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Paulo H. F. Lucinda
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Family Poeciliidae
(Livebearers)
Paulo H. F. Lucinda
Poeciliids can be characterized by: (1) a highly-inserted pectoral fin, (2) pelvic fins that migrate anteriorly during growth,
(3) recessed supraorbital pores 2b through 4a, and (4) pleural ribs on the first several haemal arches as well as a series of
other internal synapomorphies (see details in Parenti, 1981; Costa, 1998; Ghedotti, 2000).
Poeciliids are small and laterally compressed cyprinodontiform fishes. Body form ranges from extremely elongate (e.g.
Tomeurus) to deep-bodied (e.g. Phallichthys, Carlhubbsia). Size is extremely variable ranging from the tiny Fluviphylax
palikur (maximum adult recorded size: 13.9 mm) to the giant of the group, Belonesox belizanus, which reaches 200 mm.
Poeciliids comprise 299 valid species inhabiting the fresh and brackish waters of American and African continents (but are
reported to occur in salt waters in coastal areas).
Species of Poeciliidae occur in North and Middle America, Caribbean, South America to Northern Argentina, Congo basin
and the African rift lakes, Dar es Salaam and Madagascar (Parenti, 1981). Following Ghedotti (2000), fishes of the family
Poeciliidae are included in three subfamilies: Poeciliinae (220 valid species), Procatopodinae (78 valid species)
Aplocheilichthyinae (one species).
The subfamily Poeciliinae is broadly distributed throughout the Americas. Poeciliines are characterized by: (1) the uniquely
derived possession of a gonopodium formed by the modified male anal-fin rays 3, 4, and 5 (Parenti, 1981), (2) internal
fertilization, (3) viviparity (Tomeurus gracilis possess facultative viviparity). Some species with all-female individuals,
their eggs capable of developing when stimulated by the sperm of another species without fertilization. Poeciliinae is the
sister group of the Procatopodinae, a group composed of the South-American Fluviphylax 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 includes well-known aquarium fishes such as the guppies, mosquito fishes, swordtails, platys and the
mollies, being very familiar to the non-scientific public. On the other hand, poeciliines are well known from several
biological standpoints, being object of study for ecologists, anatomists, embryologists and many others biologist
researchers. Notwithstanding, this fish assemblage is disappointingly ill-studied from the perspective of systematics.
Intrageneric diversity and intergeneric relationships of the Poeciliinae are poorly known, regardless of its huge distribution
range and notoriety. Similarly, phylogenetic hypotheses for most of genera are still wanting. Taxonomic revisions and
phylogeny hypotheses have provided insights 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; Ghedotti, 2000)
or have dealt with higher taxa (Costa, 1996 and 1998; Parenti, 1981). The only comprehensive study is the classic revision
of Poeciliidae by Rosen & Bailey (1963), which did not deal with cladistic methodology. Nonetheless, Rosen & Bailey
(1963) is the basis for current inner classification of Poeciliinae. Later, Parenti & Rauchenberger (1989) modified the
classification of Rosen & Bailey (1963) in order to accommodate it into the taxonomic rank of subfamily proposed by
Parenti. Following Rosen & Bailey (1963) and Parenti & Rauchenberger (1989), Tomeurus alone is the supertribe
Tomeurini. The remaining genera form the supertribe Poeciliini, which is subdivided in the tribes Poeciliini,
Cnesterodontini, Gambusini, Scolichthyini, Girardinini, Heterandrini and Xenodexini. Later, Ghedotti (2000) proposed
another Poeciliinae classification based in his phylogenetic study of the Poecilioidea despite the fact that only 12 genera
have been used in his cladistic analysis.
ALFARO
Petalosoma Regan, 1908: 458. Type species: Petalosoma
cultratum Regan, 1908. Preoccupied in Coleoptera. Type by
monotypy. Gender: neuter.
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 replacement
558
BELOESOX
Belonesox Kner, 1860: 419, 422. Type species: Belonesox
belizanus Kner, 1860. Type by monotypy. Gender: masculine.
BRACHYRHAPHIS
Brachyrhaphis Regan, 1913a: 997. Type species: Brachyrhaphis punctifer (Hubbs, 1926)
Trigonophallus punctifer Hubbs, 1926: 49. Type locality: Guibari
Gambusia rhabdophora Regan, 1908. Type by Creek, a tributary of the Rio Cricamola below Conquantu, on the
monotypy. Gender: feminine. See Mojica et al. Caribbean slope of western Panama. Holotype: UMMZ 72573.
