumption of this derived condition
as synapomorphic for a clade containing Xenodexia,
Poecilia, Limia, Pamphorichthys, Micropoecilia, and
“Poecilia” Clade [104], with two reversals: (1) in Poecilia
and (2) in Micropoecilia, and “Poecilia” clade [Clade 87].
Character 116 - Number of subdistal retrorse spines on R4p:
(0) zero; (1) eight or more; (2) four to seven.
Among studied taxa, members of the outgroup, Alfaro,
Tomeurus
, Xenodexia, and Xenophallus lack subdistal ret-
rorse spines on R4p (state 0). Gambusia, Belonesox,
Carlhubbsia, Micropoecilia, “Poecilia”, Cnesterodon, and
Phallotorynus possess four to seven subdistal retrorse
spines on R4p (state 2). Remaining studied taxa exhibit eight
or more spines (state 1). Global parsimony of character states
support the following sequence of evolutive suppositions:
(1) state 1 is synapomorphic for Clade 125; (2) state 2 ap-
peared independently in: (a) the ancestor of Gambusia and
Belonesox; (b) the ancestor of Cnesterodon, Phalloceros
and Phallotorynus, with a reversal to state 1 in Phalloceros;
(c) the ancestor of “Poecilia” and Micropoecilia; and (d) in
Carlhubbsia; (3) reversals to state 0 occurred in Xenodexia,
and Xenophallus.
Character 117 - Short, dorsal protuberance close to base of
R4p: (0) absent; (1) present.
Phalloptychus species are unique among
cyprinodontiforms by the possession of a short, dorsal pro-
tuberance close to base of R4p (Fig. 27). Our phylogenetic
analysis supports the hypothesis that this feature represents
a synapomorphy for Phalloptychus species (state 1).
Character 118 - Elongate, dorsal protuberance just behind
retrorse spines series of R4p (Rosen, 1979: fig. 6, 26): (0) ab-
sent; (1) present.
An elongate, dorsal protuberance just behind retrorse
spines series of R4p (state 1, Rosen, 1979: fig. 6, 26) in present
in Priapella, Heterandria, Girardinus, Phallichthys,
Quintana, and Phalloceros. Remaining cyprinodontiforms
lack this structure. The phylogenetic analysis supports as-
suming the absence of such protuberance as plesiomorphic
(state 0) and its presence as apomorphic (state 1). The pres-
ence of this protuberance is herein interpreted as
synapomorphic for the members of Clade 123, with subse-
quent reversals at nodes 120, 110 and 115. Within Clade 115,
Quintana and Phalloceros independently reacquired state 1
condition.
Fig. 27. Base of gonopodium of Phalloptychus iheringii,
MCP 11054. DP = dorsal protuberance. Scale bar 1 mm.
P. H. F. Lucinda & R. E. Reis
31
PROOFS
Character 119 - Keel on posterior ventral surface of R5 formed
by the projection of R5 toward R4p (Rodriguez, 1997: fig. 5 F):
(0) absent; (1) present.
Rodriguez (1997) reported a keel on posterior ventral sur-
face of R5 formed by the projection of R5 toward R4p as
synapomorphic for a clade comprising Poecilia,
Pamphorichthys, and Limia. We observed this keel in these
taxa as well as in Xenodexia. Our phylogenetic framework
supports the absence of this keel in most cyprinodontiforms
as plesiomorphic (state 0), and its presence as apomorphic
(state 1). Thus, the presence of such keel is interpreted as a
synapomorphy for a clade embracing Xenodexia, Poecilia,
Limia, Pamphorichthys, Micropoecilia, and “Poecilia”
[Clade 104] with a reversal in Micropoecilia + “Poecilia”
clade.
Character 120 - R5 (Rodriguez, 1997: fig. 5F): (0) not bending
into R4; (1) bending into R4.
Rodriguez (1997) stated that Limia is unique among
poeciliines by the R5 bending into R4, which was confirmed
by our observations. Therefore it is herein considered a
synapomorphy for Limia species.
Character 121 - Groove dorsal to R5: (0) narrow or absent; (1)
present and wide.
Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” share the presence of a wide
groove dorsal to R5. Our phylogenetic framework supports
the absence of this groove in most cyprinodontiforms as
plesiomorphic (state 0), and its presence as apomorphic (state
1). Thus, the presence of a groove dorsal to R5 is interpreted
as a uniquely derived and unreversed synapomorphy for a
clade containing Xenodexia, Poecilia, Limia,
Pamphorichthys, Micropoecilia, and “Poecilia” [Clade 104]
in the light of present evidence.
Character 122 - Distal segment at tip of R5a: (0) normal; (1)
transformed in retrorse triangular spine; (2) hook-like.
Rosa & Costa (1993) suggested the distal segment at tip
of R5a modified into a retrorse triangular spine as a putative
synapomorphy for Cnesterodon species, whereas
Rauchenberger (1989) recognized a hook on tip of R5a as a
synapomorphy shared by Gambusia and Belonesox. In most
cyprinodontiforms, the distal segment at tip of R5a is similar
in shape to remaining segments (state 0). Cnesterodon spe-
cies are unique among poeciliines by having the distal seg-
ment at tip of R5a modified into a retrorse triangular spine
(state 1; Ghedotti, 2000: fig. 15A), which is herein hypoth-
esized as synapomorphic for this genus. In Gambusia and
Xiphophorus this segment is modified into a hook (state 2;
Rauchenberger, 1989: fig. 20). However, following our hypoth-
esis of relationships, these derived features are interpreted
as independently acquired in Gambusia
and Xiphophorus.
Character 123 - Hook on R5a contacting the segments of R4p
(Rauchenberger, 1989: fig. 20, 23): (0) absent; (1) present.
Rauchenberger (1989) proposed the presence a hook on
R5a contacting the segments of 4p as a synapomorphy unit-
ing Gambusia and Belonesox. We have confirmed this and
also observed this derived feature in Xiphophorus. Thus, the
presence of a hook on 5a contacting the segments of R4p is
interpreted as independent acquisitions by Xiphophorus and
by the ancestor of Gambusia and Belonesox.
Character 124 - Dorsal expansion of R5p (Fig. 28): (0) absent;
(1) present.
Rosa & Costa (1993) suggested the dorsal expansion of
R5p as a putative synapomorphy for Cnesterodon species.
However this feature is shared by Tomeurus, Heterandria,
Pseudopoecilia, Neoheterandria, Scolichthys, Xenodexia,
and Cnesterodon (Fig. 28). This structure is absent in re-
maining cyprinodontiform fishes. Our phylogenetic frame-
work supports the absence of this expansion in most
cyprinodontiforms as plesiomorphic (state 0), and its pres-
ence as apomorphic (state 1). According to the present hy-
pothesis, this derived feature is independently acquired by
Tomeurus, Heterandria, the ancestor of [Pseudopoecilia +
Neoheterandria + Scolichthys], Xenodexia and
Cnesterodon.
Character 125 - Serrae on R5p: (0) absent; (1) present.
Most cyprinodontiforms lack serrae on R5p (state 0).
Girardinus, Quintana, and Carlhubbsia are unique among
poeciliines by the possession of serrae on R5p (state 1). This
derived feature is interpreted as independent acquisitions by
Girardinus and by the ancestor of Quintana and Carlhubbsia.
Character 126 - Degree of fusion between lower and upper
branches of sixth anal-fin ray in adult males (R6a and R6p): (0)
absent; (1) partial; (2) total.
Fig. 28. Gonopodium of Cnesterodon n. sp. A, paratype, MZUSP 54978. Scale bar 1 mm.
Systematics of the subfamily Poeciliinae Bonaparte
32
PROOFS
In members of the outgroup, Alfaro, Brachyrhaphis,
Pseudopoecilia, Pamphorichthys, Micropoecilia, and “
Poecilia” R6a and R6p are free from each other (state 0).
Remaining poeciliines present various degrees of partial fu-
sion between these elements (state 1). These branches are
totally fused in Poeciliopsis and Phalloptychus (state 2).
Our results allow the assumption that state 1 appeared inde-
pendently in Tomeurus and in the ancestor of members of the
Clade 124, with two subsequent reversals: (1) in
Pseudopoecilia and (2) in the node Pamphorichthys +
Micropoecilia + “Poecilia” [Clade 92]. State 2 is interpreted
as synapomorphic for Poeciliopsis and Phalloptychus [Clade
105].
Character 127 - Degree of fusion between more distal ele-
ments of branches of sixth anal-fin ray in adult males (R6):
(0) not fused; segmented; (1) partially fused; (2) totally
fused.
In members of the outgroup, Alfaro, Brachyrhaphis,
Belonesox, Pseudopoecilia, Pamphorichthys,
Micropoecilia, and “Poecilia” the more distal elements of
R6 branches are not fused (state 0). In males of Tomeurus,
Priapella, Priapichthys, Heterandria, Gambusia,
Phallichthys, Quintana, Carlhubbsia, Xiphophorus,
Poecilia, Limia, Phallotorynus, and some species of
Phalloceros more distal elements of R6 branches are partially
fused (state 1). These elements are totally fused (state 2) in
remaining taxa studied. Although this character contributed
to the resolution of the present topology, it presented several
independent acquisitions and reversals during the history of
the Cyprinodontiformes.
Character 128 - Distal portion of R6: (0) not expanded; (1)
expanded.
The distal portion of R6 of most cyprinodontiform fishes
is not expanded (state 0). Neoheterandria, Phallichthys,
Carlhubbsia, Cnesterodon, and Phallotorynus are unique
among poeciliines by the possession of an expanded distal
portion of R6 (state 1; Ghedotti, 2000: fig. 14D). Following our
results, this feature is interpreted as independently acquired
in all genera above.
Character 129 - Size of lower branch of R6: (0) longer than
upper branch; (1) as long as upper branch.
In most cyprinodontiforms the lower branch of R6 is longer
than the upper branch (state 0). In Phalloptychus species the
lower branch is as long as the upper (state 1), which is as-
sumed as synapomorphic for Phalloptychus species
Character 130 - Distal portion of R6 and seventh anal-fin ray
in adult males (R7): (0) not fused; (1) fused.
In most cyprinodontiforms distal portion of R6 and R7 are
not free (state 0). In Phallotorynus n. sp. A and Phallotorynus
n. sp. B distal portion of R6 and R7 are fused to each other
(state 1). This is hypothesized as synapomorphic for a clade
containing these two species.
Caudal Fin
Character 131 - Hypural plate: (0) completely fused; (1) par-
tially fused with an elongate aperture; (2) bipartite; (3) almost
bipartite, very large aperture.
Ghedotti (2000) employed this character recognizing the
states 0 and 2 above. Ghedotti (2000) reported a bipartite
hypural plate in Gambusia, Alfaro, Poecilia, and
Phallichthys. Hypural plate is completely fused (state 0) in
Fluviphylax, Cyprinodon, Fundulus, Brachyrhaphis,
Priapella, Pseudopoecilia, Scolichthys, Xenodexia, Limia,
Pamphorichthys hollandi, and Cnesterodon. In Procatopus,
Gambusia, Girardinus, Heterandria, Carlhubbsia,
Xiphophorus, Micropoecilia, Phallotorynus, and
Phalloceros (except Phalloceros n. sp. G) hypural plate is
partially fused with an elongate aperture (state 1). A bipartite
hypural plate (state 2) is present in Belonesox, Xenophallus,
Poeciliopsis, Phalloptychus, Poecilia, and Pamphorichthys
scalpridens. Phalloceros n. sp. G possesses a hypural plate
almost bipartite, with a very large aperture (state 3). Although
this character contributed to the resolution of the present
topology, it presented several independent acquisitions and
reversals during the history of the Cyprinodontiformes.
Character 132 - Number of caudal-fin rays in contact with the
hypural plate: (0) less than nine; (1) nine or more.
In Aplocheilichthys, Fluviphylax, Procatopus,
Tomeurus,
Brachyrhaphis, Belonesox, Gambusia, Xenophallus,
Phalloptychus eigenmanni, Pamphorichthys, Micropoecilia,
“Poecilia”, Cnesterodon septentrionalis, Phallotorynus,
and Phalloceros (except Phalloceros n. sp. G and Phalloceros
n. sp. I) the number of caudal-fin rays in contact with the
hypural plate is less than nine (state 0). In remaining taxa
studied there is nine or more (state 1) caudal-fin rays in con-
tact with the hypural plate. Although this character contrib-
uted to the resolution of the present topology, it presented
several independent acquisitions and reversals during the
history of the Cyprinodontiformes.
Pigmentation
Character 133 - Elongate vertical bars on lateral surface of
body (Fig. 29): (0) absent; (1) present.
Poeciliopsis and Quintana are unique among poeciliines
by the possession of elongate vertical bars reaching dorsal
and ventral profiles plus short bars peduncle caudal. Our
results support the hypothesis that this derived feature is
independently acquired by Girardinus and by the ancestor
of Quintana and Poeciliopsis.
Character 134 - Spot on median region of flank: (0) absent; (1)
rounded; (2) elliptical; (3) squared; (4) elongate forming a bar
vertical reaching dorsal and ventral profiles; (5) typically
densely pigmented rectangle-like lateral spot located on the
14th or 15th (very rarely 16th) scale of longitudinal line.
Most cyprinodontiforms lack a spot on median region of
flank (state 0). Such a spot is present in some individuals of
Neoheterandria (coded “-”), Scolichthys, and Phalloceros
(except Phalloceros n. sp. A and Phalloceros n. sp. G). This
P. H. F. Lucinda & R. E. Reis
33
PROOFS
spot is round (state 1, Fig. 30a) in Phalloceros n. sp. B,
Phalloceros n. sp. C and Phalloceros n. sp. D, and elliptical
in most species of Phalloceros (state 2, Fig. 30b). An ellipti-
cal spot appeared independently in Phalloceros and in the
clade Neoheterandria + Scolichthys.
Phalloceros n. sp. P possesses an autapomorphic-squared
spot (state 3, Fig. 30c). In Phalloceros n. sp. H the spot is
elongate forming a bar vertical reaching dorsal and ventral pro-
files (state 4, Fig. 30d) and it is autapomorphic for this species.
Phalloceros n. sp. F also possesses an autapomorphic spot,
which is typically densely pigmented rectangle-like lateral spot
more anterior located (on the 14
th
or 15
th
, very rarely 16
th
, scale
of longitudinal line) (state 5; Fig. 30e).
Character 135 - Number of round to elliptical dark blotch along
ventral half of flank: (0) zero [absent] (Fig. 31a); (1) one (Fig.
31b); (2) two (Fig. 31c); (3) four or more (Fig. 31d).
Phallotorynusspecies, with the exception of P. fasciolatus
are unique among cyprinodontiforms by the possession of
round to elliptical dark blotches along ventral half of flanks.
Therefore, this feature is interpreted as synapomorphic for a
clade comprising Phallotorynus jucundus, Phallotorynus
victoriae, Phallotorynus n. sp. A, and Phallotorynus n. sp.
B. Phallotorynus jucundus exhibits four to seven blotches,
and it is considered as an autapomorphy. Phallotorynus n.
sp. A presents two blotches, and this state is considered
autapomorphic for this species.
Character 136 - Pigmentation of dorsal fin: (0) slightly pig-
mented with black; (1) moderately pigmented with black; (2)
densely pigmented with black.
Most cyprinodontiforms possess the dorsal fin slightly
pigmented with black (state 0, Fig. 32a). In Cyprinodon,
Brachyrhaphis, Poecilia, Xenophallus, Micropoecilia, and
Phallotorynus dorsal fin is moderately pigmented with black
(state 1, Fig. 32b), and this feature seems to be independently
acquired in each of these genera. An autapomorphic dorsal
fin densely pigmented with black (state 2, Fig. 32c) is present
in Phallotorynus jucundus.
Fig. 30. (a) Phalloceros n. sp. B, female, MCP 30548, 24.4
mm SL; (b) Phalloceros n. sp. Q, female, MCP 31141, 29.9
mm SL; (c) Phalloceros n. sp. P, female, MCP 30511, 31.8
mm SL; (d) Phalloceros n. sp. H, female, MCP 12603, 41.3
mm SL; (e) Phalloceros n. sp. F, female, MNRJ 22509, 33.7
mm SL.
Character 137 - Dark stripe on median portion of dorsal fin
(Fig. 32a): (0) absent; (1) present.
Most cyprinodontiforms lack a dark stripe on median
portion of dorsal fin (state 0, Figs. 29, 34). Such a stripe is
present in Brachyrhaphis, Priapichthys, Heterandria, Gam-
busia, Pseudopoecilia, Neoheterandria, Scolichthys,
Xiphophorus, Poecilia, Limia, Pamphorichthys,
Micropoecilia, “Poecilia”, Phallotorynus, and
Phalloceros (state 1, Fig. 32a). Our results support the hy-
pothesis that the derived condition appeared at the ances-
tor of Clade 125, with subsequent reversals in Belonesox,
Priapella, Xenodexia and re-acquisitions in Clades 112, 106
and Quintana.
Fig. 29. Phalloptychus iheringii. (a), male, 18.35 mm SL. (b),
female, 25.67 mm SL.
Systematics of the subfamily Poeciliinae Bonaparte
34
PROOFS
Fig. 31. (a) Phallotorynus fasciolatus. MZUSP 41373, female,
28.5 mm SL; (b) Phallotorynus n. sp. B. NRM 33530, female,
25.25 mm SL; (c) Phallotorynus n. sp. A, female, NRM 41848,
18.1 mm SL; (d) Phallotorynus jucundus, Female, MCP 25415,
21.9 mm SL.
Character 138 - Dark patch of pigmentation along R3: (0) ab-
sent; (1) present.
Most cyprinodontiforms lack a dark patch of pigmenta-
tion along R3 (state 0, Fig. 34). Scolichthys, Girardinus,
Phallichthys, Xenophallus, Phalloptychus, Xiphophorus,
Phallotorynus, and Phalloceros exhibit a dark patch of pig-
mentation along R3 (state 1, Fig. 33). According to the present
analysis, this feature evolved independently in (1) Scolichthys,
(2) Xiphophorus, (3) in the ancestor of Phallotorynus and
Phalloceros; and (4) in the ancestor of Girardinus,
Phallichthys, Xenophallus, Phalloptychus, and Poeciliopsis
[Clade 116], with a reversal in Poeciliopsis.
Character 139 - Dark spot posterior to anal-fin base of males
continuous ventrally side by side and continuous with ven-
tral median line of caudal peduncle (Fig. 34): (0) absent; (1)
present.
Rosa & Costa (1993) suggested that the presence of a
dark spot posterior to anal-fin base of males continuous
ventrally side by side and continuous with ventral median
line of caudal peduncle could be a synapomorphy for the
species of Cnesterodon. This is confirmed by our phyloge-
netic study.
Fig. 32. (a) Phalloceros n. sp. Q, male, 26.4 mm SL, MCP
31142; (b) Female, MCP 31141, 29.9 mm SL; (c)
Phallotorynus n. sp. B, male, 15.9 mm SL. MNHNP 4621;
(d) Female, NRM 33530, 25.2 mm SL; (e) Phallotorynus
jucundus, male, MCP 30467, 18.4 mm SL; (f) Female, MCP
25415, 21.9 mm SL.
Character 140 - Ground pigmentation of anal fin of females:
(0) slightly speckled with black, or hyaline, not forming a
distinct stripe on first rays; (1) moderately speckled with
black, with chromatophores more concentrated anteriorly
and forming a dark stripe on first rays; (2) black.
P. H. F. Lucinda & R. E. Reis
35
PROOFS
Fig. 34. Cnesterodon brevirostratus. MCP 26050. (a), male,
25.2 mm SL. (b), female, 33.7 mm SL.
Most female cyprinodontiforms possess anal fin slightly
speckled with black, or hyaline, not forming a distinct stripe
on first rays (state 0, Fig. 32a). Females of Phallotorynus
species are unique among cyprinodontiforms by the pos-
session of a anal fin moderately to densely speckled with
black, with chromatophores more concentrated anteriorly
and forming a dark stripe on first rays (states 1 and 2, Fig.
32b,c). This condition is herein interpreted as synapomorphic
for the genus and the possession of a black anal fin (state 2,
Fig. 32c) is considered as a derived autapomorphy for
Phallotorynus jucundus.
Miscellaneous
Character 141 - Viviparity: (0) absent; (1) present.
Viviparity among cyprinodontiform fishes has long been
discussed (e.g. Rosen & Bailey, 1963; Parenti, 1981; Meyer &
Lydeard, 1993; Ghedotti, 2000). Among cyprinodontiforms
viviparity evolved independently in the Goodeidae,
Anablepidae, and Poeciliinae. Tomeurus was coded “-” be-
cause it exhibits facultative viviparity. Since Tomeurus is the
most basal poeciliine according to the present phylogenetic
hypothesis, the facultative viviparity could be viewed in two
different ways: (1) as a preliminary “testing” stage of vivipar-
ity towards “true viviparity” which was achieved by the an-
cestor of remaining poeciliines; or (2) as an autapomorphic
specialized condition of viviparity adaptable for different en-
vironmental conditions.
Character 142 - Position of urogenital papilla of females: (0)
along of median ventral line; (1) turned to the right; (2) turned
to the left.
Females of most cyprinodontiforms possess the urogeni-
tal papilla on the median ventral line (state 0). Urogenital pa-
pilla of females is turned to the right (state 1) and is inter-
preted as synapomorphic for [Phalloceros n. sp. N +
Phalloceros n. sp. R + Phalloceros n. sp. I + Phalloceros n.
sp. O + Phalloceros n. sp. P + Phalloceros n. sp. M +
Phalloceros n. sp. H + Phalloceros n. sp. Q + Phalloceros n.
sp. J + Phalloceros n. sp. L] [Clade 76]. Urogenital papilla of
females is turned to the left (state 2) and is interpreted as
synapomorphic for [Phalloceros n. sp. E + Phalloceros n. sp.
G + Phalloceros n. sp. F] [Clade 81].
Character 143 - Orbital bones: (0) absent; (1) anterior and
posterior; (2) anterior only.
In Fluviphylax, Fundulus, Jenynsia, Brachyrhaphis,
Tomeurus, Phalloptychus, Xenodexia, and Cnesterodon (except
Cnesterodon n. sp. A) orbital osseous plates are absent (state
0). Pseudopoecilia and Quintana exhibit one anterior orbital
bones (state 2). Remaining taxa studied possess two orbital os-
seous plates (anterior and posterior ones) (state 1). Although
this character contributed to the resolution of the present topol-
ogy, it presented several independent acquisitions and rever-
sals during the history of the Cyprinodontiformes.
Phylogenetic reconstruction and synapomorphy list
The phylogenetic analysis yielded 96 equally most parsi-
monious trees with length (L) = 758 steps (including TSA in
the outgroup), consistency index (CI) = 0.35, and retention
index (RI) = 0.75. A strict consensus tree is shown in Figs. 1,
2, and 3. Synapomorphy list is presented in the “Taxonomic
Account” section and in the Appendix III. Fits of individual
characters are summarized in Appendix IV.
Taxonomic Account
The current phylogenetic study supports the proposal of
a new classification for the subfamily Poeciliinae. These modi-
fications are necessary in order to make groups natural (mono-
phyletic). Diagnoses are provided for suprageneric clades.
Nevertheless, diagnoses for monotypic tribes are only pre-
liminary because this study focused on the search for de-
rived features uniting genera rather than on generic
autapomorphies. Thus, generic diagnoses were not fully sur-
veyed during this study. Oncoming studies will possibly re-
veal several other diagnostic features for these monotypic
tribes. Some authors (e.g. Regan, 1913; Hubbs, 1924; Rosen,
Fig. 33. Phalloceros n. sp. U. (a), male, 19.9 mm SL, MCP
30468. (b), female, MCP 30023, 31.5 mm SL.
Systematics of the subfamily Poeciliinae Bonaparte
36
PROOFS
1952; Rosen, 1979; Rosen & Bailey, 1963; Rauchenberger,
1989; Rodriguez, 1997) already provided some insight on di-
agnostic characters for some of these tribes but not neces-
sarily on the light of a phylogenetic framework. Rather than
writing down the diagnoses and discussions of these mono-
typic clades, we refer the reader to the articles above for a
detailed study.
Thus, the following classification is proposed (summa-
rized in Table 3):
Table 3. Proposed Classification of Poeciliinae
Subfamily Poeciliinae Bonaparte, 1831
Tribe Tomeurini Eigenmann, 1912
Tomeurus
Tribe Alfarini Hubbs, 1924
Alfaro
Tribe Brachyrhaphini, new
Brachyrhaphis
Tribe Priapichthyini, new
Priapichthys
Tribe Priapellini Ghedotti, 2000
Priapella
Tribe Heterandriini Hubbs, 1924
Heterandria
Tribe Gambusiini Gill, 1893
Gambusia, Belonesox, Neoheterandria, Scolichthys,
Pseudopoecilia
Supertribe Poeciliini Bonaparte, 1831
Tribe Poeciliini Bonaparte, 1831
Poecilia, Quintana, Carlhubbsia, Limia, Xiphophorus,
Xenodexia, Pamphorichthys, Micropoecilia
Tribe Girardinini Hubbs, 1924
Girardinus, Poeciliopsis, Xenophallus, Phalloptychus,
Phallichthys
Tribe Cnesterodontini Hubbs, 1924
Cnesterodon, Phalloceros, Phallotorynus
Subfamily Poeciliinae Bonaparte
[Clade 129]
Poecilini Bonaparte, 1831: 94, unavailable name; preoccupied
in Coleoptera.
Poeciliini Bonaparte, 1831. Type-genus: Poecilia Bloch &
Schneider, 1801.
Diagnosis. Poeciliines species share the following uniquely
derived and unreversed features: (1) ventral portion of proxi-
mal anal-fin radials 6 to 10 in adult males not laterally com-
pressed without anterior and posterior flanges [82-1**] (a
condition found in other members of the Superfamily
Poeciloidea, except for Fluviphylax); and (2) anal-fin rays 3,
4, and 5 in adult males modified in copulatory structure [86-
1**].
Additionally, poeciliines can be diagnosed by the follow-
ing not uniquely derived and/or reversed features: (1) six
branchiostegal rays [22-1]; (2) anterior process of anterior
ceratohyal not extending ventral to ventral hypohyal [24-1*];
(3) females with 10 anal-fin rays [65-1]; (4) second, third, and
fourth gonactinosts into a gonactinost complex [67-1*]; (5)
complete fusion of second and third gonactinosts [72-1]; fu-
sion of anal-fin posterior median radials (5
th
to last one) in adult
males to respective proximal radials [77-1]; (6) twelve anal-fin
rays of males [85-1*]; (7) squared and antrorse spines on
subdistal segments of R3 [108-1]; (8) nine or more caudal-fin
rays in contact with the hypural plate [132-1]; (9) anterior and
posterior orbital bones (143-1); and (10) viviparity [141-1*].
Composition. Tribes Tomeurini, Alfarini, Brachyrhaphini,
Priapichthyini, Priapellini, Heterandriini, Gambusiini, Poeciliini,
and Cnesterodontini.
Distribution. North America through Central America, the
Caribbean, through South America to Argentina.
Remarks. The family name Poecilini has already been used
by Bonaparte (1831), however it appeared to be preoccupied
in Coleoptera. The family-group name based on Poecilus
Bonelli (Carabidae) was created by Bonelli in 1810. He called
the group “Poecilii”, which is typically taken to be a family-
group name. When it is used these days, it is either as a tribe
(Poecilini) or a subtribe (Poecilina) (David Maddison, in litt.).
Later, Bonaparte (1846) added one “i” to the name differing it
from the Coleoptera family-group name (even if the differ-
ences between two family-group names is only one letter
they are not homonyms - article 55.4 of the ICZN, 1999).
However, Poeciliini Bonaparte, 1846 is an unjustified
emendation for Poecilini Bonaparte, 1831 (article 32.5.3 of
the ICZN, 1999) but it is in prevailing usage. So, it is attrib-
uted to the original author and date and is deemed to be a
justified emendation following the article 33.2.3.1 of the
ICZN (1999).
Tribe Tomeurini Eigenmann
[Clade 63]
Tomeurini Eigenmann, 1912: 460. Type-genus: Tomeurus
Eigenmann, 1909.
Diagnosis. Tomeurins can be diagnosed by the following
uniquely derived features: (1) preopercular canal partially
closed, only canals between pores 11-12, and 12-U closed
(sometimes canal U-V also closed) [8-4*]; (2) three pelvic-
fin rays in females [44-2*]; (3) haemal arch and spine of
vertebrae 13-17 in adult males absent [47-1*]; (4) first proxi-
mal radial of dorsal fin in adult males located between neu-
ral spines of 23
rd
and 24
th
or 24
th
and 25
th
vertebrae [62-3*];
(5) adult females with first proximal radial of dorsal fin lo-
cated between neural arches of vertebrae 6; 23 and 24 or 24
and 25 [63-6*]; and (6) six dorsal-fin rays (males and fe-
males) [64-4*].
Additionally, tomeurins can be diagnosed by the follow-
ing not uniquely derived and/or reversed features: (1) ante-
rior margin of frontals extend anterior by between nasals [1-
0]; (2) parietals short restricted to the epiotic region, not reach-
ing sphenotic anteriorly [2-1]; (3) absence of an epiotic pro-
P. H. F. Lucinda & R. E. Reis
37
PROOFS
cess [3-3]; (4) posterior section of posterior remnant of infraor-
bital system opened into a groove [7-1]; (5) preorbital canal ab-
sent or opened, forming a very shallow groove [9-2]; (6) man-
dibular canal absent or opened, forming a very shallow groove
[10-1]; (7) anterior process of anterior ceratohyal extending ven-
tral to ventral hypohyal present [24-0]; (8) interarcual cartilage
absent [25-1]; (9) three pelvic-fin rays in males [33-3]; (10) pelvic
girdle of males located below pectoral girdle, posterior border of
basipterygium anterior to posterior border of cleithrum [35-3];
(11) dorsolateral process of pelvic fin in adult males large [36-1];
(12) anterior tip of basipterygium in adult males clearly pointed
[37-1]; (13) width of first pelvic-fin ray in adult males decreasing
abruptly at distal portion, distal slender portion short [41-2]; (14)
second pelvic-fin ray in adult males unbranched [42-1]; (15) dis-
tal portion of third and fourth gonactinosts separate, except by
tip of gonactinost [74-2]; (16) eight anal-fin rays in males [85-3];
(17) dorsal expansion of ray 5p of anal fin in adult males present
[124-1]; (18) R6a and R6p partially fused [126-1]; (19) more distal
elements of R6 branches partially fused [127-1]; and (20) orbital
bones absent [143-0].
Composition.Genus Tomeurus.
Distribution. As for Tomeurus.
GenusTomeurus Eigenmann
Tomeurus Eigenmann, 1909: 53. Genus masculine. Type-spe-
cies: Tomeurus gracilis Eigenmann, 1909. Type by
monotypy and original designation.
Composition.Tomeurus gracilis Eigenmann.
Distribution.Tomeurus gracilis occurs in small coastal drain-
ages of the Venezuelan departments Delta Amacuro,
Monagas, Territorio Federal, and in Brazilian states of Amapá,
and Pará. The species also inhabits the drainages of rio Guamá
and rio Tocantins in Brazil, the drainages of the Cuyuni,
Mazaruni, and Essequibo Rivers in the Guyana and
Courantyne rivier drainage in Suriname.
Tribe Alfarini + Tribe Brachyrhaphini +
Tribe Priapichthyini + Tribe Priapellini +
Tribe Heterandriini + Tribe Gambusiini +
Supertribe Poeciliini
[Clade 126]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features: (1)
posterior supraorbital canal (2b, 3, 4a) opened, forming a sinu-
ous depression over the frontal (supraorbital bone) [5-1*];
(2) anterior section of posterior remnant of infraorbital sys-
tem (canal 4b, 5, 6a) opened, pores confluent forming a major
sinuous depression above and slightly behind the orbit [6-
1]; (3) medial surface of ascending process of premaxilla
slightly angled laterally [11-1*]; (4) anterior border of ventral
maxilla concave [14-1]; (5) pelvic girdle of males posteriorly
located; posterior border of cleithrum approximately aligned
with center of basipterygium (or more posterior) [35-1]; (6)
ligastyle with one axis [46-1]; and (7) adult males with anterior
process on base of fifth middle anal-fin radial pointed and
upward directed [78-1].
Tribe Alfarini Hubbs
[Clade 64]
Alfarini Hubbs, 1924: 11.Type-genus: Alfaro Meek, 1912.
Diagnosis. Alfarins can be diagnosed by the following not
uniquely derived and/or reversed features: (1) pleural ribs
associated with haemal arches in males [60-1]; (2) pleural ribs
associated with haemal arches in females [61-1]; (3) adult fe-
males with first proximal radial of dorsal fin located between
neural arches of 14
th
and 15
th
vertebrae; (4) 12 or more anal-fin
rays in females [65-0]; (5) gonactinost complex approximately
perpendicular to body longitudinal axis [68-1]; (6) distal por-
tion of second and third gonactinosts fused [71-1]; (7) distal
portion of third and fourth gonactinosts completely fused
[74-1]; (8) lateral process on base of fifth middle anal-fin ra-
dial in adult males minute [79-4]; and (9) presence of a palp in
subdistal segments of R3 [88-1].
Composition.Genus Alfaro.
Distribution. As for Alfaro.
Alfaro Meek
Petalosoma Regan, 1908: 458. Type species: Petalosoma
cultratum Regan, 1908. Type by monotypy. Gender: neu-
ter. Preoccupied by Petalosoma Lewis, 1903 in Coleoptera.
Alfaro Meek, 1912: 72. Type species: Alfaro acutiventralis
Meek, 1912. Type by monotypy. Gender: masculine.
Petalurichthys Regan, 1912: 494 [footnote]. Type species:
Petalosoma cultratum Regan, 1908. Type by being a re-
placement name. Gender: masculine.
Furcipenis Hubbs, 1931: 1. Type species: Priapichthys huberi
Fowler, 1923. Type by original designation. Gender: mas-
culine.
Composition.Alfaro cultratus (Regan) and A. huberi (Fowler)
Distribution. Southern Guatemala, Honduras, Costa Rica,
Nicaragua, and Western Panama.
Tribe Brachyrhaphini + Tribe Priapichthyini +
Tribe Priapellini + Tribe Heterandriini +
Tribe Gambusiini + Supertribe Poeciliini
[Clade 125]
Diagnosis.Members of this clade share the following uniquely
derived and unreversed features: (1) haemal arch and spine
Systematics of the subfamily Poeciliinae Bonaparte
38
PROOFS
of vertebrae 13-17 in adult males modified into gonapophyses
[47-2**]; and (2) lateral process on base of fifth middle anal-
fin radial in adult males large [79-1**].
Additionally, they can be diagnosed by the following not
uniquely derived and/or reversed features: (1) epiotic process
longer than exoccipital process but not reaching first pleural rib [3-
1*]; (2) preorbital canal partially closed bearing two upper pores
and a lower deep groove [9-1]; (3) three well-developed
gonapophyses [48-1*]; (4) functional gonapophyses located on
vertebrae 14, 15, and 16 [49-1*]; (5) gonactinost complex inclined
forward, forming an angle wider than 90
º
relative to the body lon-
gitudinal axis [68-2*]; (6) basal process on first gonactinost small
[69-1*]; (7) second and third gonactinosts partially fused [72-2*];
(8) anal-fin posterior median radials (5
th
to last one) in adult males
not fused to respective proximal radials [77-0]; (9) one series of
subdistal retrorse spines on R4p [114-1*]; (10) eight or more
subdistal retrorse spines on R4p [116-1]; and (11) dark stripe present
on median portion of dorsal fin [137-1].
Tribe Brachyrhaphini, new
[Clade 61]
Type-genus: Brachyrhaphis Regan, 1913.
Diagnosis. Brachyrhaphins can be diagnosed by the following
not uniquely derived and/or reversed features: (1) anterior mar-
gin of frontals extend anterior by between nasals [1-0]; (2) ante-
rior process of anterior ceratohyal extending ventral to ventral
hypohyal present [24-0]; (3) pelvic girdle of males very posterior,
anterior border of basipterygium posterior to the posterior bor-
der of cleithrum [35-0]; (4) second gonapophysis approximately
perpendicular to vertebral column [54-4]; (5) first proximal radial
of dorsal fin in adult males located between neural spines of 11
th
and 12
th
vertebrae [62-6]; (6) adult females with first proximal
radial of dorsal fin located between neural arches of 11
th
and 12
th
vertebrae [63-3]; (7) nine dorsal-fin rays (males and females) [64-
1]; (8) anterior process on base of fifth middle anal-fin radial in
adult males hardly developed and round [78-2]; (9) ninth
gonactinost bearing wing-like lateral projections [84-1]; (10) males
with 10 anal-fin rays [85-0]; (11) R5a, R5p, R4a, and R4p directed
upwards [112-1]; (12) less than nine caudal-fin rays in contact
with the hypural plate [132-0]; (13) dorsal fin moderately pig-
mented with black [136-1]; (14) ground pigmentation of anal fin
of females moderately speckled with black, with chromatophores
more concentrated anteriorly and forming a dark stripe on first
rays [140-1]; and (14) orbital bones absent [143-0].
Composition.Genus Brachyrhaphis.
Distribution. As for Brachyrhaphis.
Brachyrhaphis Regan
Brachyrhaphis Regan, 1913: 997. Type species: Gambusia
rhabdophora Regan, 1908. Type by monotypy. Gender:
feminine.
Trigonophallus Hubbs, 1926: 48. Type species:
Trigonophallus punctifer Hubbs, 1926. Type by original
designation. Gender: masculine.
Plectrophallus Fowler, 1932: 384. Proposed as new subge-
nus of Panamichthys Hubbs. Type species: Panamichthys
tristani Fowler, 1932. Type by original designation. Gen-
der: masculine.
Composition. Brachyrhaphis cascajalensis (Meek &
Hildebrand), B. episcopi (Steindachner), B. hartwegi Rosen
& Bailey, B. hessfeldi Meyer & Etzel, B. holdridgei Bussing,
B. parismina (Meek), B. punctifer (Hubbs), B. rhabdophora
(Regan), B. roseni Bussing, B. roswithae Meyer & Etzel, B.
terrabensis (Regan).
Distribution.Mexico (southern Pacific drainages), Costa Rica
(Pacific and Atlantic drainages), Guatemala, Panama (central
and western).