(1997) for phylogenetic relationships inferred Maximum length: 3 cm TL
Distribution: Central America: Panama.
from DNA sequences.
Countries: Panama
Trigonophallus Hubbs, 1926: 48. Type species: Trigonophallus
punctifer Hubbs, 1926. Type by original designation. Gender:
masculine.
Plectrophallus Fowler, 1932: 384. Proposed as new subgenus of
Panamichthys Hubbs. Type species: Panamichthys tristani
Fowler, 1932. Type by original designation. Gender: masculine.
559
CARLHUBBSIA
Allophallus Hubbs, 1936: 232. Name preoccupied in Diptera.Type
species: Allophallus kidderi Hubbs, 1936. Type by original
designation. Gender: masculine.
Carlhubbsia Whitley, 1951: 67. Type species: Allophallus kidderi
Hubbs, 1936. Type by being a replacement name. Gender:
feminine.
CESTERODO
Cnesterodon Garman, 1895: 43. Type species: Poecilia
decemmaculata Jenyns, 1842. Type by original designation.
Gender: masculine. See Rosa & Costa (1993) for taxonomic
revision.
Gulapinnus Langer, 1913: 207. Type species: Poecilia
decemmaculata Jenyns, 1842. Type by original designation.
Gender: masculine.
560
GAMBUSIA
Gambusia Poey, 1854: 382. Type species: Gambusia punctata
Poey, 1854. Type by subsequent designation. Gender: feminine.
Gambusia alvarezi Hubbs & Springer, 1957: 310, fig 14. Type
locality: El Ojo de San Gregorio, Chihuahua, Mexico. Holotype:
UMMZ 168979.
561
Gambusia atrora Rosen & Bailey, 1963: 102, fig. 43. Type
locality: ferry acrosss Ro Axtla to a shallow area 150 yards
upstream, Xilitla, San Luis Potos, Mexico. Holotype: UMMZ
179999.
Maximum length: 3 cm TL
Distribution: North America: Mexico.
Countries: Mexico
Gambusia hurtadoi Hubbs & Springer, 1957: 307, fig. 13. Type
locality: El Ojo de la Hacienda Dolores, seven miles south of
Jiminez, Chihuahua, Mexico. Holotype: UMMZ 168975.
Maximum length: 3.1 cm TL
Distribution: North America: Chihuahua (only known from type
locality and proximities).
Countries: Mexico
Gambusia beebei Myers, 1935: 305, fig. 274. Type locality: tang
de Miragone, Haiti, S. W. end of lake, from Aux Cayes road.
Neotype: USNM 203161.
Maximum length: 3.3 cm SL
Distribution: Caribbean Islands: tang de Miragoane in Tiburon
Peninsula, southwestern Haiti.
Countries: Haiti
562
563
Gambusia wrayi Regan, 1913a: 988, pl. 99 (fig. 3-4); fig. 168b.
Type locality: Jamaica. Lectotype: BMNH 1912.12.20.7,
designated by Fink (1971a: 61).
Gambusia gracilior Regan, 1913a: 989, pl. 99 (fig. 5-6); Fig. 168c.
Type locality: Jamaica. Syntypes: (13) BMNH 1969.9.8.1-14 (14
now 6).
Maximum length: 3.22 cm SL
Distribution: Caribbean Islands: Jamaica Island.
Countries: Jamaica
Species inquirenda
Heterophallus milleri Radda, 1987a: 127, Fig. 2. Type locality:
Ro Teapa at Teapa, W side of river, under bridge; Tabasco state
[Mexico]. Holotype: UMMZ 184705.
GIRARDIUS
Girardinus Poey, 1854: 383. Type species: Girardinus metallicus
Poey, 1854. Type by monotypy. Gender: masculine.
Glaridodon Garman, 1895: 40. Type species: Girardinus
uninotatus Poey, 1861. Type by original designation. Not
available name preoccupied in fossil Reptilia. Gender: masculine.
Glaridichthys Garman, 1896: 232. Type species: Girardinus
564
HETERADRIA
Heterandria anzuetoi Rosen & Bailey in Rosen, 1979: 324, fig. 11.
Type locality: Ro Achuelo, south side of Gualn, Ro Motagua
drainage, Zacapa, Guatemala. Holotype: AMNH 36319.