Tribe Priapichthyini + Tribe Priapellini +
Tribe Heterandriini + Tribe Gambusiini +
Supertribe Poeciliini
[Clade 124]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features: (1)
anterior tip of basipterygium in adult males clearly pointed
[37-1]; (2) third gonapophysis forming an angle of 35-70 de-
grees relative to vertebral column [55-1]; (3) absence of an
anterior convex expansion of second gonactinost [70-1]; (4)
males with eleven anal-fin rays [85-2]; (5) R6a and R6p par-
tially fused [126-1]; and (6) more distal elements of R6 branches
partially fused [127-1].
Tribe Priapichthyini, new
[Clade 55]
Type-genus: Priapichthys
Regan, 1913
Diagnosis. Priapichthyins share the following uniquely de-
rived and unreversed features: (1) mandibular canal present
and partially closed bearing six pores [10-3*]. Addition-
ally, this tribe can be diagnosed by the following not
uniquely derived and/or reversed features: (1) ascending
process of premaxilla elongate, distal tip pointed [12-1];
(2) second gonapophysis forming an angle of 45-70 de-
grees relative to vertebral column [54-1]; (3) absence of a
basal process on first gonactinost [69-0]; (4) distal portion
of second and third gonactinosts fused [71-1]; (5) com-
plete fusion of second and third gonactinosts [72-1]; (6)
distal portion of third and fourth gonactinosts completely
fused [74-1]; (7) eighth gonactinost bearing wing-like lat-
eral projections [83-1]; and (8) ninth gonactinost bearing
wing-like lateral projections [84-1].
Composition.Genus Priapichthys.
P. H. F. Lucinda & R. E. Reis
39
PROOFS
Distribution. As for Priapichthys.
Priapichthys Regan
Priapichthys Regan, 1913: 991. Type species: Gambusia
annectens Regan, 1907. Gender: masculine.
Diphyacantha Henn, 1916: 113. Type species: Diphyacantha
chocoensis Henn, 1916. Type by monotypy. Gender: femi-
nine.
Darienichthys Hubbs, 1924: 8 [footnote]. Type species: Gam-
busia darienensis Meek & Hildebrand, 1913. Type by origi-
nal designation. Gender: masculine.
Panamichthys Hubbs, 1924: 8 [footnote]. Type species:
Priapichthys panamensis Meek & Hildebrand, 1916. Type
by original designation. Gender: masculine.
Alloheterandria Hubbs, 1924: 9 [footnote]. Type species:
Gambusia nigroventralis Eigenmann & Henn, 1912. Type
by original designation. Gender: feminine.
Composition. Priapichthys annectens (Regan), P. caliensis
(Eigenmann & Henn), P. chocoensis Henn, P. darienensis
(Meek & Hildebrand), P. nigroventralis (Eigenmann & Henn),
P. panamensis Meek & Hildebrand, P. puetzi Meyer & Etzel.
Distribution. Costa Rica (Pacific and Atlantic drainages),
Panama (Pacific drainages), Ecuador and Colombia.
Tribe Priapellini + Tribe Heterandriini +
Tribe Gambusiini + Supertribe Poeciliini
[Clade 123]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features:
(1) ligastyle triangular [46-2]; (2) curvature of
gonapophysis forming an angle of 16-45 degrees relative
to vertebral column [53-1*]; (3) pleural ribs 7, 8, and 9 in
adult males curved forward converging to the same point
towards pelvic girdle [59-1*]; (4) females with eleven anal-
fin rays [65-2]; and (5) elongate and dorsal protuberance
present along R4p (just behind retrorse spines series) of
anal fin in adult males [118-1].
Tribe Priapellini Ghedotti
[Clade 62]
Priapellini Ghedotti, 2000: 39. Type genus: Priapella Regan,
1913
Diagnosis. Priapellins share the following uniquely derived
and unreversed features: (1) mesethmoid: cartilaginous [0-
1*]; and (2) mandibular canal present and entirely closed,
bearing five pores [10-2*].
Additionally, priapellins can be diagnosed by the follow-
ing not uniquely derived and/or reversed features: (1) ante-
rior margin of frontals straight or slightly cleft medially [1-1];
(2) absence of an epiotic process [3-3]; (3) preorbital canal
present and entirely closed, bearing four pores [9-0]; (4) ante-
rior border of ventral maxilla straight [14-0]; (5) interarcual
cartilage absent [25-1]; (6) pleural ribs associated with haemal
arches in males [60-1]; (7) pleural ribs associated with haemal
arches in females [61-1]; (8) absence of an anterior convex
expansion of second gonactinost [70-0]; and (9) absence of a
dark stripe on median portion of dorsal fin [137-0].
Composition. Genus Priapella.
Distribution. As for Priapella.
Priapella
Regan
Priapella Regan, 1913: 992. Type species: Gambusia bonita
Meek, 1904. Type by monotypy. Gender: feminine.
Composition.Priapella bonita (Meek), P.compressa Alvarez,
P.intermedia Alvarez & Carranza, P. olmecae Meyer & Perez.
Distribution. Southern Mexico.
Tribe Heterandriini + Tribe Gambusiini +
Supertribe Poeciliini
[Clade 122]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features: (1)
anterior margin of frontals extend anterior by between nasals
[1-0]; (2) posterior supraorbital canal (2b, 3, 4a) absent or
opened, forming a shallow groove [5-0]; (3) third
gonapophysis forming an angle of 10-32 degrees relative to
vertebral column [55-2*]; (4) first proximal radial of dorsal fin
in adult males located between neural spines of 13
th
and 14
th
vertebrae [62-1]; and (5) hypural plate partially fused with an
elongate aperture [131-1].
Tribe Heterandriini Hubbs, new usage
[Clade 54]
Heterandriini Hubbs, 1924: 7. Type-genus: Heterandria
Agassiz, 1853.
Diagnosis. Heterandriins share the following uniquely de-
rived and unreversed feature: (1) distal portion of third and
fourth gonactinosts completely fused, except by a small
notch [74-3*]. Heterandriins can also be diagnosed by the
following not uniquely derived and/or reversed features:
(1) epiotic process shorter than exoccipital process [3-2];
(2) anterior process of anterior ceratohyal extending ventral
to ventral hypohyal present [24-0]; (3) dorsolateral process
of pelvic fin in adult males large [36-1]; (4) width of first
pelvic-fin ray in adult males decreasing abruptly at distal
portion, distal slender portion long [41-1]; (5) first proximal
radial of dorsal fin in adult males located between neural
spines of 11
th
and 12
th
vertebrae [62-6]; (6) absence of a basal
Systematics of the subfamily Poeciliinae Bonaparte
40
PROOFS
process on first gonactinost [69-0]; (7) distal portion of sec-
ond and third anal-fin gonactinosts fused [71-1]; (8) mem-
branous tip anterior to R4 and R5 curved downwards [110-
1]; (9) membranous tip anterior to R4 and R5 small [111-1];
and (10) dorsal expansion of ray 5p of anal fin in adult males
present [124-1].
Composition.Genus Heterandria.
Distribution. As for Heterandria.
Heterandria Agassiz
Heterandria Agassiz, 1853: 135. Type species: Heterandria
formosa Girard, 1859. Gender: feminine. Type by subse-
quent designation by Bailey (1952).
Pseudoxiphophorus Bleeker, 1860: 440. Type species:
Xiphophorus bimaculatus Heckel, 1848. Gender: mascu-
line.
Poeciliodes Steindachner, 1863: 176 [15]. Type species:
Poeciliodes bimaculatus Steindachner, 1863. Type by
monotypy. Gender: masculine.
Composition. Heterandria anzuetoi Rosen & Bailey, H.
attenuata Rosen & Bailey, H. bimaculata (Heckel), H. cataractae
Rosen, H. dirempta Rosen, H. formosa Girard, H. jonesii
(Günther), H. litoperas Rosen & Bailey, and H. obliqua Rosen.
Distribution.Southern USA to Nicaragua (Atlantic and Gulf
drainages) and Guatemala (Pacific drainage).
Tribe Gambusiini + Supertribe Poeciliini
[Clade 121]
Diagnosis. Members of this clade can be diagnosed by the fol-
lowing not uniquely derived and/or reversed features: (1) ante-
rior section of posterior remnant of infraorbital system absent or
opened, forming a shallow groove [6-0]; (2) mandibular canal
absent or opened, forming a very shallow groove [10-1]; (3)
medial surface of ascending process of premaxilla angled later-
ally at proximal end, forming a triangle space between proximal
ends of ascending processes [11-2]; (4) anterior border of ven-
tral maxilla straight [14-0]; (5) ascending process of
parasphenoids in adults short, not reaching pterosphenoids [20-
1]; (6) adult females with first proximal radial of dorsal fin located
between neural arches of 13
th
and 14
th
vertebrae [63-1]; and (7)
nine dorsal-fin rays (males and females) [64-1].
Tribe Gambusiini Gill, new usage
[Clade 120]
Gambusiini Gill, 1893: 133. Type-genus: Gambusia Poey,
1854.
Diagnosis. Gambusiins can be diagnosed by the following
not uniquely derived and/or reversed features: (1) preorbital
canal absent or opened, forming a very shallow groove [9-2];
(2) ascending process of premaxilla elongate, distal tip pointed
[12-1]; (3) absence of an elongate and dorsal protuberance
along R4p (just behind retrorse spines series) of anal fin in
adult males [118-0]; and (4) more distal elements of R6
branches not fused [127-0].
Composition. Genera Gambusia, Belonesox, Pseudopoecilia,
Neoheterandria, Scolichthys.
Distribution. Northern USA to Peru (Inland, Gulf, Atlantic
and Pacific drainages) including the Caribbean Islands.
Gambusia + Belonesox
[Clade 118]
Diagnosis. Gambusia and Belonesox share the following
uniquely derived and unreversed features: (1) gonactinost 2,
3, and 4 fused into a column [73-1**]; and (2) lateral flanges
on ventral portion of anal-fin radial 4 in adult males present
and continuous, without dorsal cleft [75-1**].
Additionally, this clade can be diagnosed by the following
not uniquely derived and/or reversed features: (1) posterior
section of posterior remnant of infraorbital system opened into
a groove [7-1]; (2) pelvic girdle of males very posterior, anterior
border of basipterygium posterior to the posterior border of
cleithrum [35-0]; (3) anterior tip of basipterygium in adult males
approximately triangular and round [37-0]; (4) pleural ribs in
adult males almost straight, slightly curving forward and not
converging to the same point towards pelvic girdle [59-0]; (5)
pleural ribs associated with haemal arches in males [60-1]; (6)
absence of an anterior convex expansion of second gonactinost
[70-0]; (7) complete fusion of second and third gonactinosts
[72-1]; (8) distal portion of third and fourth gonactinosts com-
pletely fused [74-1]; (9) lateral process on base of fifth middle
anal-fin radial in adult males minute [79-4]; (10) R5a, R5p, R4a,
and R4p directed upwards [112-1]; (11) four to seven subdistal
retrorse spines on R4p [116-2]; (12) hook on 5a contacting the
segments of 4p [123-1]; and (13) less than nine caudal-fin rays
in contact with the hypural plate [132-0].
GambusiaPoey
Gambusia Poey, 1854: 382. Type species: Gambusia punctata
Poey, 1854. Type by subsequent designation. Gender: femi-
nine.
Paragambusia Meek, 1904: 133. Type species: Gambusia
nicaraguensis Günther, 1866. Type by original designa-
tion. Gender: feminine.
Heterophallus Regan, 1914: 65. Type species: Heterophallus
rachovii Regan, 1914. Type by monotypy. Gender: mas-
culine.
Arthrophallus Hubbs, 1926: 38. Proposed as subgenus of
Gambusia. Type species: Heterandria patruelis Baird &
Girard, 1853. Type by original designation. Gender: mas-
culine.
P. H. F. Lucinda & R. E. Reis
41
PROOFS
Heterophallina Hubbs, 1926: 26. Proposed as subgenus of
Gambusia. Type species: Gambusia regani Hubbs, 1926.
Type by original designation. Gender: feminine.
Schizophallus Hubbs, 1926: 40. Proposed as subgenus of
Gambusia. Type species: Gambusia holbrookii Girard,
1859. Type by original designation. Gender: masculine.
Dicerophallus Alvarez, 1952: 95. Type species: Dicerophallus
echeagarayi Alvarez, 1952. Type by original designation.
Gender: masculine.
Flexipenis Hubbs in Rivas, 1963: 334. Type species: Gambu-
sia vittata Hubbs, 1926. Type by original designation.
Gender: masculine.
Orthophallus Rivas, 1963: 339. Type species: Gambusia
lemaitrei Fowler, 1950. Type by original designation. Gen-
der: masculine.
Composition.Gambusia affinis (Baird & Girard); G. alvarezi
Hubbs & Springer, G. amistadensis Peden, G. atrora Rosen
& Bailey, G. aurata Miller & Minckley, G. beebei Myers, G.
bucheri Rivas, G. clarkhubbsi Garrett & Edwards, G.
dominicensis Regan, G. echeagarayi (Alvarez), G. eurystoma
Miller, G. gaigei Hubbs, G. geiseri Hubbs & Hubbs, G.
georgei Hubbs & Peden, G. heterochir Hubbs,G. hispaniolae
Fink, G. hurtadoi Hubbs & Springer, G. krumholzi Minckley,
G. lemaitrei Fowler, G. longispinis Minckley, G. luma Rosen
& Bailey, G. marshi Minckley & Craddock, G. melapleura
(Gosse); G. myersi Ahl, G. nicaraguensis Günther, G. nobilis
(Baird & Girard), G. panuco Hubbs,G. pseudopunctata Rivas,
G. punctata Poey, G. manni Hubbs, G. monticola Rivas, G.
puncticulata Poey, G. rachowi (Regan), G. regani Hubbs, G.
rhizophorae Rivas, G. senilis Girard, G. sexradiata Hubbs,
G. speciosa Girard, G. vittata Hubbs, G. wrayi Regan, G.
xanthosoma Greenfield, and G. yucatana Regan.
Distribution. Northern USA to Colombia (Inland, Gulf and
Pacific drainages) including the Caribbean Islands.
Belonesox Kner
Belonesox Kner, 1860: 419, 422. Type species: Belonesox
belizanus Kner, 1860. Type by monotypy. Gender: mascu-
line.
Composition. Belonesox belizanus Kner.
Distribution. Central America: from laguna San Julian, north-
east of Ciudad Veracruz in Mexico to Costa Rica. Southern Gulf
of Mexico, southern Yucatán and along Central American coast
south to Nicaragua. Introduced in freshwater in Florida.
Pseudopoecilia + Neoheterandria + Scolichthys
[Clade 117]
Diagnosis. Pseudopoecilia + Neoheterandria + Scolichthys
share the following not uniquely derived and/or reversed fea-
tures: (1) medial surface of ascending process of premaxilla
approximately straight [11-0]; (2) ligastyle absent [46-0]; (3)
10 or more dorsal-fin rays (males and females) [64-0]; (4) males
with nine anal-fin rays [85-4]; (5) spines on subdistal seg-
ments of R3 retrorse [108-2]; (6) dorsal expansion of ray 5p of
anal fin in adult males present [124-1]; and (7) hypural plate
completely fused [131-0].
Pseudopoecilia Regan
Pseudopoecilia Regan, 1913: 995. Type species: Poecilia festae
Boulenger, 1898. Type by monotypy. Gender: feminine.
Composition. Pseudopoecilia austrocolumbiana Radda, P.
festae (Boulenger), P. fria (Eigenmann & Henn)
Distribution.Pacific drainages of Ecuador, Peru, and Colombia.
Neoheterandria + Scolichthys
[Clade 114]
Diagnosis. Neoheterandria and Scolichthys share the fol-
lowing uniquely derived and unreversed feature: (1) first and
second branchiostegal rays united at the base [23-1**]. Ad-
ditionally, Neoheterandria plus Scolichthys can be diagnosed
by the following not uniquely derived and/or reversed fea-
tures: (1) females with 10 anal-fin rays [65-1]; (2) ventral pro-
jection of R4a towards R3 [113-1]; and (3) more distal ele-
ments of R6 branches totally fused [127-2].
Neoheterandria Henn
Neoheterandria Henn, 1916: 117. Type species:
Neoheterandria elegans Henn, 1916. Type by monotypy.
Gender: feminine.
Allogambusia Hubbs, 1924: 8. Type species: Gambusia
tridentiger Garman, 1895. Type by original designation.
Gender: feminine.
Composition. Neoheterandria cana (Meek & Hildebrand),
N. elegans Henn, N. tridentiger (Garman).
Distribution.Nicaragua (Atlantic drainages), Costa Rica (At-
lantic drainages), Panama (central, Atlantic and Pacific drain-
ages), and Colombia (Atlantic drainages).
Scolichthys Rosen
Scolichthys Rosen, 1967: 2. Type species: Scolichthys
greenwayi Rosen, 1967. Type by original designation.
Gender: masculine.
Composition. Scolichthys greenwayi
Rosen and S. iota
Rosen.
Distribution. Río Chixoy, río Chajmaic, and río Salinas sys-
tem in Alta Verapaz, Guatemala.
Systematics of the subfamily Poeciliinae Bonaparte
42
PROOFS
Supertribe: Poeciliini Bonaparte, new
[Clade 119]
Poecilini Bonaparte, 1831: 94, unavailable name; preoccupied
in Coleoptera.
Poeciliini Bonaparte, 1831. Type-genus: Poecilia Bloch &
Schneider, 1801.
Diagnosis. Members of the supertribe Poeciliini share the
following uniquely derived and unreversed features: (1) as-
cending process of premaxilla short and pointed [12-2**]; (2)
presence of a curved and forward directed process on ventral
surface of dentary [15-1**]; (3) tooth plates of third and fourth
pharingobranchials fused, forming a elongate structure with
teeth regularly distributed [26-1**]; and (4) fifth
ceratobranchial wide and bearing teeth regularly distributed
[28-1**].
Additionally, they can be diagnosed by the following not
uniquely derived and/or reversed features: (1) premaxillary
symphysis elevated [13-1*]; (2) teeth compressed [21-1*];
(3) five branchiostegal rays [22-0]; (4) toothless fourth
ceratobranchial [27-1]; (5) absence of spines on subdistal
segments of R3 [108-0]; and (6) absence of a dark stripe on
median portion of dorsal fin [137-0].
Composition. Tribes: Poeciliini, Girardinini and
Cnesterodontini
Distribution. North, Central and South America.
Tribe Girardinini Hubbs, new usage
[Clade 116]
Girardinini Hubbs, 1924: 9. Type-genus: Girardinus Poey, 1854.
Diagnosis. Girardinins can be diagnosed by the following
not uniquely derived and/or reversed features: (1) width of
first pelvic-fin ray in adult males decreasing abruptly at distal
portion, distal slender portion long [41-1]; (2) distal portion
of second and third gonactinosts fused [71-1]; (3) more distal
elements of R6 branches totally fused [127-2]; and (4) dark
patch of pigmentation along R3 [138-1].
Composition.Genera Girardinus, Phallichthys, Xenophallus,
Poeciliopsis, Phalloptychus.
Distribution.USA, Mexico, Cuba, Belize, Guatemala, Hondu-
ras, Costa Rica, Panama, Colombia, and Brazil.
Girardinus Poey
Girardinus Poey, 1854: 383. Type species: Girardinus
metallicus Poey, 1854. Type by monotypy. Gender: mas-
culine.
Glaridodon Garman, 1895: 40. Type species: Girardinus
uninotatus Poey, 1861. Type by original designation. Not
available; name preoccupied in fossil Synapsida. Gender:
masculine.
Glaridichthys Garman, 1896: 232. Type species: Girardinus
uninotatus Poey, 1861. Type by being a replacement name
for Glaridodon Garman, 1895 preoccupied in fossil Rep-
tilia. Gender: masculine.
Toxus Eigenmann, 1903: 226. Type species: Toxus riddlei
Eigenmann, 1903. Type by original designation. Gender:
masculine.
Allodontium Howell Rivero & Rivas, 1944: 17. Type species:
Heterandria cubensis Eigenmann, 1904. Type by original
designation. Gender: neuter.
Dactylophallus Howell Rivero & Rivas, 1944: 15. Type spe-
cies: Girardinus denticulatus Garman, 1895. Type by origi-
nal designation. Gender: masculine.
Composition. Girardinus creolus Garman, G. cubensis
(Eigenmann), G
. denticulatus Garman, G. falcatus
(Eigenmann), G. metallicus Poey, G. microdactylus Rivas, G.
uninotatus Poey.
Distribution.Cuba.
Phallichthys + Xenophallus +
Poeciliopsis + Phalloptychus
[Clade 113]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features: (1)
parietals short restricted to the epiotic region, not reaching
sphenotic anteriorly [2-1]; (2) epiotic process long extending
beyond first pleural rib [3-0]; (3) ascending process of
parasphenoids in adults long, contacting pterosphenoids [20-
0]; (4) first proximal radial of dorsal fin in adult males located
between neural spines of 10
th
and 11
th
vertebrae [62-4]; (5)
adult females with first proximal radial of dorsal fin located
between neural arches of 10
th
and 11
th
vertebrae [63-2]; (6)
absence of a basal process on first gonactinost [69-0]; (7)
complete fusion of second and third gonactinosts [72-1]; (8)
lateral process on base of fifth middle anal-fin radial in adult
males asymmetrical [79-3]; (9) middle anal-fin radials 5, 6, and
7 in adult males asymmetrical (right lateral projection more
compressed and much larger than left one) [80-1]; (10) anal
fin asymmetrical in adult males [87-1]; (11) more distal ele-
ments of R6 branches partially fused [127-1]; and (12) hypural
plate bipartite [131-2].
Phallichthys Hubbs
Phallichthys Hubbs 1924: 10. Type species: Poeciliopsis
isthmensis, Regan 1913. Type by original designation. Gen-
der: masculine.
Composition. Phallichthys amates (Miller); P. fairweatheri
Rosen & Bailey, P. quadripunctatus Bussing, P. tico Bus-
sing.
P. H. F. Lucinda & R. E. Reis
43
PROOFS
Distribution. Belize and northern Guatemala to Costa Rica
and western Panama (Atlantic drainages). Introduced on Pa-
cific Slope of Costa Rica.
Xenophallus + Poeciliopsis + Phalloptychus
[Clade 110]
Diagnosis. Xenophallus, Poeciliopsis, and Phalloptychus
share the following uniquely derived and unreversed fea-
ture: (1) convergent anal-fin proximal radials of females
[66-2**].
Additionally, this clade can be diagnosed by the following not
uniquely derived and/or reversed features: (1) width of first pel-
vic-fin ray in adult males decreasing abruptly at distal portion,
distal slender portion long [41-1]; (2) ligastyle with one axis [46-1];
(3) females with 10 anal-fin rays [65-1]; (4) distal portion of third
and fourth gonactinosts completely fused [74-1]; (5) absence of
an anterior process on base of fifth middle anal-fin radial in adult
males [78-0]; (6) males with 10 anal-fin rays [85-0]; (7) absence of
an elongate and dorsal protuberance along R4p (just behind ret-
rorse spines series) of anal fin in adult males [118-0]; and (8) more
distal elements of R6 branches totally fused [127-2].
XenophallusHubbs
Xenophallus Hubbs, 1924: 10 [footnote]. Type species: Gam-
busia umbratilis Meek, 1912. Type by original designa-
tion. Gender: masculine.
Composition. Xenophallus umbratilis (Meek)
Distribution. Costa Rica (Atlantic Drainages).
Poeciliopsis + Phalloptychus
[Clade 105]
Diagnosis. Poeciliopsis and Phalloptychus share the follow-
ing uniquely derived and unreversed features: (1) ventral pro-
cess of anguloarticular short, not extending anterior to where
anguloarticular overlaps dentary; [18-1**]; (2) gonapophysis
of vertebra 14 very curved in adult males [52-1**]; and (3)
branches of R6a and R6p completely fused [126-2**].
Additionally, this clade can be diagnosed by the following
not uniquely derived and/or reversed features: (1) anterior tip
of basipterygium in adult males clearly round and keeled [37-
3]; (2) presence of a lateral keel in basipterygium in adult males
[38-1]; (3) first proximal radial of dorsal fin in adult males lo-
cated between neural spines of 12
th
and 13
th
vertebrae [62-0];
and (4) adult females with first proximal radial of dorsal fin
located between neural arches of 12
th
and 13
th
vertebrae [63-0].
Poeciliopsis Regan
Poeciliopsis Regan, 1913: 996. Type species: Poecilia
presidionis Jordan & Culver, 1895. Type by subsequent
designation. Gender: feminine.
Leptorhaphis Regan, 1913: 998. Type species: Gambusia
infans Woolman, 1894. Type by monotypy. Gender:
feminine.
Aulophallus Hubbs, 1926: 64, 69. Type species: Poecilia
elongata Günther, 1866. Type by original designation.
Gender: masculine.
Poecilistes Hubbs, 1926: 63, 68. Type species: Heterandria
lutzi Meek, 1904. Type by original designation. Gender:
masculine.
Composition. Poeciliopsis baenschi Meyer, Radda, Riehl
& Feichtinger, P. balsas Hubbs, P. catemaco Miller, P.
elongata (Günther); P. fasciata (Meek); P. gracilis
(Heckel), P. hnilickai Meyer & Vogel, P. infans (Woolman),
P. latidens (Garman), P. lucida Miller, P. lutzi (Meek); P.
monacha Miller, P. occidentalis (Baird & Girard), P.
paucimaculata Bussing, P. presidionis (Jordan & Culver),
P. prolifica Miller, P. retropinna (Regan), P. scarlli Meyer,
Riehl, Dawes & Dibble, P
. turneri Miller, P. turrubarensis
(Meek); P. viriosa Miller.
Distribution. Southern USA to Colombia (Pacific drainages),
and southeastern Mexico, Guatemala, and Honduras (Atlan-
tic drainages).
Phalloptychus Eigenmann
Phalloptychus Eigenmann, 1907: 426, 430. Type species:
Girardinus januarius Hensel, 1868. Type by original des-
ignation. Gender: masculine.
Composition. Phalloptychus eigenmanni Henn, P. januarius
(Hensel), P. iheringii (Boulenger).
Distribution. Brazil: rio Catu in Alagoinhas (Bahia) and
coastal drainages from Rio de Janeiro to Rio Grande do
Sul.
Tribe Poeciliini + Tribe Cnesterodontini
[Clade 115]
Diagnosis. Poeciliins and Cnesterodontins can be diag-
nosed by the following not uniquely derived and/or re-
versed features: (1) parietals absent [2-2]; (2) anterior tip
of basipterygium in adult males approximately triangular
and round [37-0]; (3) ligastyle with one axis [46-1]; (4)
pleural ribs associated with haemal arches in males [60-
1]; (5) pleural ribs associated with haemal arches in fe-
males [61-1]; (6) absence of an anterior convex expansion
of second gonactinosts [70-0]; (7) absence of an anterior
process on base of fifth middle anal-fin radial in adult
males [78-0]; (8) males with 10 anal-fin rays [85-0]; and (9)
absence of an elongate and dorsal protuberance along
R4p (just behind retrorse spines series) of anal fin in adult
males [118-0].
Systematics of the subfamily Poeciliinae Bonaparte
44
PROOFS
Tribe Poeciliini Bonaparte, new usage
[Clade 112]
Poecilini Bonaparte, 1831: 94, unavailable name; preoccupied
in Coleoptera.
Poeciliini Bonaparte, 1831. Type-genus Poecilia Bloch &
Schneider, 1801.
Diagnosis. Poeciliins can be diagnosed by the following not
uniquely derived and/or reversed features: (1) notch on dentary
[16-1]; (2) ascending process of parasphenoids in adults long,
contacting pterosphenoids [20-0]; (3) second gonapophysis form-
ing an angle of 45-70 degrees relative to vertebral column [54-1];
(4) third gonapophysis forming an angle of 35-70 degrees relative
to vertebral column [55-1]; (5) first proximal radial of dorsal fin in
adult males located between neural spines of 11
th
and 12
th
verte-
brae [62-6]; (6) adult females with first proximal radial of dorsal fin
located between neural arches of 8
th
and 9
th
vertebrae [63-7*]; (7)
10 or more dorsal-fin rays (males and females) [64-0]; and (8) dark
stripe present on median portion of dorsal fin [137-1].
Composition. Genera Poecilia, “Poecilia” reticulata,
Xiphophorus, Limia, Pamphorichthys, Micropoecilia,
Quintana, Carlhubbsia, and Xenodexia.
Distribution. North, Central and South America.
Quintana + Carlhubbsia
[Clade 109]
Diagnosis. Quintana and Carlhubbsia share the following
not uniquely derived and/or reversed features: (1) parietals
short restricted to the epiotic region, not reaching sphenotic
anteriorly [2-1]; (2) epiotic process long extending beyond
first pleural rib [3-0]; (3) anal-fin proximal radials divergent in
females [66-1]; (4) anal fin asymmetrical in adult males [87-1];
and (5) serrae on R5p [125-1].
Quintana Hubbs
Quintana Hubbs, 1934: 2. Type species: Quintanaatrizona Hubbs,
1934. Type by original designation. Gender: feminine.
Composition. Quintana atrizona, Hubbs.
Distribution. Western Cuba.
Carlhubbsia Whitley
Allophallus Hubbs, 1936: 232. Type species: Allophallus
kidderi Hubbs, 1936. Type by original designation. Gen-
der: masculine.
Carlhubbsia Whitley, 1951: 67. Type species: Allophallus
kidderi Hubbs, 1936. Type by being a replacement name.
Gender: feminine.
Composition. Carlhubbsia kidderi (Hubbs), and C. stuarti
Rosen & Bailey.
Distribution. Carlhubbsia is recorded only from the
drainage of the río Polochic and Lake Izabal (Guatemala),
whereas C. kidderi is known to occur in the río
Champotón (Mexico) and in the drainages of río San
Pedro de Mártir and río de la Pásion (Guatemala) (Rosen
& Bailey, 1959).
Xiphophorus + Xenodexia +Poecilia + Limia +
Pamphorichthys + Micropoecilia + “Poecilia”
Clade [108]
Diagnosis. Members of this clade share the following
uniquely derived and unreversed feature: (1) Lateral wings
on segments of R3 symmetrical [109-1**]. Additionally,
members of this clade can be diagnosed by the following
not uniquely derived and/or reversed features: (1) pelvic-
fin in adult males long, second ray extending beyond the
end of anal-fin base [34-1]; (2) pelvic girdle of males very
posterior, anterior border of basipterygium posterior to the
posterior border of cleithrum [35-0]; (3) anterior tip of
basipterygium in adult males clearly pointed [37-1]; (4) pleu-
ral ribs in adult males almost straight, slightly curving for-
ward and not converging to the same point towards pelvic
girdle [59-0]; and (5) spines on subdistal segments of R3
retrorse [108-2].
Xiphophorus Heckel
Xiphophorus Heckel, 1848: 291. Type species: Xiphophorus
hellerii Heckel, 1848. Type by subsequent designation.
Gender: masculine.
Platypoecilus Günther, 1866: 350. Type species: Platypoecilus
maculatus Günther, 1866. Type by monotypy. Gender:
masculine.
Composition. Xiphophorus alvarezi Rosen, X. andersi
Meyer & Schartl, X. birchmanni Lechner & Radda, X.
clemenciae Alvarez, X. continens Rauchenberger, Kallman
& Morizot, X. cortezi Rosen, X. couchianus (Girard), X.
evelynae Rosen, X. gordoni Miller & Minckley, X. hellerii
Heckel, X. kallmani Meyer & Schartl, X. kosszanderi Meyer
& Wischnath, X. maculatus (Günther); X. malinche
Rauchenberger, Kallman & Mozirot, X. mayae Meyer &
Schartl, X. meyeri Schartl & Schröder, X. milleri Rosen, X.
mixei Kallman, Walter, Morizot & Kazianis, X. montezumae
Jordan & Snyder, X. monticolus Kallman, Walter, Morizot &
Kazianis. X. multilineatus Rauchenberger, Kallman &
Morizot, X. nezahualcoyotl Rauchenberger, Kallman &
Morizot, X. nigrensis Rosen, X. pygmaeus Hubbs & Gor-
don, X. signum Rosen & Kallman, X. variatus (Meek); X.
xiphidium (Gordon).
Distribution. Mexico to Belize (Atlantic drainages).
P. H. F. Lucinda & R. E. Reis
45
PROOFS
Xenodexia +Poecilia + Limia +
Pamphorichthys + Micropoecilia + “Poecilia”
Clade [104]
Diagnosis. Members of this clade share the following
uniquely derived and unreversed feature: (1) wide groove
dorsal to R5 [121-1**].
Additionally, this clade can be diagnosed by the following
not uniquely derived and/or reversed features: (1) preorbital ca-
nal present and entirely closed, bearing four pores [9-0]; (2) adult
females with first proximal radial of dorsal fin located between
neural arches of 10
th
and 11
th
vertebrae [63-2]; (3) absence of a
basal process on first gonactinost [69-0]; (4) fusion of anal-fin
posterior median radials (5
th
to last one) in adult males to respec-
tive proximal radials [77-1]; (5) distal segments of R4p posterior
to serrae deeper than wide [115-1*]; (6) keel on posterior ventral
surface of R5 formed by the projection of R5 toward R4 [119-1*];
and (7) hypural plate completely fused [131-0].
Xenodexia Hubbs
Xenodexia Hubbs, 1950: 8. Type species: Xenodexia ctenolepis
Hubbs, 1950. Type by original designation. Gender: feminine.
Composition. Xenodexia ctenolepis Hubbs.
Distribution. Guatemala, Alta Vera Paz in río Finca, tributary
to the río Negro (río Chixoy) which lower down becomes the
río Salinas of the río Usumacinta basin.
Poecilia + Limia + Pamphorichthys +
Micropoecilia + “Poecilia”
Clade [99]
Diagnosis. Members of this clade share the following uniquely
derived and unreversed features: (1) two (rarely one) well-
developed gonapophyses [48-2**]; and (2) functional
gonapophyses located between vertebrae 13 to 15, but never
on vertebra 16 [49-2**].
Additionally, this clade can be diagnosed by the follow-
ing not uniquely derived and/or reversed features: (1) ligastyle
absent [46-0]; (2) Hollister’s foramen on first or second
gonapophysis [51-1*]; (3) eight dorsal-fin rays (males and
females) [64-2]; (4) nine anal-fin rays in females [65-3*]; and
(5) presence of a palp in subdistal segments of R3 [88-1].
Poecilia + Limia
Clade [93]
Diagnosis. Poecilia and Limia share the following not uniquely
derived and/or reversed features: (1) parietals short restricted to
the epiotic region, not reaching sphenotic anteriorly [2-1]; (2) pos-
terior supraorbital canal closed [5-2]; (3) width of first pelvic-fin
ray in adult males decreasing abruptly at distal portion, distal
slender portion long [41-1]; and (4) second gonapophysis form-
ing an angle of 15-45 degrees relative to vertebral column [54-0].
Poecilia Bloch & Schneider
Poecilia Bloch & Schneider, 1801: 452. Type species: Poecilia
vivipara Bloch & Schneider, 1801. Type by subsequent
designation. Gender: feminine.
Mollienesia Lesueur, 1821: 3. Type species: Mollienesia
latipinna Lesueur, 1821. Type by monotypy. Gender:
feminine.
Alazon Gistel, 1848: X. Type species: Poecilia vivipara Bloch
& Schneider, 1801. Type by being a replacement name.
Gender: masculine.
Allopoecilia Hubbs, 1924: 11. Type species: Girardinus
caucanus Steindachner, 1880. Type by original designa-
tion. Gender: feminine.
Neopoecilia Hubbs, 1924: 11. Type species: Neopoecilia
holacanthus Hubbs, 1924. Type by original designation.
Gender: feminine.
Psychropoecilia Myers, 1935: 311. Type species:
Platypoecilus dominicensis Evermann & Clark, 1906. Type
by original designation. Gender: feminine.
? Lembesseia Fowler, 1949: 267. Type species: Lembesseia
parvianalis Fowler, 1949. Type by original designation.
Gender: feminine.
Curtipenis Rivas & Myers, 1950: 289. Type species:
Mollienesia elegans Trewavas, 1948. Type by original
designation. Gender: masculine.
Composition. Poecilia amazonica Garman, P. boesemani
Poeser, P. butleri Jordan, P. catemaconis Miller, P. caucana
(Steindachner), P. caudofasciata (Regan), P. chica Miller, P.
elegans (Trewavas), P. formosa (Girard), P. gillii (Kner); P.
hispaniolana Rivas, P. koperi Poeser, P. kykesis Poeser, P.
koperi Poeser, P. latipinna (Lesueur), P. latipunctata Meek,
P. marcellinoi Poeser, P. maylandi Meyer, P. mechthildae
Meyer, Etzel & Bork, P. mexicana Steindachner, P. orri
Fowler, P. petenensis Günther, P. salvatoris Regan, P.
sphenops Valenciennes, P. sulphuraria (Alvarez), P. teresae
Greenfield, P. vandepolli Van Lidth de Jeude, P. velifera
(Regan); P. vivipara Bloch & Schneider, P. wandae Poeser,
and P. zonata Nichols.
Distribution. North, Central, and South America.
Limia Poey
Limia Poey, 1854: 382, 390. Type species: Limia cubensis Poey,
1854. Type by subsequent designation. Gender: feminine.
Acropoecilia Hilgendorf, 1889: 52. Type species: Poecilia
tridens Hilgendorf, 1889. Type by monotypy. Gender: femi-
nine.
Odontolimia Rivas, 1980: 29. Type species: Limia grossidens
Rivas, 1980. Type by original designation. Gender: femi-
nine.
Pseudolimia Poeser 2002: 54. Type species: Limia heterandria
Regan 1913. Type by original designation (also mono-
typic). Gender: feminine.
Systematics of the subfamily Poeciliinae Bonaparte
46
PROOFS
Composition. Limia caymanensis Rivas & Fink, L.
dominicensis (Valenciennes); L. fuscomaculata Rivas, L.
garnieri Rivas, L. grossidens Rivas, L. heterandria Regan,
L. immaculata Rivas, L. melanogaster (Günther), L.
melanonotata Nichols & Myers, L. miragoanensis Rivas, L.
nicholsi Myers, L. nigrofasciata Regan, L. ornata Regan, L.
pauciradiata Rivas, L. perugiae (Evermann & Clark), L. rivasi
Franz & Burgess, L. sulphurophilia Rivas, L. tridens
(Hilgendorf), L. versicolor (Günther), L. vittata (Guichenot),
and L. yaguajali Rivas.
Distribution. Cayman Islands, Haiti, Dominican Republic,
Haiti, Jamaica, Cuba, and Venezuela.