Maximum length: 6 cm TL
Distribution: Central America: Basin of Motagua River in
Departments of El Progreso, Zacapa, Chiquimula, and Izabal,
Guatemala, and Copn, Honduras, and basin of Lempa River in
the Departments of Chiquimula and Jutiapa, Guatemala.
Countries: Guatemala, Honduras
565
LIMIA
Limia Poey, 1854: 383. Type species: Limia cubensis Poey, 1854.
Type by subsequent designation. Gender: feminine. The genus
Limia originally proposed by Poey (1854), was later listed as a
subgenus of Poecilia by Rosen and Bailey (1963) and Parenti
and Rauchenberger (1989). Rivas (1978; 1980) ressurected the
genus. spliting in sugenera and species-groups. Farr (1984)
survey of courtship display behavior. Chambers (1987) studied
the structural variation of gonopodium. Hamilton (2001) studied
the intrageneric relationships based on molecular data. Rodriguez
(1997) discussed the phylogentic position.
Acropoecilia Hilgendorf, 1889: 52. Type species: Poecilia tridens
Hilgendorf, 1889. Type by monotypy. Gender: feminine.
Odontolimia Rivas, 1980: 29. Type species: Limia grossidens
Rivas, 1980. Type by original designation. Gender: feminine.
Limia ornata Regan, 1913a: 1016, pl. 101 (fig. 7). Type locality:
Haiti. Syntypes: (5) BMNH 1912.9.4.12-14 plus BMNH
1913.3.6.34 (6, in 1 jar).
Maximum length: 4 cm SL
Distribution: Caribbean Islands: Lake Miragoane, southwestern
Haiti.
Countries: Haiti
Limia grossidens Rivas, 1980: 29, fig. 1b. Type locality: North end
566
MICROPOECILIA
Micropoecilia Hubbs, 1926: 73. Type species: Poecilia vivipara
parae Eigenmann, 1894. Type by original designation. Gender:
feminine.
567
EOHETERADRIA
PAMPHORICHTHYS
Pamphoria Regan, 1913a: 1003. Type species: Cnesterodon
scalpridens Garman, 1895. Type by monotypy. Gender:
feminine.
Pamphorichthys Regan, 1913a: 1003. Type species: Heterandria
minor Garman, 1895. Type by original designation. Gender:
masculine.
Parapoecilia Hubbs, 1924: 11 [footnote]. Type species: Limia
hollandi Henn, 1916. Type by original designation. Gender:
feminine.
Limia hollandi Henn, 1916: 138. Pl. 19 (fig. 3). Type locality:
Penedo, Rio San Francisco. Holotype: FMNH 55861 [ex CM
4643a].
Maximum length: 2.1 cm SL
Distribution: South America: So Francisco River basin and one
record from the Parnaba River basin (Figueiredo, 1997).
Countries: Brazil
Common names: Barrigudinho (Brazil), Guaru (Brazil)
PHALLICHTHYS
Phallichthys Hubbs, 1924: 10 [footnote]. Type species:
Poeciliopsis isthmensis Regan, 1913. Type by original
designation. Gender: masculine.
568
PHALLOCEROS
PHALLOTORYUS
PHALLOPTYCHUS
Phalloptychus Eigenmann, 1907: 430. Type species: Girardinus
januarius Hensel, 1868. Type by original designation. Gender:
masculine.
569
POECILIA
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. Originally
spelt Mollinesia but emmendation is justified (Bailey & Miller,
1950). Taxonomy and phylogenetic hypotheses of subgenus
Mollienensia: Hubbs (1933); Schultz & Miller (1971); Miller
(1975, 1983); Rauchenberger (1989).
Alazon Gistel, 1848: X. Type species: Poecilia vivipara Bloch &
Schneider, 1801. Type by being a replacement name. Gender:
masculine.
Lebistes De Filippi, 1861: 69. Type species: Lebistes poecilioides
De Filippi, 1861. Type by monotypy. Gender: masculine.
Acanthophacelus Eigenmann, 1907: 426 [footnote]. Type species:
Poecilia reticulata Peters, 1859. Type by original designation.
Gender: masculine.
Allopoecilia Hubbs, 1924: 11 [footnote]. Type species: Girardinus
caucanus Steindachner, 1880. Type by original designation.