Pamphorichthys + Micropoecilia + “Poecilia”
Clade [92]
Diagnosis. Members of this clade can be diagnosed by the
following not uniquely derived and/or reversed features: (1)
absence of an epiotic process [3-3]; (2) absence of a notch on
dentary [16-0]; (3) gonactinost complex approximately per-
pendicular to body longitudinal axis [68-1]; (4) absence of an
anterior convex expansion of second gonactinost [70-1]; (5)
R6a and R6p not fused [126-0]; (6) more distal elements of R6
branches not fused [127-0]; and (7) less than nine caudal-fin
rays in contact with the hypural plate [132-0].
Pamphorichthys Regan
Pamphorichthys Regan, 1913: 1003. Type species: Heterandria
minor Garman, 1895. Type by monotypy. Gender: masculine.
Pamphoria Regan, 1913: 1003. Type species: Cnesterodon
scalpridens Garman, 1895. Type by monotypy. Gender: femi-
nine.
Parapoecilia Hubbs, 1924: 11. Type species: Limia hollandi
Henn, 1916. Type by original designation. Gender: feminine.
Composition. Pamphorichthys araguaiensis Costa, P.
hasemani (Henn); P. hollandi (Henn); P. minor (Garman); P.
scalpridens (Garman).
Distribution. Drainages of the rio Tocantins basin, rio Xingu.
rio Paraguai, rio São Francisco, rio Parnaíba, rio Amazonas,
and rio Tapajós (Figueiredo, 1997).
Micropoecilia + “Poecilia” reticulata
Clade [87]
Diagnosis. Micropoecilia and “Poecilia” share the follow-
ing not uniquely derived and/or reversed features: (1) preor-
bital canal present and partially closed bearing two upper
pores and a lower deep groove [9-1]; (2) anterior tip of
basipterygium in adult males approximately triangular and
round [37-0]; (3) first gonapophysis forming an angle of 5-15
degrees relative to vertebral column [53-2]; (4) second
gonapophysis forming an angle of zero-15 degrees relative to
vertebral column [54-3]; (5) distal portion of third and fourth
gonactinosts separate, except by tip of gonactinost [74-2];
(6) gonactinost 5 fused to gonactinost complex [81-1]; (7)
spines on subdistal segments of R3 retrorse [108-2]; (8) distal
segments of R4p posterior to serrae wider than deep [115-0];
(9) four to seven subdistal retrorse spines on R4p [116-2];
(10) absence of a keel on posterior ventral surface of R5 formed
by the projection of R5 toward R4p [119-0]; and (11) hypural
plate partially fused with an elongate aperture [131-1].
Micropoecilia Hubbs
Micropoecilia Hubbs, 1926: 73. Poecilia vivipara parae
Eigenmann, 1894. Type by original designation. For pur-
poses of the type species, the subspecies parae is el-
evated to species level; type not P. vivipara.
Recepoecilia Whitley, 1951: 68. Poecilia vivipara parae
Eigenmann, 1894. Type by being a replacement name. For
purposes of the type species, the subspecies P. v. parae
is elevated to species level; type not P. vivipara. Un-
needed replacement for Micropoecilia Hubbs, 1926 not
preoccupied by Micropoecila Kraatz, 1880 in Coleoptera.
Composition. Micropoecilia minima (Costa & Sarraf), M.
picta (Regan),
M. parae (Eigenmann), M. bifurca
(Eigenmann), and M. branneri (Eigenmann).
Distribution. Drainage of the rio Amazonas, rio Guamá basin,
coastal drainages of Brazil, French Guyana, Guyana, and
Suriname; Trinidad and Tobago.
Remarks. The tree topology placed “Poecilia” reticulata
as the sister group of the Micropoecilia clade and far from
the type-species of the genus, Poecilia vivipara. This fact
brings taxonomic and nomenclatural implications, since the
generic name Poecilia cannot be applied to the reticulata
species. The most parsimonious action should be the merg-
ing of “Poecilia” reticulata with the genus Micropoecilia
Hubbs, 1926.
“Poecilia” reticulata was originally described as Poecilia
reticulata Peters, 1859 and Lebistes poecilioides De Fillipi,
1861 is considered a junior synonym. Lebistes poecilioides,
whose types are lost, was described as a new genus and a
new species. Since the name Lebistes is older than
Micropoecilia it has priority and Micropoecilia should be
considered a junior synonym of Lebistes. However, Poeser &
Isbrücker (2002) suggested, based on evidence on the origi-
nal description of De Fillipi, that L. poecilioides is not equal
to Poecilia reticulata. If it is the case, Lebistes cannot be a
synonym of Micropoecilia. However, Eigenmann (1907)
erected the genus Acanthophacelus for Poecilia reticulata,
thus if Micropoecilia species and “Poecilia” reticulata are
merged under the same generic name, Acanthophacelus
Eigenmann, 1907 has priority over MicropoeciliaHubbs, 1926
and therefore should be resurrected and revalidated. Given
that the types of Lebistes poecilioides are lost and that De
P. H. F. Lucinda & R. E. Reis
47
PROOFS
Fillipi’s description is probably not acurate and reliable, any
nomenclatural action is premature and also depends upon a
more inclusive phylogenetic analysis among members of the
tribe Poeciliini. This analysis could address the question more
thoroughly. At the moment, unless Lebistes poecilioides
types or unquestionable evidence are found Lebistes
poecilioides and Poecilia reticulata should be regarded
Incertae Sedis in Poeciliinae.
Tribe Cnesterodontini Hubbs, new usage
[Clade 111]
Cnesterodontini Hubbs, 1924: 8. Type-genus: Cnesterodon
Garman, 1895
Diagnosis. Cnesterodontins share the following uniquely
derived and unreversed features: (1) males with five pelvic-
fin rays [33-1**]; (2) pedicel in R3 united to R4 [90-1**]; (3)
pedicel at tip of R3 [91-1**]; and (4) membranous appendix at
tip of R3 [92-1**].
Additionally, the tribe Cnesterodontini can be diagnosed
by the following not uniquely derived and/or reversed fea-
tures: (1) absence of an epiotic process [3-3]; (2) posterior
section of posterior remnant of infraorbital system opened
into a groove [7-1]; (3) preorbital canal absent or opened,
forming a very shallow groove [9-2]; (4) post-temporal un-
branched [31-1]; (5) females with five pelvic-fin rays [44-1];
(6) pleural ribs 6, 7, and 8 in adult males curved forward con-
verging to the same point towards pelvic girdle [59-2*]; (7)
eight dorsal-fin rays (males and females) [64-2]; (8) adult males
with anterior process on base of fifth middle anal-fin radial
pointed and upward directed [78-1]; (9) membranous tip ante-
rior to R4 and R5 curved downwards [110-1]; (10) small mem-
branous tip anterior to R4 and R5 [111-1]; (11) four to seven
subdistal retrorse spines on R4p [116-2]; and (12) expanded
distal portion of R6 [128-1].
Composition. Genera Cnesterodon, Phallotorynus, and
Phalloceros.
Distribution. Southern South America.
CnesterodonGarman
Cnesterodon Garman, 1895: 43. Type–species: Poecilia
decemmaculata Jenyns, 1842. Type by original designa-
tion. Gender masculine.
Gulapinnus Langer, 1913: 207. Type–species: Poecilia
decemmaculata Jenyns, 1842. Type by monotypy. Gen-
der masculine.
Composition. Nine species: Cnesterodon decemmaculatus
(Jenyns), C. carnegiei Haseman, C. brevirostratus Rosa &
Costa, C. septentrionalis Rosa & Costa, C. omorgmatos
Lucinda & Garavello, C. hypselurus Lucinda & Garavello, C.
raddai Meyer & Etzel, Cnesterodon n. sp. A (a new species
being described by Lucinda [in prep.]), and Cnesterodon n.
sp. B (a new species being described by Anza et al. [in prep.]).
Distribution. Rio Uruguai drainage, laguna dos Patos sys-
tem, río Negro, río Salado, western drainages of Argentina
and small coastal drainages of Uruguay and Argentina, up-
per portions of the rio Iguaçu and its upper tributaries, the
headwaters of the rio Maquiné in the Tramandaí system, and
the headwaters of rio Itajaí-Açu drainage, upper rio Araguaia
drainage, rio Paranapanema basin, rio Paraguai and lower rio
Paraná drainages, upper rio Iporanga tributary of the rio
Ribeira de Iguape.
Phallotorynus + Phalloceros
Clade [106]
Diagnosis. Phallotorynus and Phalloceros share the fol-
lowing not uniquely derived and/or reversed features: (1)
halves of supraoccipital process bifid, outer half larger than
inner half [4-2*]; (2) ascending process of premaxilla short
and truncate [12-3]; (3) premaxillary symphysis not elevate
[13-0]; (4) gonactinost complex approximately perpendicu-
lar to body longitudinal axis [68-1]; (5) lateral flanges on
ventral portion of anal-fin radial 4 in adult males present and
cleft dorsally forming separate dorsally directed processes
[75-2]; (6) less than nine caudal-fin rays in contact with the
hypural plate [132-0]; (7) dark stripe present on median por-
tion of dorsal fin [137-1]; and (8) dark patch of pigmentation
along R3 [138-1].
Phalloceros Eigenmann
Phalloceros Eigenmann, 1907: 427, 431. Type species:
Girardinus caudimaculatus Hensel, 1868. Type by origi-
nal designation. Gender: masculine.
Composition.Phalloceros caudimaculatus (Hensel), plus 21
new species being described by Lucinda (in prep.).
Distribution. Upper rio Tocantins drainage, coastal and in-
land drainages from Bahia (Brazil), southward to Uruguay,
Argentina, and Paraguay.
Phallotorynus Henn
Phallotorynus Henn, 1916: 126. Type species: Phallotorynus
fasciolatus Henn, 1916. Type by monotypy. Gender: mas-
culine.
Composition. Phallotorynus fasciolatus Henn, P. jucundus
Ihering, P. victoriae Oliveros, plus three new species being
described by Lucinda et al. (in prep.).
Distribution. Rio Paraíba do Sul drainage and rio Paraná-
Paraguay drainage.
Systematics of the subfamily Poeciliinae Bonaparte
48
PROOFS
Discussion
The classification of the subfamily Poeciliinae. The classifi-
cation of the subfamily Poeciliidae has suffered many modifi-
cations even before the establishment as a distinct subfamily
(see historical accounts). Preterit authors attempted to define
the Poeciliinae in the absence of a cladistic hypothesis.
Eigenmann (1907) and Regan (1911) were the first to define
the Poeciliinae based on exclusive characters: absence of
exoccipital condyles and male anal-fin rays modified into a
gonopodium. Non-cladistic diagnoses and non-cladistic
speculations of relationships were provided by different au-
thors (e.g. Hubbs, 1924, 1926; Howell Rivero & Hubbs, 1936;
Rosen, 1952; Rosen & Gordon, 1953; Rosen & Bailey, 1959;
Rosen, 1967; Costa, 1991; Meyer, 1993; Meyer & Etzel, 1996,
1998; Meyer & Radda, 2000; Meyer & Etzel, 2001a, 2001b;
Meyer & Schartl, 2002). Non-cladistic classifications were also
proposed (e.g. Hubbs, 1924, 1926; Rosen & Bailey, 1963).
However, these classifications did not necessarily communi-
cate phylogenetic relationships.
Hubbs (1924) was the first to present a formal classifica-
tion for the subfamily Poeciliinae. Hubbs’ (1924) classifica-
tion is somewhat congruent with the classification proposed
herein. The naturalness of the groups Tomeurini, Alfarini,
Gambusiini, Poeciliopsinae, and Poeciliinae proposed by
Hubbs are corroborated by our phylogenetic analysis. The
classification of Hubbs (1924) has suffered subsequent, little
modifications, which mainly accounted for the inclusion of
new genera on the tribes (Hubbs, 1926; Hubbs, 1931; Hubbs,
1936; Hubbs & Howell Rivero, 1936; Rosen, 1950 and Rosen,
1952). The most striking partial congruence among these clas-
sifications and the classification herein proposed is that of
Poeciliopsinae Hubbs and our tribe Girardinini. Hubbs (1924)
suggest that Leptorhaphis [=Poeciliopsis], Phallichthys,
Xenophallus, Poeciliopsis, and Phalloptychus were closed
related, creating the name Poeciliopsinae for these genera.
Hubbs (1926) added Poecilistes [=Poeciliopsis], and
Aulophallus [=Poeciliopsis] to this group. The subfamily
Poeciliopsinae proposed by Hubbs (1924, 1926) is very simi-
lar to our tribe Girardinini, differing only by the inclusion of
the genus Girardinus in our tribe (we employed the name
Girardinini Hubbs, 1924: 9; which has page priority over
Poeciliopsini Hubbs, 1924: 10, although this is not necessar-
ily obligatory - see Article 64 on fourth edition of the ICZN,
1999). Hubbs (1924, 1926) allocated Girardinus in his new
tribe Girardinini together with Glaridichthys and Toxus (jun-
ior synonyms of Girardinus).
Contrarily to us, Rosen & Bailey (1959) sustained the idea
that Poeciliopsiinae is not a natural group and should there-
fore be dissolved. They discussed the gonopodium asymme-
try of the Poeciliopsinae and believed that characters related
to gonopodium folding were highly adaptive and may had
evolved independently more than once within the subfamily.
Theses authors also supposed that Poeciliopsis and
Phallichthys form a natural group and that Carlhubbsia is
related to Quintana and Giradinus. Our results are contrary
to Rosen & Bailey’s (1959) assumptions, except for the state-
ment that Carlhubbsia is related to Quintana. The dissolu-
tion of the Poeciliopsinae was formally put forward by Rosen
& Bailey’s (1963) new poeciliin classification. Members of
this group were allocated in two different tribes: Phalloptychus
in Cnesterodontini, Poeciliopsis, Phallichthys, and
Xenophallus in Heterandriini. Expectedly Carlhubbsia,
Quintana, and
Giradinus were unified in a tribe Girardinini.
The classification of Rosen & Bailey (1963) also differs
from ours in the position of genera Alfaro and Priapella in
the tribe Poeciliini and the position of Brachyrhaphis in the
tribe Gambusiini. We proposed Alfaro, Brachyrhaphis, and
Priapella as type-genera of monotypic, basal tribes. Addi-
tionally, the tribes Girardinini and Heterandriini of Rosen &
Bailey (1963) differ substantially from ours. This incongru-
ence can be explained by the fact that diagnoses of supra-
specific groups by Rosen & Bailey (1963) (and also the state-
ments of Rosen & Bailey, 1959) were made in the absence of
cladistic methodology and therefore not necessarily reflect
phylogenetic relationships. Rosen (1979: 278-279) was aware
of this and recognized the fragility of Rosen and Bailey’s
classification, when stated: “ (…) it should be noted that the
diagnoses of genera and other supra-specific groups in Rosen
and Bailey were designed as phenetic statements of overall
similarity. In short, little attention was paid to find unique
characters for defining the taxa and only an implicit effort was
made to interpret the different states of a character as primi-
tive or derived. (…).”
The advent of cladistic methodology has brought some
improvement towards the comprehension of poeciliine rela-
tionships. Some authors addressed the relationships of smaller
groups of the Poeciliinae (e.g. Rosen, 1979; Rauchenberger,
1989; Rosa & Costa, 1993; Rauchenberger et al.,1990; Meyer
et al., 1994; Mojica et al., 1997; Rodriguez, 1997; Ptaceck &
Breden, 1998; Marcus & McCune, 1999; Breden et al., 1999;
Hamilton, 2001; Mateos et al., 2002; Poeser, 2003; Kallmann
et al., 2004), whereas others coped with higher taxa (Parenti,
1981; Meyer & Lydeard, 1993; Costa, 1996, 1998; Ghedotti,
2000). However, no phylogenetic study has simultaneously
analyzed the relationships among representatives of all
poeciliine genera. The only comprehensive study is the clas-
sic revision of the “Poeciliidae” by Rosen & Bailey (1963),
which did not deal with cladistic methodology.
The proposed phylogeny and classification attempted to
include representatives of all poeciliine genera. They are con-
sistent with the results of the phylogenies proposed by
Breden et al. (1999), and Figueiredo (1997) for Poecilia and
its allies. On the other hand, the phylogenetic hypothesis of
Rauchenberger (1989), Rodriguez (1997), and Ghedotti (2000)
are partially congruent with the present results. Probably part
of the incongruence can be explained by the fact that these
phylogenetic studies had been performed for different sub-
units of the Poeciliinae.
Rauchenberger (1989) proposed: (1) Gambusia and
Belonesox as sister groups and (2) that clade as the sister-
group of Brachyrhaphis. The former proposal, but not the
P. H. F. Lucinda & R. E. Reis
49
PROOFS
later was corroborated by our results. Our hypothesis sup-
port Gambusia and Belonesox more closely related to
Pseudopoecilia, Neoheterandria, and Scolichthys than to
Brachyrhaphis. Thus, we modified the usage of the name
Gambusiini sensu Rauchenberger (1989), employing it to re-
fer to a clade composed of Gambusia, Belonesox,
Pseudopoecilia, Neoheterandria, and Scolichthys.
Costa (1991) suggested the monophyly of the group em-
bracing Pamphorichthys, Poecilia, Limia, Xiphophorus,
Cnesterodon, Phalloceros, Phallotorynus, Phalloptychus,
Priapichthys, Poeciliopsis, Priapella, Quintana,
Carlhubbsia, Xenodexia, and Phallichthys supported by four
putative synapomorphies (see our characters 12, 15, 26 and
28). This is partially corroborated by our results. The current
phylogenetic analysis supports these features as a uniquely
derived and unreversed synapomorphies for the supertribe
Poeciliini [Clade 119], which in addition to the genera above
(except Priapichthys and Priapella), also comprises
Girardinus, Xenophallus, and Micropoecilia.
Rodriguez’s (1997) conclusions are also consistent with ours.
This author was unable (like us) to find shared derived charac-
ters uniting Alfaro and Priapella together with Xiphophorus,
Poecilia (including Micropoecilia), Limia, and Pamphorichthys
as suggested by Rosen & Bailey (1963). Rodriguez (1997) hy-
pothesized a monophyletic Poeciliini composed of Xiphophorus,
Poecilia (including Micropoecilia), Limia, and Pamphorichthys
on the basis of three synapomorphies: (1) a wide groove dorsal
to R5; (2) long pelvic-fin in adult males, second ray surpassing
the end of anal-fin base; and (3) compressed external teeth. How-
ever, none of the synapomorphies above are exclusive to this
fish assemblage and are not useful as diagnostic characters.
The presence of a wide groove dorsal to R5 is also observed in
Xenodexia and herein interpreted as a uniquely derived and
unreversed synapomorphy for a broader clade containing
Xenodexia, Poecilia, Limia, Pamphorichthys, Micropoecilia,
and “Poecilia” [Clade 104]. A long second pelvic-fin ray sur-
passing the end of anal-fin base is also present in Xenophallus
and therefore is hypothesized to have been independently ac-
quired by Xenophallus and by the ancestor of a clade compris-
ing Xiphophorus, Xenodexia, Poecilia, Limia, Pamphorichthys,
Micropoecilia, and “Poecilia” [Clade 108]. Presence of com-
pressed teeth is herein interpreted as synapomorphic for a more
encompassing clade, i. e. the supertribe Poeciliini [Clade 119]
(with a reversal in Cnesterodon brevirostratus + C.
septentrionalis clade).
Ghedotti (2000) was the first to propose a phylogenetic
classification scheme for the group, despite relying his con-
clusions on a limited sample (representatives of twelve
poeciliine genera). This is understandable, however, given
that the main focus of Ghedotti’s paper was the relationships
among members of a more encompassing group: the super-
family Poecilioidea. Except for the recognition of Alfaro and
Priapella as basal taxa, Ghedotti’s (2000) hypothesis is only
partially harmonious with ours. It differs mainly by the place-
ment of Tomeurus as a highly derived poeciliine (in the tribe
Cnesterodontini), whereas our results support the assump-
tion that Tomeurus is the most basal poeciliine; the sister-
group of the remaining members of the subfamily. The phylo-
genetic position of Tomeurus will be discussed in more detail
below in the next section.
The goal of phylogenetic analyses is to continuously
improve our knowledge of relationships. The purpose of a
written classification is to communicate phylogenetic rela-
tionships. The proposed hypothesis of relationships and clas-
sification is only preliminary. Much of continued effort on
taxonomy and phylogeny are still required towards a fully
understanding of poeciliine history.
The Cnesterodontini. Questions related to the definition of the
Cnesterodontini have long been based on the pre-cladistic diag-
noses proposed by Hubbs (1924; 1926) and Rosen & Bailey (1963).
The tribe Cnesterodontini as originally erected by Hubbs (1924)
was composed of the genera Phalloceros, Cnesterodon,
Phallotorynus, and Diphyacantha. The cnesterodontins were
defined as poeciliines bearing “terminal segment of ray 3 forming
a more or less specialized process” (Hubbs, 1924: 9). Hubbs (1926)
added Darienichthys to the tribe. Later, Rosen & Bailey (1963)
recognized Diphyacanthaand Darienichthys as junior synonyms
of Priapichthtys and removed them from the Cnesterodontini,
placing them in the tribe Heterandriini. Rosen & Bailey (1963) also
added Phalloptychus to the Cnesterodontini. These authors pro-
vided a diagnosis for the tribe based on the following characters:
(1) absence of parietals; (2) unforked posttemporal; (3) long
gonopodium; (4) unique type of gonopodial appendix at tip of
R3; and (5) sexually dimorphic pleural ribs. Among these charac-
ters only “the unique type of gonopodium appendix” revealed
useful in diagnosing the tribe. The loss of parietals is not unique
to Cnesterodontini (sensu Rosen & Bailey, 1963); it also occurs
in Pseudopoecilia, Xenodexia,Pamphorichthys, Micropoecilia,
and Poecilia reticulata. Besides Phalloceros do possess pari-
etals. An unbranched post-temporal is exhibited by Scolichthys,
Cnesterodon, Phallotorynus, and Phalloceros, but
Phalloptychus possess a bifid post-temporal. Long
gonopodium and sexually dimorphic pleural ribs are also not
uniquely derived features shared by the Cnesterodontini
sensu Rosen & Bailey, 1963.
Ghedotti (2000) was the first to attempt a solution for the
recognition or diagnosis of a monophyletic Cnesterodontini.
He recognized Tomeurus as a member of the tribe
Cnesterodontini. Ghedotti (2000: 53) diagnosed the
Cnesterodontini by the following unique and unreversed
synapomorphies: “(1) less than six pelvic-fin rays; (2) male
pelvic girdle far anterior, under pectoral girdle; (3) paired bony
cirri on the third anal-fin ray in males; and (4) dorsoposterior
region of lachrymal very narrow. ”
Ghedotti (2000: 42) argued that “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 sus-
pensorium (T.S. 45, 65, 72, 79, 85). The principle of parsimony
supports the latter hypothesis and the homoplastic reversals
Systematics of the subfamily Poeciliinae Bonaparte
50
PROOFS
in jaw structure possibly may be explained as a developmen-
tal by-product of reduced size. ” However, our results (also
based on global parsimony) support a rather different view, i.
e. the morphology of the oral and pharyngeal jaws supports
the assumption that Tomeurus is the most basal poeciliine,
the sister-group of remaining members of the subfamily. We
have different interpretations for Ghedotti’s (2000) above
evidence for Tomeurus placement in the Cnesterodontini,
which are discussed below.
Ghedotti (2000; character 45) argued that absence of
gonapophyses in Cnesterodon and Tomeurus are evidence
of common exclusive ancestry. However, Cnesterodon spe-
cies do exhibit gonapophyses, although rudimentary (see
character 47). Following Ghedotti (2000; character 79), an-
other putative synapomorphy for Tomeurus + Cnesterodon
clade would be the posteriorly inclined position of the proxi-
mal anal-fin radials in males. Actually, in Tomeurus and
Cnesterodon the gonactinost complex is very inclined back-
wards to an angle smaller than 45
º
relative to the body longi-
tudinal axis (state 68-0). However, on the basis of the present
hypothesis of relationships, the condition in Cnesterodon is
interpreted as a synapomorphic reversal and as plesiomorphic
in Tomeurus. Ghedotti (2000; character 85) also proposed the
lateral processes on ventral portions of sixth, seventh, and
eighth proximal anal-fin radials in adult males contacting each
other as synapomorphic for a clade Tomeurus + Cnesterodon.
We believe these structures are non-homologous in both
genera. These processes are entirely fused in Tomeurus form-
ing a co-ossified structure, whereas in Cnesterodon lateral
processes are present in posterior middle anal-fin rays of males
(and not in the proximal ones), which are fused to their re-
spective proximal radials.
Ghedotti (2000; character 65) proposed the male pelvic
girdle far anterior, under pectoral girdle, as synapomorphic
for the clade ((Tomeurus, Cnesterodon) Phalloceros,
Phallotorynus)). Our analysis revealed the position of the
pelvic girdle is not a useful character in diagnosing the
Cnesterodontini. We identified five states of this character,
which presented several independent acquisitions and re-
versals during the history of the Cyprinodontiformes (see
character 35).
According to Ghedotti (2000; character 72), the pres-
ence of paired bony cirri on tip of R3 is a putative
synapomorphy for a Cnesterodon + Phalloceros +
Phallotorynus + Tomeurus clade. However, the bony cirri
of our Cnesterodontini and Tomeurus seem to represent non-
homologous structures. The bony cirrus or pedicel of R3 of
cnesterodontins is attached to R4 (character 90), whereas in
Tomeurus, such structure is only attached to R3. Besides
the pedicel of cnesterodontins is associated with a membra-
nous appendix, whereas a membranous appendix associ-
ated with R3 is lacking in Tomeurus.
Ghedotti (2000) recognized the reduced number of pelvic
fin-rays (less than six) as synapomorphic for Phalloceros,
Phallotorynus, Cnesterodon, and Tomeurus. Our results sup-
port the hypothesis that such a reduction independently ap-
peared in Tomeurus and in the ancestor of Phalloceros,
Phallotorynus, and Cnesterodon.
The naturalness of a tribe Cnesterodontini composed of
Phalloceros, Phallotorynus, and Cnesterodon, has already
been suggested by Rosen (1959: 496): “(…) the morphologi-
cal details of the osteocranium and some post-cranial bones
of Cnesterodon nevertheless show conclusively that this
genus and two others, Phalloceros and Phallotorynus, form
a tightly knit group (…).” Rosen & Kallman (1959) suggested
that axial and appendicular skeleton resemblances between
Tomeurus and Cnesterodon may be attributed to parallel evo-
lution. These authors experimentally demonstrated that the
loss or extreme reduction of gonapophyses in Tomeurus and
Cnesterodon, respectively, is ontogenetically related to (and
can be attributed) the far anterior position of the male pelvic
girdle. These conclusions are very congruent to our results,
which interpreted the position of the male pelvic girdle and
the loss or extreme reduction of gonapophyses in Tomeurus
and Cnesterodon as independently acquired.
The phylogenetic position of Tomeurus as the sister-group
of remaining poeciliines brings to discussion the evolution of
viviparity in the subfamily. Facultative viviparity in Tomeurus
could be viewed as (1) a preliminary stage of viviparity towards
true viviparity, a stage achieved by the ancestor of all remaining
poeciliines; or (2) as an autapomorphic specialized condition of
viviparity adaptable for different environmental conditions.
Most generic diagnostic characters for Cnesterodon,
Phalloceros, and Phallotorynus provided by different au-
thors (Eigenmann, 1907; Henn, 1916; Rosen & Bailey, 1963;
Oliveros, 1983; Rosa & Costa, 1993) have been corroborated
by this study. These generic characters as well as the rela-
tionships within the genera of the Cnesterodontini will be
discussed in detail on oncoming studies that include the taxo-
nomic revisions and phylogeny of the genera Cnesterodon,
Phalloceros, Phallotorynus, and Phalloptychus (see Intro-
duction for further details).
Biogeography of poeciliines. A clear biogeographic explana-
tion for poeciliine distribution is not available, however, few
considerations can be speculated. The subfamily Poeciliinae
is distributed throughout the Americas. The most basal
poeciliine Tomeurus gracilis occurs in coastal drainages of
northern South America. The tribes Alfarini, Brachyrhaphini,
Priapichthyini, and Priapellini have a mainly Central Ameri-
can distribution, whereas the Heterandriini, and Gambusiini
have also invaded North American drainages. The genera
Priapichthys, Pseudopoecilia, and Neoheterandria are dis-
tributed along Pacific drainages of Panama, Ecuador, Peru,
and Colombia, which may represent an endemism area in
South America. These genera may have differentiated in iso-
lation provided by the elevation of the Andes. Girardinins are
Central American fishes, except for the South American
Phalloptychus. An explanation for the distribution of the
Girardinini requires the assumption of either a broader pret-
erit distribution, followed by extinction events or a
dispersionist hypothesis. Poeciliins are distributed through-
P. H. F. Lucinda & R. E. Reis
51
PROOFS
out the Americas. The most basal poeciliins (Quintana,
Carlhubbsia, Xiphophorus, and Xenodexia) are Central-
American. Poecilia is widespread throughout the Americas,
and Limia is confined to the Caribbean and Venezuela. Mem-
bers of the Clade 92 (Micropoecilia, “Poecilia” reticulata,
and Pamphorichthys) are restricted to South America. It
seems reasonable to suppose that members of the tribe
Poeciliini derived from a widespread ancient poeciliin, which
suffered local differentiations resultant of local vicariant events
giving rise to Quintana (in Cuba), Carlhubbsia (Guatemala
and Mexico), Xiphophorus (Mexico to Belize), and Xenodexia
(Guatemala), Limia (in the Caribbean and Venezuela) and
members of Clade 92 in South America. Micropoecilia and
“Poecilia” reticulata are distributed along northern coastal
South American drainages of Venezuela, Guyana, Surinam,
French Guyana and Brazil (Amapá and Pará States), whereas
Pamphorichthys is distributed in rio Paraguai basin and north-
ern drainages of Brazil (Tocantins, Xingu, São Francisco,
Parnaíba, Amazonas, and Tapajós). Since Micropoecilia +
“Poecilia” reticulata and Pamphorichthys are sister groups,
it can be hypothesized that the ancestor of these fishes in-
habit a huge ancient area in northern South American which
was split in two areas, which represent the current distribu-
tion of Micropoecilia and Pamphorichthys. Cnesterodontins
are endemic to southern South America.
The pattern of distribution of subsets of the subfamily
Poeciliinae may help to tentatively identify putative areas of
endemism for poeciliines in the American continent. Those
areas are: (1) southern North America; (2) Cuba; (3) the Carib-
bean and Venezuela; (4) inland Central America; (5) pacific
drainages of Panama, Colombia, Ecuador and Peru; (6) coastal
drainages of northern South America along the Venezuela,
Guyana, Suriname, French Guyana, and Brazil (Amapá and
Pará States); (7) Paraguay River basin and northern drain-
ages of Brazil (Tocantins, Xingu, São Francisco, Parnaíba,
Amazonas, and Tapajós); and (8) southern South American
drainages.
Acknowledgements
For loan of specimens we are deeply indebted to Anne-
Marie Hine (BMNH), Barbara Brown (AMNH), Barry Chernoff
(FMNH), Carl Ferraris, David Catania (CAS), Edward O. Wiley
(KU), Eldredge Bermingham (STRI), James Albert (UF), Hugo
López (MLP), James Maclaine (BMNH), José Pezzi da Silva
(MCP), José Lima Figueiredo (MZUSP), Kate Shaw (KU),
Marcelo Loureiro (ZVC-P), Mary Anne Rogers (FMNH), Os-
car Shibatta (MZUEL), Osvaldo Oyakawa (MZUSP), Paulo
Buckup (MNRJ), Ricardo Rosa (UFPB), Richard Vari (USNM),
Rigoberto Gonzales (STRI), Scott Schaefer (AMNH), Susan
Jewett (USNM), Sven Kullander (NRM), Vinícius Abilhôa
(MHNCI), William Saul (ANSP), and William Fink (UMMZ).
David Maddison (University of Arizona) provided informa-
tion on Coleopteran taxa. We are grateful to José Pezzi da
Silva for curatorial help in MCP. Edson Pereira (MCP) photo-
graphed specimens. We owe special thanks to Carlos
Figueiredo (MNRJ), Carlos Lucena (MCP), Luiz Malabarba
(MCP / UFRGS), Márcio Martins (FZB), Ricardo Rosa (UFPB),
and a anonymous reviewer for comments and criticism on
various versions of this manuscript. This study was devel-
oped during a doctoral program at the Pontifícia Universidade
Católica do Rio Grande do Sul (PUCRS) and was supported
by the Universidade do Tocantins (UNITINS), the
Universidade Federal do Tocantins (UFT), and the Fundação
Coordenação de Aperfeiçoamento de Pessoal de Nível Supe-
rior (CAPES, PICDT doctoral fellowship). RER is partially
supported by CNPq through process 305344/87-0.
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Systematics of the subfamily Poeciliinae Bonaparte
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PROOFS
Appendix I
Examined Material. Aplocheiloidei: Fundulidae: Fundulus heteroclitus,
MCP 23910, 9/2*, USA. New York. Cyprinodontoidei: Cyprinodontidae:
Cyprinodon macularius, MCP 23894, 9/2*, USA. Anablepidae: Jenynsia
multidentata, Brazil. Rio Grande do Sul. MCP 11447, 76/1*, rio Camaquã,
and adjacent ponds near Camaquã. - MCP 15473, 33/1*, lagoa de
Tramandaí. Poeciliidae: Aplocheilichthyinae. Aplocheilichthys
spilauchen.USNM 303703, 50+/4*, Cameroon, Lower Ndian system, S
of Korup to seacoast; mangrove channels around Isangele village.
Fluviphylax pygmaeus. MZUSP 29373, 60/4*, Brazil. Rondônia, rio
Machado, lago de Paracaúba. Procatopus gracilis. USNM 303352, 15/
4*, Cameroon, upper Ndian river bordering Korup; stream junction with
Ndian. Poeciliinae. Alfaro huberi, UMMZ 190567, 30/4*, Guatemala.
Zacapa, río Pasabien, 2 km above NNW ca. hwy 9, 10 km WSW of río
Hondo. Belonesox belizanus UF 116114, 7/2*, USA, Florida. UF 34775,
30/2*, United States. Florida, Dade Everglades. Brachyrhaphis
rhabdophora, ANSP 144181, 26/1* and ANSP 144173, 23/1*, Costa
Rica, Puntarenas, Peninsula de Osa, Corcovado National Park. Carlhubssia
kidderi. UMMZ 184619, 30/4*, Mexico. Tabasco, río Chilapa, ca.6
miles by boat NW of Ciudad Pemex landing. Cnesterodon brevirostratus.
MCP 13950, 27/4*, Brazil. Rio Grande do Sul. Bom Jesus, rio Manuel
Leão. MCP 19785, 324/6*, São Francisco de Paula, arroio da Serraria in
road Potreiro Velho - Pró-Mata. Cnesterodon carnegiei. MHNCI 7609,
9/1*, Brazil. Paraná. Almirante Tamandaré, rio Passaúna. USNM 304991,
12/2*, stream in fazenda Lara Maria, tributary of ribeirão Amola-Faca,
near Balsa Nova. Cnesterodon decemmaculatus. MCP 10397, 47/ 4*,
Uruguay, Sierra Mahoma. MCP 11404, 30/4*, Quaraí, arroio Garupá,
between Quaraí and Alegrete. Cnesterodon hypselurus. MCP 12593, 31/
6*, Brazil. Paraná. Jaguariaíva, rio Cilada, bridge of road BR 151, rio
Paranapanema drainage. – MCP 22744, 9/2*, paratopotypes. Cnesterodon
omorgmatos. MCP 22742, 21/4*, Brazil. Paraná. Pinhão, rio das Torres.
Cnesterodon raddai. UMMZ 207503, 12/1*, Paraguay. Misiones, small
stream at bridge on dirt road to Ayolas (ca. 51.2 km S of San Patricio), ca.
2 km ENE of Ayolas; río Paraná drainage. 27
º
22’12”S 56
º
43’12”W.
Neembucu. UMMZ 207748, 30/2* of 102, Paraguay. Misiones, small
pool on S side of dirt road between San Ignacio and Pilar, ca. 114 km
WNW of San Ignacio (ca. 8 km WNW of San Juan) 26
º
40’24”S
57
º
55’12”W. Cnesterodon septentrionalis. MZUSP 41370, 25/1*,
paratypes, Brazil. Mato Grosso. Alto Araguaia, swamp near córrego do
Rancho fountainhead. MZUSP 69687, 21/4* of 42, Alto Araguaia, ribeirão
do Sapo, 464.04 km from ferrovia Ferronorte. Cnesterodon n. sp. A,
MZUSP 54978, 12/2*, Brazil. São Paulo. Apiaí, creek on headwaters of
rio Iporanga, inside Mineradora Oxical, 24
º
24’42”S 48
º
39’25” W. Gam-
busia holbrooki. MZUSP 46100, 7/2*, USA, North Carolina, Lumber
river at US Route 74 crossing near Boardman, Robeson-Columbus coun-
ties (border). Girardinus uninotatus, AMNH 96301, 30/4*, Cuba. Pinar
Del Río, río Taco Taco. Heterandria jonesii, UMMZ 210999, 30/4*,
Mexico. Veracruz, spring on Rancho Santa Anita, 6 km NNW of Potrero
Viejo. Limia vittata, AMNH 96567, 30/4*, Cuba. Sancti Spiritus, Cuyuji,
17 km Sof Iznaga. Micropoecilia branneri, MCP 22040, 5/1*, Brazil
Pará. Castanhal, igarapé Apeú, on road Belém - Brasília. Micropoecilia
sp., MCP 22055, 209/4*, Brazil. Maranhão. Peritoró, rio Peritoró.
Neoheterandria tridentiger. STRI 2335, 10/4*, Panamá. río Anton, stream
through thermal. STRI 2351, 10, río Santa Maria, Quebrada on the Divisa.
Pamphorichthys hollandi, MCP 16851, 104/4*, Brazil. Bahia. Guanambi,
rio da Olaria, 30 km E of Malhada, on road to Guanambi. Pamphorichthys
scalpridens, MCP 15386, 300/6*, Brazil. Pará. Itaituba, braço morto of
rio Tapajós, at bairro da Piracuna, Itaituba. Phallichthys fairweatheri,
UMMZ 196466, 30/4*, Mexico. Quintana, laguna Caobas, 2 km S of
Hwy 186, from which turnoff is 3 km and of road to San Antonio.