Gender: feminine.
eopoecilia Hubbs, 1924: 11 [footnote]. Type species:
eopoecilia holacanthus Hubbs, 1924. Type by original
designation. Gender: feminine.
Psychropoecilia Myers, 1935: 311. Proposed as subgenus of
Poecilia. 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.
Recepoecilia Whitley, 1951: 68. Type species: Poecilia vivipara
parae Eigenmann, 1894. Type by being a replacement name.
Gender: feminine.
Countries: Mexico
Poecilia dauli Meyer & Radda, 2000
Poecilia dauli Meyer & Radda, 2000: 77, fig. 1. Type locality:
brook near Miranda, Venezuela. Holotype: NWM-94540.
Maximum length: 3.9 cm TL
Distribution: South America: Coastal drainages of Venezuela
Countries: Venezuela
Poecilia elegans (Trewavas, 1948)
Mollienesia elegans Trewavas, 1948: 409, pl. 1 (fig. 4); figs. 1-2.
Type locality: Jarabocoa, San Domingo, Haiti, West Indies.
Holotype: BMNH 1947.11.27.4-5 (1 of 2).
Maximum length: 3.6 cm SL
Distribution: Caribbean Islands: Eastern Hispaniola Island.
Countries: Dominican Republic
570
Maximum length: 11 cm SL
Distribution: North and Central America: From San Juan River,
Mexico to Guatemala. At least one country reports adverse
ecological impact after introduction.
Countries: Belize, Guatemala, Mexico, Panama (?)
571
572
POECILIOPSIS
Hemixiphophorus Bleeker, 1860: 440. Type species: Xiphophorus
gracilis Heckel, 1848. Type by subsequent monotypy. Gender:
masculine. Appeared first in key, without included species. One
species added by Bleeker (1860: 485). Senior synonym of
Poeciliopsis Regan, 1913 but Poeciliopsis is placed in the official
list of generic names in zoology (opinion 375) of the ICZN (see
Rosen & Bailey, 1963:131 and Melville & Smith, 1987).
Poeciliopsis Regan, 1913a: 996. Type species: Poecilia presidionis
Jordan & Culver, 1895. Type by subsequent designation. Gender:
feminine.
Leptorhaphis Regan, 1913a: 998. Type species: Gambusia infans
Woolman, 1895. Type by monotypy. Gender: feminine.
Aulophallus Hubbs, 1926: 69. Type species: Poecilia elongata
Gnther, 1866. Type by original designation. Gender: masculine.
Poecilistes Hubbs, 1926: 68. Type species: Heterandria lutzi
Meek, 1904. Type by original designation. Gender: masculine.
Countries: Mexico
573
Countries: Mexico
PRIAPELLA
Priapella Regan, 1913a: 992. Type species: Gambusia bonita
Meek, 1904. Type by monotypy. Gender: feminine.
574
PRIAPICHTHYS
Countries: Colombia
PSEUDOPOECILIA
Pseudopoecilia Regan, 1913a: 995. Type species: Poecilia festae
Boulenger, 1898. Type by monotypy. Gender: feminine.
Countries: Panama
Priapichthys nigroventralis (Eigenmann & Henn,
1912)
Gambusia nigroventralis Eigenmann & Henn in Eigenmann, 1912:
26. Type locality: Rio San Juan at Itsmina, Colombia. Holotype:
FMNH 56045 [ex CM 4835].
Gambusia caudovittata Regan, 1913b: 471. Type locality: Rio
Condoto [Colombia]. Holotype: BMNH 1913.10.1.68.
Maximum length: 3 cm TL
Distribution: South America: San Juan River basin.
Countries: Colombia
QUITAA
Quintana Hubbs, 1934: 2. Type species: Quintana atrizona Hubbs,
1934. Type by original designation. Gender: feminine.
SCOLICHTHYS
Scolichthys Rosen, 1967: 2. Type species: Scolichthys greenwayi
Rosen, 1967. Type by original designation. Gender: masculine.
575
TOMEURUS
Tomeurus Eigenmann, 1909: 53. Type species: Tomeurus gracilis
Eigenmann, 1909. Type by monotypy. Gender: masculine.
XEOPHALLUS
Xenophallus Hubbs, 1924: 10 [footnote]. Type species: Gambusia
umbratilis Meek, 1912. Type by original designation. Gender:
masculine.
XIPHOPHORUS
576
577
Xiphophorus
nezahualcoyotl
Kallman & Morizot, 1990
Rauchenberger,
GEERA IQUIREDAE
Arizonichthys Nichols, 1940: 1. Type species: Arizonichthys
psammophilus Nichols, 1940. Type by monotypy. Gender:
masculine.
Gambusia (Toluichthys) Dahl in Dahl & Medem, 1964: 80. Type
species: Gambusia (Toluichthys) meadi Dahl, 1964. Type by
monotypy. Gender: masculine.
Hubbsichthys Schultz, 1949: 95. Type species: Hubbsichthys
laurae Schultz, 1949. Type by original designation. Gender:
masculine.
SPECIES IQUIREDAE
Arizonichthys psammophilus Nichols 1940: 1. Type locality: 3 1/2
miles east of Tanque Verde, Pima County, Arizona, altitude 2750
ft. Holotype: AMNH 15373 (not found in 1996).
Fundulus capensis Garman 1895: 113, Pl. 3 (fig. 2). Type locality:
False Bay, Cape of Good Hope, South Africa [apparently in
error]. Holotype: MCZ 6454.
Gambusia (Toluichthys) meadi Dahl in Dahl & Medem, 1964: 80,
fig. pg. 81. Type locality: vecindad de Tol. Holotype: not
located.
Gambusia baracoana Rivas, 1944b: 46. Type locality: small freshwater pond, near the mouth of Rio Miel, in the vicinity of the city
of Baracoa, Province of Oriente [Cuba]. Holotype: USNM
203150 [ex Rivas coll. 134].
Girardinus januarius var. reticulatus Hippius 1910: 392, Figs. 3132. Unknown type locality. No types known.
Girardinus microdactylus rivasi Baru & Wohlgemuth, 1994: 250,
fig. 1. Type locality: small man-made freshwater pool near
Rancho Tesoro Hotel, Nueva Gerona, Isla de la Juventud, Cuba.
Holotype: Inst. Landscape Ecol., Acad. Sci. Czech Republic ILE
11993.
Girardinus zonatus Schreiner & Miranda Ribeiro, 1903: 73. Type
locality: Brazil ? Syntypes: MNRJ (10) uncatalogued and
missing.
Hubbsichthys laurae Schultz, 1949: 96, fig. 13. Type locality:
Near Pampn, Estado de Trujillo, Venezuela, and probably in
Ro Motatn drainage. Holotype: USNM 120999.
Lebistes poecilioides De Filippi, 1861: 70, pl. 4 (figs. 6, 6a-d).
Type locality: Barbados, West Indies. No types known. Probable
synonym of some species of Poecilia, but not P. reticulata.
Lebistes reticulatus aurata Schreitmller, 1934: 242. No type
localilty. Syntypes: ZMB 21084 (7). Name found in Eschmeyer
(1998) without reference to a publication. Availability of name
undetermined.
Mollienesia sphenops petersi Schindler, 1959: 1, fig. 1. Type
locality:Yojoa-See (NWHonduras, atlantisches Einzugsgebiet).
Holotype: ZSM 15639.
Poecilia mexicana De Filippi in Tortonese, 1940: 142. type
locality: Citt di Messico. Not available: nomen nudum also
preoccupied by Poecilia mexicana Steindachner, 1863.
Poecilia unipunctata Gurin-Mneville, 1829-38: 28, pl. 47 (fig.
3). Type locality: les eaux douce de lAmrique Mridionale, a
Rio-Janeiro. No types known.
Xiphophorus elegans Paepke & Seegers, 1986: 175. Type locality:
?. Syntypes: ZMB 31510 (2)?
Xiphophorus heckelii Weyenbergh, 1874: 292, pls. 1-2 (= figs. 131). Type locality: La Plata. Syntypes: MSNG 33704 (4)
Xiphophorus marmoratus Obregon-Barboza & ContrerasBalderas, 1988: 95, fig. 2. Type locality: Cascada La Alberca, sw
of Mzquiz, road to airport, Coahuila, Mexico. Holotype: UANL
8077.
Xiphophorus minor Weyenbergh, 1877: 20, pl. 4 (fig. 19). Type
locality: Catamarca, Argentina. Syntypes: (several) Mus.
Nacional Argentina, Buenos Aires.
Xiphophorus obscurus Weyenbergh, 1877: 18, pl. 4 (figs. 17-18).
578
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