Phalloceros caudimaculatus. MCP 20158, 16/4*, Sapiranga, arroio
Feitoria. Phalloceros n. sp. A. MCP 19831, 20/4*, Brazil. Rio Grande do
Sul, Mariana Pimentel, stream affluent to arroio dos Ratos, at Horto
Florestal Mariana. Phalloceros n. sp. B. MCP 30548, 14/4*, Brazil,
Bahia, Prado, fourth stream flowing E at 26 km from Prado on road from
Prado to Cumuruxatiba. Phalloceros n. sp. C. UFPB 2214, 14/1* of 37,
Brazil. Bahia. Porto Seguro, unnamed stream affluent to rio Camurigi, rio
João de Tiba drainage, Estação Ecológica Pau Brazil, 15 km NW from
Porto Seguro. UFPB 2748, 22/2* of 68, Brazil. Bahia. Porto Seguro,
unnamed stream, rio João de Tiba drainage, Estação Ecológica Pau Brazil,
15 km NW from Porto Seguro. Phalloceros n. sp. D. USNM 330339, 5/
2*, Brazil. Goiás, Aruanã, rio Araguaia drainage. Phalloceros n. sp. E.
MCP 20585, 26/2*, Brazil, Rio de Janeiro, Cachoeiras de Macacu, rio
Macacu, ca. 1.5 km SE of Macacu, near road RJ 116. 22
º
29’9”S
42
º
39’34”W. Phalloceros n. sp. F. MCP 30512, 6/4*, Brazil, Rio de
Janeiro, Sapucaia, rio São Francisco, tributary to rio Paquequer fazenda
São Francisco de Paula. Phalloceros n. sp. G. MCP 30509, 6/2*; Brazil.
Rio de Janeiro, rio Parati-Mirim, near km 202 of BR 101, upstream Vila
do Patrimônio. Phalloceros n. sp. H. MCP 12603, 87/4*, Brazil, Rio de
Janeiro, Parati, rio São Roque nearby road BR 101. Phalloceros n. sp. I.
MCP 13735, 11/4*, Brazil, Santa Catarina, rio Lindo, affluent to rio
Cubatão, ca. 1 km from Trevo on road Pirabeiraba - Campo Alegre (SC-
301), Pirabeiraba. Phalloceros n. sp. J. MCP 29270, 256/6*, Brazil, Rio
Grande do Sul, Itati, creek ca. 200 m from arroio do Padre. Phalloceros
n. sp. L. MCP 20579, 117/4*, creek on Iporanga exit to Apiaí, Apiaí, São
Paulo, Brazil. Phalloceros n. sp. M. MCP 12549, 4/2*, Brazil, Paraná,
Paranaguá, rio Jacareí at km 18 on road BR 277. Phalloceros n. sp. N.
MCP 25561, 3/1*, Brazil, Paraná, Antonina, flooded areas on margins of
the PR 340. MCP 27005, 2/1*, Antonina, rio Dois de Fevereiro.
Phalloceros n. sp. O. MCP12197, 87/6*, Brazil, São Paulo, Juquiá, creek
on road BR 116, near Juquiá (affluent to rio Juquiá). Phalloceros n. sp. P.
MCP 30511, 6/2*, Brazil, Minas Gerais, Lagoa Santa, córrego do Jaque,
tributary of the left margin of the rio das Velhas. Phalloceros n. sp. Q.
MHNCI 6151, 8/2*, Brazil, Paraná, Paranaguá, creek at Praia do Forte,
Ilha do Mel. MHNCI 6262, 4/2*, Brazil, Paraná, Pontal do Paraná,
Balneário de Shangri-lá. Phalloceros n. sp. R. MZUSP 35422, 7/2*,
Brazil, Santa, Catarina, Itapoá. Phalloceros n. sp. S. MCP 30510, 6/4*,
Brazil, Rio de Janeiro, Parati, córrego da Toca do Boi, near Condomínio
Laranjeiras. Phalloceros n. sp. T. MZUSP 43467, 26/3*, Brazil. Paraná.
Creek near beach, Guaratuba. USNM 297945, 13/2*, Brazil. Paraná. rio
da Praia, near Guaratuba. Phalloceros n. sp. U. MCP 30023, 90/5*,
Brazil, Espírito Santo, Itarana, córrego Limoeiro, at Praça Oito. UMMZ
215307, 31/4*, Brazil, Espírito Santo, Santa Tereza, córrego at Valsugana
Velha. Phalloceros n. sp. V. MCP 30508, 6/4*, Brazil, Rio de Janeiro,
Sapucaia, rio São Francisco, tributary to rio Paquequer, fazenda São Fran-
cisco de Paula. MCP 20569, 165/10*, Brazil, Rio de Janeiro, Teresópolis,
rio Paquequer, near bridge on km 86 on road BR 116, upstream Represa
Guinle. Phalloptychus iheringii. MCP 11054, 107/8*, Brazil. Santa
Catarina. Tubarão, rio Tubarão and lateral channels near Campo Verde.
MCP 26060, 2/1*, Brazil. Rio Grande do Sul. Rio Grande, Ilha dos
Marinheiros, Porto Rei. MCP 26061, 3/1*, Rio Grande. MCP 26062, 2/
1*, Brazil. Rio Grande do Sul. Rio Grande, Ilha dos Marinheiros, Ponta da
Marambaia. Phalloptychus januarius. MCP 8493, 65/4*, Brazil. Rio de
Janeiro, lagoa de Jacarepaguá.. MNHCI 6174, 15/4*, Brazil. Paraná,
Guaraqueçaba, Ilha de Superagui. MNHCI 6183, 27/4*, Brazil. Paraná.
Paranaguá, Ilha do Mel, Praia de Brasília. Phallotorynus n. sp. A. NRM
42823, 8/2*, arroyo Cambay where crossing about 16 km on road
Caaguazú-Yhú. Phallotorynus fasciolatus. MZUSP 41373, 16/2*, Brazil.
São Paulo. Campo Grande, Paranapiacaba. Phallotorynus jucundus. MCP
25415, 5/2*, Brazil. São Paulo. São Simão, rio Tamanduá headwaters,
affluent to rio Pardo. UFPB 2161, 21/1* of 50, Brazil. São Paulo. São
Simão, rio Tamanduá headwaters, affluent to rio Pardo. Phallotorynus n.
sp. C, MZUSP 69189, 7/1*, Brazil. Mato Grosso, Tacuru, road to Paranhos
on creek at fazenda Sossego, tributary to rio Puitá, affluent to rio Iguatemi.
Phallotorynus n. sp. B. UFPB 2165, 4/2*, Paraguay. San Pedro, río Aguaray
on dirt highway, 2.1 km N of road E to Capitan Bado, affluent to rio Jejui-
Guazu. UMMZ 240156, 80/9*, Paraguay. San Pedro, río Aguaray and
associated run-off ditch at bridge on dirt highway (route 3), 2.1 km N of
junction with road E to Capitan Bado. Phallotorynus victoriae. NRM
42907, 73/4*, Paraguay. Alto Paraná. río Acaray, km 12. “Poecilia”
reticulata. MCP 20592, 103/6*, Brazil. Rio de Janeiro, rio Macacu, ca.
1.5 km SE of Cachoeiras de Macacu. Rio de Janeiro. Poecilia vivipara,
MCP 18118, 39/6*, Brazil. Bahia. Helvécia. córrego Pau Alto (affluent
to rio Pau Alto), on road BR-418, ca. 58 km E of Nanuque. Poeciliopsis
gracilis, MCP 23916, 20/2*, Mexico, Chiapas. Priapella compressa,
UMMZ 210816, 20/4*, Mexico. Chiapas, río Misala just below bridge on
hwy to Agua Azul, at Ruiz Cortinez. Priapichthys annectens, ANSP
163139, 43/4*, Costa Rica. Limon, creek crossing new oil company road
from Bri-Bri to Suretka 4 km W of Bri-Bri. Pseudopoecilia fria, USNM
338706, 10/2*, Ecuador. West tributary to río Baba (Guayas Drainage),
300 m S of Al Balnearia road and 1 km S of San Andres. Quintana
atrizona, AMNH 96396, 30/4*, Cuba. Isla de la Juventud, 16 km SE of la
Fe, Canal de la Isabela, La Reforma District. Scolichthys greenwayi,
AMNH 32887, 30/4*, Guatemala. Alta Verapaz, source and mouth of río
Candelaria Yalicar. Tomeurus gracilis. MNRJ 15180, 20/2* of 41, Brazil.
Amapá. Mazagão. Queimada near Mazagão. MZUSP 26512, 20/4* of
30, igarapé Inó, Furo de Panaquera. AMNH 72910, 26/5* of 31, Guyana.
Cuyuni-Mazaruni. Kartabo point, confluence Mazaruni and Cuyuni Riv-
ers. Xenodexia ctenolepis, AMNH 32137, 30/4*, Guatemala.
Huehuetenango, just above Todos Santos, río Ixcan at channel on E side
of island. Xenophallus umbratilis, ANSP 169120, 30/4*, Costa Rica.
Guanacaste, North shore of lago Arenal about 22 km E of Nuevo Arenal.
Xiphophorus sp., MNRJ 17932, 33/4*, Brazil. Rio de Janeiro. Rio Claro,
ribeirão das Lages or rio Pires, near fazenda Lorena.
P. H. F. Lucinda & R. E. Reis
57
PROOFS
Appendix II. Character state data matrix of 145 characters for 65 poeciliine taxa and 6 outgroup taxa. All transformation series
were considered unordered. Question marks indicate that character state could not be checked due to lack of available
specimens or structures. Dashes indicate both inapplicable codings and polymorphisms.
0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000
0000000000 1111111111 2222222222 3333333333 4444444444 5555555555 6666666666 7777777777
0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123456789
Phalloceros
n. sp.
B
0013200152 1230010221 –100101111 ?111020100 0100101211 0041020002 111–––0111 0020020001
Phalloceros
n. sp.
C
0013200152 1230010001 01001?1111 ?111010000 0100101211 0042320002 1111220111 0020020001
Phalloceros
n. sp.
D
0013200152 1230010000 11001?1011 1111010?00 0100101211 0041020002 1111210111 0020120001
Phalloceros
n. sp.
G
0013200152 1230010000 01001?1111 1111020000 0100101211 0041020002 1111220111 0020020001
Phalloceros
n. sp.
F
0013200152 1230010000 0100101111 1111020100 0100101211 0041020002 1111220111 0020120001
Phalloceros
n. sp.
E
0013200152 1230010000 0100101111 0111020100 0100101211 0041020002 1111220111 0020020001
Phalloceros
n. sp.
J
0013200152 1230010220 0100101111 0111010100 0100101211 0021020002 11–4220111 0020120001
Phalloceros
n. sp.
I
0013200152 1230010220 01001?1111 0111020100 0100101211 0041020102 1111220111 0020020001
Phalloceros
n. sp.
P
0013200152 1230010000 –100101111 0111020100 0100101211 0041020002 1114210111 0020020001
Phalloceros
n. sp.
H
0013200152 1230010220 01001?1111 1111020100 0100101211 0041020002 11–12–0111 0110120001
Phalloceros
n. sp.
O
0013200152 1230010220 0100101111 0111020100 0100101211 0041020002 117–2–0111 0020020001
Phalloceros
n. sp.
R
0013200152 1230010220 0100101111 01110?0?00 0000–01213 0041020?0? 11712201?? ???0??????
Phalloceros
n. sp.
M
0013200152 1230010000 0100101111 0111020100 010010???? 00???????? 1??4–101?? ???0??????
Phalloceros
n. sp.
Q
0013200152 1230010220 0100101111 0111020100 0100101211 0041020002 1111210111 0020120001
Phalloceros
n. sp.
L
0013200152 1230010220 0100101111 0111020100 01001012–– 0041020002 11112–0111 0020020001
Phalloceros
n. sp.
N
00?3200152 1230010??? 0100??1?1? ?111020100 010010???? 0????????? 1??12–01?? ???0??????
Phalloceros
n. sp.
T
0013200152 1230010220 0100101111 0111021100 0??0101211 0041020002 1111220111 0020020001
Phalloceros
n. sp.
S
0013200152 1230010220 0100101111 0111021100 0100101211 0041020002 1111120111 0020020001
Phalloceros
n. sp.
U
0013200152 1230010000 0100101111 1111021100 0100101211 0041020002 117–220111 0020020001
Phalloceros
n. sp.
V
0013200152 1230010221 0100101111 0111020100 0100101211 0041020002 1111–20111 0020020001
Phalloceros
n. sp.
A
0013200152 1230010000 01001–1111 0111020100 0100101211 0041020002 1111–20110 0110020001
Phalloceros caudimaculatus
0013200152 1230010000 0100101111 0111020100 0100101211 0041020002 111–220111 0020020001
Phalloptychus eigenmanni
?????00102 1????????? ?1???????? ??1??????? ????01???? ?????????? ????–1?1?? ??????????
Phalloptychus januarius
0023000102 1221010010 0100101111 0–12012310 1301113211 0013230011 1100112130 0110100003
Phalloptychus iheringii
0023000102 1221010010 0100101111 0–12012310 1301113211 0013230011 110–112130 0110100003
Phallotorynus fasciolatus
00?3000022 1030010000 ?100??1?1? ?111010000 00–0101211 0001020002 1115220111 1020021021
Phallotorynus jucundus
0023200022 1230010000 1100101111 0111010000 0000101211 0001020002 1111011111 1020022021
Phallotorynus victoriae
0023200022 1230010000 1100101111 0111010000 0000101211 0001020002 1111221111 1020021021
Phallotorynus
n. sp.
A
0023200022 1230010000 0100101111 0111010000 0000101211 0001020002 1014–10111 1220122021
Phallotorynus
n. sp.
B
0023200022 1230010000 0100101111 0111010000 0000101211 0001020002 11–1211111 1220122022
Cnesterodon decemmaculatus
0023000112 1221010000 1100101111 011–030201 021010–300 003___1102 110––10102 0020200112
Cnesterodon carnegiei
0023000112 1221010000 1100101011 0112030201 0210100300 003–––1102 1010210102 0020200112
Cnesterodon brevirostratus
0023000112 10–1010000 1000101011 0112030201 0210100300 003–––1102 1110210102 0020200112
Cnesterodon septentrionalis
0023000112 1021010000 1000101111 0112030201 0200100300 003–––1–02 0063210102 0020200112
Cnesterodon omorgmatos
0023000112 1221010000 1100101011 0112030201 0210100300 003–––1102 1110210102 0020200112
Cnesterodon hypselurus
0023000112 1021010000 1100101011 0111030201 02?0100300 003–––1102 11––210102 0020200112
Cnesterodon
n. sp.
B
0023000112 1251010000 1100101111 0111030201 0210100300 003–––1102 1001210102 0020000112
Cnesterodon
n. sp.
A
0023000112 1021010000 11001?1011 0111030201 0210100300 003–––1102 1111210102 0120000112
Cnesterodon raddai
0023000112 1221010000 1100101111 ?11303???? 0210100?00 00?–––1?00 ?0??2101?? ???0?0????
“Poecilia”reticulata
0022000001 1221011000 0100101111 0010100000 0000000222 01023–0000 1103–10110 1020200104
Micropoecilia
sp 0023000001 1221010000 0100101111 0010100000 00000002–2 01023–0000 1111–30110 1000000101
Micropoecilia branneri
0023000001 1221010000 0100101111 0010100000 0000–00222 01032–0000 11––2–?110 1020200101
Pamphorichthys hollandi
0023100112 1221010000 0100101111 0010120110 0000000222 11032–0000 1142330110 1110100104
Pamphorichthys scalpridens
0023100112 1221010000 1100101111 0010110110 00000002–2 11032–0000 1142330110 0020000104
Poecilia vivipara
0011020000 4221011000 0100101111 0010100100 0100000222 01010–0000 116–230110 0020100104
Limia vittata
0011020000 1221011000 0110101111 0010140100 01000002–– 00010–0000 1142–10120 0110000101
Xenodexia ctenolepis
0022000000 6–21011000 2100101111 0010100100 0000001234 0000110000 1162020020 0000000103
Xiphophorus sp.
0001000001 1221011000 0100101011 0010100100 0000001211 0024400000 1127010121 0020000011
Poeciliopsis gracilis
0010000001 1221011010 0100101111 0010011310 0100001211 0012020001 0000212120 1110100003
Girardinus uninotatus
0001120000 5241110000 1100101111 0010000100 0100002211 0001020001 0011120121 1120020011
Xenophallus umbratilis
0010000000 1221010000 0110101111 0010140100 0100001211 0001020001 0042112120 1110100003
Quintana atrizona
0010000001 1231010000 0100101111 0010010310 0000002211 0000110001 0163121120 0020000031
Phallichthys fairweatheri
00100–0001 1221011000 0100101111 0010010000 0000002211 0000010001 0042020120 1110000013
Carlhubbsia kidderi
0010021001 1221011000 0100101111 0010010000 0000001211 0001110001 1127021121 1020000001
Heterandria jonesi
0002000001 0100100000 0010000001 0010011100 0100002211 0001020001 0060020120 1120300011
Priapichthys annectens
0101011001 3110100000 0010100001 0010010100 0000001211 0000110000 000–010120 1110100011
Neohetrandria tridentiger
0001000002 1010000000 0011100001 0010021000 0100003211 0001020001 0024–10120 1110000011
Pseudopoecilia fria
0022000002 1000100000 1010100001 0010010100 0000000211 0002020001 0011020121 1020000001
Scolichthys greenwayi
0001000112 1010000000 1011100101 0110040010 0000000211 0001020001 1011110111 0020000011
Gambusia holbrooki
0001000132 1210000000 1010100001 0010000100 0000001211 0002340000 1114220120 0011110014
Belonesox belizanus
0100000112 12–0000000 0010100001 0010001100 0000002211 0001010000 1071110121 0111110024
Brachyrhaphis rhabdophora
0001011001 0100100000 0010000001 0010000000 0000001211 0000400000 0063110121 0020000021
Priapella compressa
1103011000 2100000000 0010110001 0010010100 0000002211 0001010001 1100020121 0020000011
Tomeurus gracilis
0013000142 1000000000 001–010001 0013031100 0210200100 0–––––0000 0036420100 0–10200100
Alfaro huberi
0100011000 0100100000 0010100001 0010010000 0000001000 00––––0000 11–4000110 0110100114
Fluviphylax pygmaeus
1023000062 1000000220 10–0100100 0010000000 0000000000 00––––0000 0055–400–0 0000000000
Procatopus gracilis
1123011000 2000000000 0000100000 0010040100 0000000000 00––––0000 00––0000–0 0000000100
Aplocheilichthys spilauchen
0120000000 0000000000 0000000001 0010000000 0000000000 00––––0000 00–02000–0 0000000000
Jenynsia unitaenia
0000022000 0000000000 0100000001 0000000000 0000000000 00––––0000 004?11–0–0 0000000000
Cyprinodon macularius
0003022003 0010000110 1100100101 0000000100 0000001000 00––––0000 00820000–0 0000000000
Fundulus heteroclitus
0000022000 0010000110 0000100000 0100000000 0000000–00 00––––0000 00000100–0 0000000000
Systematics of the subfamily Poeciliinae Bonaparte
58
PROOFS
Appendix II. Cont.
0000000000 0000000000 1111111111 1111111111 1111111111 1111111111 1111
TAXA 8888888888 9999999999 0000000000 1111111111 2222222222 3333333333 4444
0123456789 0123456789 0123456789 0123456789 0123456789 0123456789 0123
Phalloceros
n. sp.
B
0010041000 1111000000 000–0–––00 1300101010 0000001200 0–?0100110 010?
Phalloceros
n. sp.
C
0010001000 1111000000 000–0–––00 1300101010 0000001200 0100100110 0101
Phalloceros
n. sp.
D
0010001000 1111000000 000–0–––00 1000101010 0000001200 0–?0100110 010?
Phalloceros
n. sp.
G
0010041000 1111101010 000–0–––00 1300101010 0000001100 0310000110 0121
Phalloceros
n. sp.
F
0010041000 1111101010 000–0–––00 1300101010 0000001100 0100500110 0121
Phalloceros
n. sp.
E
0010001000 1111101010 000–0–––00 1300101010 0000001100 0100200110 0121
Phalloceros
n. sp.
J
0010001000 1111–120–0 000–0–––00 1300101010 0000001100 0100200110 0112
Phalloceros
n. sp.
I
0010001000 1111200110 000–0–––00 1300101010 0000001200 0110200110 0111
Phalloceros
n. sp.
P
00100?1000 1111211020 000–0–––00 1300101010 0000001200 0100300110 0111
Phalloceros
n. sp.
H
0010041000 1111211020 000–0–––00 1300101010 0000001200 0100400110 0111
Phalloceros
n. sp.
O
0010041000 1111211020 000–0–––00 1300101010 0000001200 01–0200110 0111
Phalloceros
n. sp.
R
0?10041000 1111200110 000–0–––00 1300101010 0000001200 01?0200110 0111
Phalloceros
n. sp.
M
??10041000 1111201020 000–0–––00 1300101010 0000001200 0100200110 0111
Phalloceros
n. sp.
Q
0010041000 1111211020 000–0–––00 1300101010 0000001200 0100200110 0111
Phalloceros
n. sp.
L
0010041000 1111211020 000–0–––00 1300101010 0000001100 0100200110 0111
Phalloceros
n. sp.
N
??10041000 1111200110 000–0–––00 1300101010 0000001200 01?0200110 011?
Phalloceros
n. sp.
T
0010001000 ?111200010 000–0–––00 1300101010 0000001200 0100200110 0101
Phalloceros
n. sp.
S
0010001000 1111200010 000–0–––00 1300101010 0000001200 0100200110 0101
Phalloceros
n. sp.
U
0010001000 1111200010 000–0–––00 1300101010 0000001200 0100200110 0101
Phalloceros
n. sp.
V
0010041000 1111200010 000–0–––00 1300101010 0000001100 0–00–00110 0101
Phalloceros
n. sp.
A
0010001000 1111000000 000–0–––00 1300101010 0000001100 01–0000110 0101
Phalloceros caudimaculatus
00100–1000 1111000000 000–0–––00 1300101010 0000001100 0100200110 0101
Phalloptychus eigenmanni
??1???110? ????–––––0 ????????0? ????1?1??0 0??0?0???? ?201000??? 010?
Phalloptychus januarius
1010041100 0000–––––0 00––0–––00 0000101100 0000002201 0211000010 0100
Phalloptychus iheringii
1010031100 0000–––––0 00––0–––00 0000101100 0000002201 0211000010 0100
Phallotorynus fasciolatus
0010031001 1110–––––0 10?–????00 ??00102000 0000001110 0100001110 1101
Phallotorynus jucundus
0010041001 1110–––––0 101–110100 2100102000 0000001110 0100032110 2101
Phallotorynus victoriae
0010031001 1110–––––0 101–100000 2100102000 0000001110 0100011110 1101
Phallotorynus
n. sp.
A
0010041001 1110–––––0 101–201200 2200102000 0000001110 1100021110 1101
Phallotorynus
n. sp.
B
0010041001 1110–––––0 101–100000 2200102000 0000001110 1100011110 1101
Cnesterodon decemmaculatus
0010041000 1110–––––0 01000–––00 0000102000 001010–210 0010000001 0100
Cnesterodon carnegiei
0010041000 1110–––––0 01010–––00 1100102000 0010101210 0010000001 0100
Cnesterodon brevirostratus
0010041000 1110–––––0 010–0–––00 1100102000 0010101210 0010000001 0100
Cnesterodon septentrionalis
0010041000 1110–––––0 01010–––00 1100102000 0010101210 0000000001 0100
Cnesterodon omorgmatos
0010041000 1110–––––0 01010–––00 1100102000 0010101210 0010000001 0100
Cnesterodon hypselurus
0010041000 1110–––––0 01010–––00 110010–000 0010101210 0010000001 0100
Cnesterodon
n. sp.
B
0010041000 1110–––––0 01000–––00 1100102000 0010101210 0010000001 0100
Cnesterodon
n. sp.
A
0010041000 1110–––––0 01010–––00 1100102000 0010101210 00–0000001 0100
Cnesterodon raddai
??100?1000 1110–––––0 01000–––00 1100102000 001010??10 0––0000001 0101
“Poecilia”reticulata
0110001010 0000–––––0 00––0–––21 0000102000 0100000000 0?00000100 0101
Micropoecilia
sp 0010041010 0000–––––0 00––0–––21 0000102000 0100000000 0100001100 0101
Micropoecilia branneri
0110001010 0000–––––0 00––0–––21 0000102000 0100000000 0100001100 0101
Pamphorichthys hollandi
0010031010 0000–––––0 00––0–––01 0000111001 0100000000 0000000100 0101
Pamphorichthys scalpridens
0010041010 0000–––––0 00––0–––01 0000211001 0100000000 0200000100 0101
Poecilia vivipara
0010001010 0000–––––0 00––0–––21 0000101001 0100001100 0210001100 0101
Limia vittata
0110001010 0000–––––0 00––0–––01 1100111001 1100001100 0010000100 0101
Xenodexia ctenolepis
1010021100 0000–––––0 00––0–––11 0000010001 010010––0– 0010000000 0100
Xiphophorus sp.
0010001000 0000–––––0 00––0–––21 00002010–0 0021001100 0110000110 0101
Poeciliopsis gracilis
1010001100 0000–––––0 00––0–––00 0000101000 0000002200 0–10000?00 0101
Girardinus uninotatus
0010021000 0000–––––1 00––0–––00 0000101010 0000011200 0110000010 0101
Xenophallus umbratilis
1011101100 0000–––––0 00––0–––00 0000000000 0000001200 0200001110 0101
Quintana atrizona
0010001100 0000–––––1 00––0–––00 0000101010 0000011100 0–11000000 0102
Phallichthys fairweatheri
1010021100 0000–––––0 00––0–––00 0000101010 0000001110 0–10000010 0101
Carlhubbsia kidderi
0010021100 0000–––––0 00––0–––02 0000102000 0000011110 0110000000 0101
Heterandria jonesi
0010021000 0000–––––0 00––0–––10 1100101010 0000101100 0110000100 0101
Priapichthy sannectens
0011121000 0000–––––0 00––0–––10 0000101000 0000001100 0–10000100 0101
Neohetrandria tridentiger
0010141000 0000–––––0 00––0–––20 0001101000 0000101210 0–10–00100 0101
Pseudopoecilia fria
0010041000 0000–––––0 00––0–––20 0000101000 0000100000 0010000100 0102
Scolichthys greenwayi
0010001000 0000–––––0 00––0–––00 0001101000 0000101200 0010200110 0101
Gambusia holbrooki
0010021000 0000–––––0 00––0–––10 0011102000 0021001100 0100000100 0101
Belonesox belizanus
0010021000 0000–––––0 00––0–––10 0010102000 0001001000 0200000000 0101
Brachyrhaphis rhabdophora
0010101000 0000–––––0 00––0–––10 0010101000 0000000000 0000001100 1100
Priapella compressa
0010021000 0000–––––0 00––0–––10 0000101010 0000001100 0010000000 0101
Tomeurus gracilis
0010031000 0000–––––0 00––0–––10 0000000000 0000101100 ––00000000 0–00
Alfaro huberi
0010011010 0000–––––0 00––0–––10 0000000000 0000000000 0–10000000 0101
Fluviphylax pygmaeus
0000030000 0000–––––0 00––0–––00 0000000000 0000000000 0000000000 0000
Procatopus gracilis
0000050000 0000–––––0 00––0–––00 0000000000 0000000000 0100000000 0001
Aplocheilichthys spilauchen
0000050000 0000–––––0 00––0–––00 0000000000 0000000000 0200000000 0001
Jenynsia unitaenia
0–000?0000 0000–––––0 00––0–––00 0000000000 0000000000 0?10000000 0100
Cyprinodon macularius
0000010000 0000–––––0 00––0–––00 0000000000 0000000000 0010001000 0001
Fundulus heteroclitus
0000000000 0000–––––0 00––0–––00 0000000000 0000000000 0010000000 0000
P. H. F. Lucinda & R. E. Reis
59
PROOFS
Appendix III.
Synapomorphies (*Uniquely derived; ** uniquely derived and
unreversed. Numbers before colons are node numbers on the cladogram
of Fig. 1)
71: 17-0; 18-0; 63-4.
72: 74-1.
73: 127-1.
74: 65-1; 95-1*; 96-1; 98-2**.
75: 97-1**.
76: 142-1**.
77: 19-1; 134-1.
78: 36-0; 85-4.
79: 30-1; 85-4; 134-0 (134-5).
80: 17-2*; 18-2*.
81: (94-1**); 96-1; 142-2**.
82: 36-1; (94-2*); 127-2.
83: 98-1*; 94-1** (94-2*).
84: 20-0; 71-2**; 74-1; 111-2**; 130-1**.
85: 21-0.
87: (9-1); 37-0; (53-2); (54-3); 63-1 (63-3); 74-2; 81-1; 108-2; 115-
0; 116-2; 119-0; 131-1.
88: 4-1; 7-1; 8-1; (9-2); 35-1 (35-2); 38-1; 50-1**; (53-3); (54-2);
(62-4); 64-3**; 79-4; 85-3 (85-4); 108-0.
89: 134-2.
90: 65-1; 76-2**; 85-4.
91: 33-2.
92: 3-3; 9-1 (9-2); 16-0; 53-2 (53-3); 62-0 62-1 (62-4); 68-1; 70-1;
126-0; 127-0; 132-0.
93: 2-1; 5-2; 41-1; 54-0.
94: 44-1.
95: 30-1; 127-2; 134-1.
96: 20-0; 35-2; 37-1.
97: 66-1; 135-1**.
98: 63-0; 74-2.
99: 46-0; (48-2**); (49-2**); 51-1*; (54-2 (54-3); 64-2; 65-3*; 88-
1.
100: 2-2; 3-3; 7-1; 9-2; 33-2; (36-2**); 40-1**; 41-3**; 43-1**; 45-
1**; 46-3; (53-3); 54-2; 55-3**; 58-1**; 60-1; 61-1; 68-3**;
70-0; 85-3 (85-4); 117-1**; 129-1**; 133-1; 143-0.
101: 2-1; 8-5**; 41-1; 52-4*; (78-0); 93-1**; 111-3*; 116-1; 118-1;
128-0.
102: 7-0; 8-2**; 70-1; 76-1**; (78-2); 85-3; 89-1**; 100-1**; 102-
1**; 104-1**; 110-2**; 136-1; 140-1.
103: 27-0; 62-1; 103-1**.
104: 48-2** (48-3); 49-2** (49-4*);9-0; 63-2; 69-0; 77-1; 115-1*;
119-1*; 121-1**; 131-0.
105: 18-1**; 36-1 (36-2**); 37-3; 38-1; 52-1**; 53-2 (53-3); ; 62-
0; 63-0; 126-2**.
106: 4-2*; (8-2**)(8-5**); 12-3; 13-0; (68-1); 75-2; 78-0 (78-2);
132-0; 137-1; 138-1.
107: (8-1); 35-3; (37-2**); 39-1**; 41-2; 42-1; 46-0; 47-3**; 48-0;
49-0; 52-3**; 57-1; 56-1**; 62-0; (68-0); 69-2**; 77-1; 79-2;
85-4; 101-1**; 122-1**; 124-1; 127-2; 131-0; 139-1**; 143-0.
108: 34-1; 35-0; 37-1; 59-0; 108-2; 109-1**.
109: 2-1; 3-0; 37-0 (37-3); 66-1; (78-3*); 87-1; 125-1.
110: 41-1; 46-1; 65-1; 66-2**; 68-0 (68-1); 74-1; 78-0; 85-0; 118-
0; 127-2.
111: 3-3; 7-1; 8-1 (8-2**)(8-5**); 9-2; 31-1; 33-1**; 37-1 (37-2);
44-1; 59-2*; (64-2); 78-1; 90-1**; 91-1**; 92-1**; 110-1; 111-
1; 116-2; 128-1.
112: 16-1; 20-0; 54-1; 55-1; 62-6; 63-7*; (64-0); 137-1.
113: 2-1; 3-0; 20-0; 62-4; 63-2; 69-0; 72-1; 79-3; 80-1; 87-1; 127-1;
131-2.
114: 23-1**; 35-2 (35-4); 65-1; 113-1; (127-2).
115: 2-2; 37-0; 46-1; 60-1; 61-1; 64-0 (64-2); 70-0; 78-0; 85-0; 118-
0.
116: 42-0; 71-1; 127-2; 138-1.
117: 46-0; 64-0; 85-4; 108-2; 124-1; 127-0 (127-2); 131-0.
118: 7-1; 8-1 (8-3*); 35-0; 37-0; 55-1 (55-4*); 59-0; 60-1; 64-1 (64-
2); 70-0; 72-1; 73-1**; 74-1; 75-1**; 79-4; 112-1; 116-2; 123-
1; 132-0.
119: (12-2**); 13-1*; 15-1**; 21-1*; 22-0; 26-1**; 27-1; 28-1**;
108-0; 137-0.
120: 9-2; (12-1); 118-0; 127-0.
121: 12-1 (12-2**); 10-1; 11-2; 14-0; 20-1; (62-0); 63-1; 64-1.
122: 1-0; 5-0; 55-2*; 62-0 (62-6); 131-1.
123: 46-2; 53-1*; 59-1*; 65-2; 118-1.
124: 37-1; 55-1; 70-1; 85-2; 126-1; 127-1.
125: 3-1*; 9-1; 47-2**; 48-1*; 49-1*; 68-1 (68-2*); 69-1*; 72-2*;
77-0; (79-1**); 114-1*; 116-1; 137-1.
126: 5-1*; 6-1; 11-1*; 14-1; (35-1); 46-1; (68-2*); 78-1; (79-1**)
(79-4).
129: 22-1; 24-1*; [29-1*]; 35-1 (35-3); 65-1; 67-1*; 72-1; 77-1; 82-
1**; 85-1*; 86-1**; 108-1; 132-1; 141-1*; 143-1.
Autapomorphies (* = uniquely derived)
0: 94-0; 98-0.
1: 19-1; 20-0; 53-2; 54-3.
2: 27-0; 65-1; 74-1; 111-0.
3: 37-0; 131-3*; 132-1; (134-0).
4: 74-1; (134-5*).
6: 35-1; 52-2; 63-4; 65-2; 74-1; 85-0; 96-2*; 143-2.
7: 57-1; 85-0; 132-1.
8: 134-3*.
9: 30-1; 71-1; 72-1; 134-4*.
10: 62-7.11: 41-0; 49-3*; 62-7.
12: 95-0.
17: 64-1.
18: 30-1; 62-7.
20: 69-0; 71-1; 72-1.
22: 132-0.
23: (85-4).
24: (85-3).
25: 4-0; 11-0; 63-5*.
26: 64-0; 105-1*; 107-1*; 135-3*; 136-2*; 140-2*.
28: 61-0; 63-4; 66-0; 104-2*; 106-1*; 107-2*; 135-2*.
29: 79-2.
30: 74-2; 110-0; 111-0.
31: 61-0.
33: 27-1; 42-0; 60-0; 61-0; 62-6; 63-3; 132-0.
36: 12-5*; 61-0.
37: 71-1.
38: 33-3; 59-0; 61-0; 143-1.
39: 3-2; 16-1; (62-0); (63-3); 65-1; 79-4.
40: (62-1); (63-1); 72-0; 74-0; 81-0; 136-1.
41: 53-3; 54-2; 136-1.
42: (35-2); 71-1; 72-1; 74-1; (85-3).
43: 20-1; (35-1); 70-0; (85-4); 114-2; 131-2.
44: 10-4*; 79-4; 68-1; 74-1; 115-0; 131-2; 136-1.
45: 22-1; 35-4; 51-0; 62-4; 65-1; 71-1; 72-1; 81-1; 108-0; 110-1;
111-1; 120-1*.
46: 3-2; 10-6*; 20-2*; (48-3); (49-4*); 53-0; 67-0; 72-0; 79-3; 80-1;
85-2; 87-1; 108-1; 114-0; 116-0; 124-1; 137-0; 143-0.
47: 2-0; 27-0; 52-2; 53-4*; 54-4; 55-0; 62-2; 65-1; 78-1; 114-2;
122-2; 123-1; 138-1.
48: 16-1; (36-1); (53-2); 64-2; 138-0.
49: 4-1; 5-2; 9-0; 10-5*; 12-4*; 14-1; 35-0; 75-2; 99-1; 125-1.
50: 9-0; 22-1; 34-1; 35-4; 83-1; 84-1; 114-0; 116-0; 132-0; 136-1.
51: 12-3; 16-0; (37-3); 38-1; 46-2; 53-0; 60-0; 64-1; 69-0; (78-3*);
99-1; 118-1; 133-1; 137-1; 143-2.
52: 16-1; 37-0; 53-0; 55-1; 64-0; 128-1.
53: 5-2; 6-1; (37-0); 62-2; 70-1; 85-2; 109-2*; 116-2; 128-1.
54: 3-2; 24-0; 36-1; 41-1; (62-6); 69-0; 71-1; 74-3*; 110-1; 111-1;
124-1.
55: 10-3*; 12-1; 54-1; 69-0; 71-1; 72-1; 74-1; 83-1; 84-1.
56: 20-0; (35-2); 36-1; 41-1; 46-3; 62-2; 63-4; 69-0; 71-1; 72-1; 84-
1; 128-1.
57: 2-2; 3-2; 12-0; 14-1; 53-2; (127-0); 143-2.
58: 7-1; 8-1; 27-1; 31-1; (35-4); 38-1; 60-1; 68-1; 70-0; 85-0; 134-
2; 138-1.
59: (8-3*); 46-1; 53-2; 54-3; (55-4*); 61-1; 63-4; (64-2); 69-0; 113-
1; 122-2; 127-1.
60: 1-1; 3-0; (8-1); 20-0; 36-1; (55-1); 62-7; (64-1); 71-1; 78-2;
131-2; 137-0.
61: 1-0; 24-0; 35-0; (54-4); 62-6; 63-3; 64-1; 78-2; 84-1; 85-0; 112-
1; 132-0; 136-1; 140-1; 143-0.
62: 0-1*; 1-1; 3-3; 9-0; 10-2*; 14-0; 25-1; 60-1; 61-1; 70-0; 137-0.
63:1-0; 2-1; 3-3; 7-1; 8-4*; 9-2; 10-1; 24-0; 25-1; 33-3; (35-3); 36-
1; 37-1; 41-2; 42-1; 44-2*; 47-1*; 62-3*; 63-6*; 64-4*; 65-2;
74-2; 85-3; 124-1; 126-1; 127-1; 132-0; 143-0.
64: 60-1; 61-1; 63-4; 65-0; (68-1); 71-1; 74-1; (79-4); 88-1.
Systematics of the subfamily Poeciliinae Bonaparte
60
PROOFS
Appendix IV
Fits of individual characters in strict consensus tree (L = 758, CI = 0.35, RI = 0.75; number of steps include transformation
series in the outgroup) (C – character; S – steps; ci; ri).
