A New Species of Poecilia from Honduras (Teleostei: Poeciliidae) Fred N. Poeser1
Copeia 2011, No. 3, 418–422
A New Species of Poecilia from Honduras (Teleostei: Poeciliidae) Fred N. Poeser1 A new species of Poecilia from the Atlantic slopes of Honduras is described and assigned to the subgenus Mollienesia. Poecilia hondurensis, new species, differs from all congeners in the combination of having tricuspid teeth on the inner of both jaws and lacking a prominent spine or hook on the gonopodium. Eleven geographically close congeners are treated in some detail, and a key is provided to identify all species occurring south of Mexico.
T
HE Neotropical genus Poecilia is a diverse assembly of species subdivided in at least five subgenera (Rosen and Bailey, 1963; Poeser et al., 2005). All species of Poecilia from Central America are allocated to the subgenus Mollienesia sensu Miller (1975), except for P. caucana, a peripheral species from Panama that belongs to the subgenus Allopoecilia Hubbs, 1924. Within Mollienesia, the P. sphenops complex is a group of species that shows remarkable morphological similarity considering its wide distribution, from northern Mexico (P. mexicana and P. butleri) to Venezuela (P. koperi), which caused Rosen and Bailey (1963) to lump as many as 34 nominal taxa as synonyms of P. sphenops. In Poecilia, many morphometric features are more variable within species than between them. Thus, the validity of describing new species based on proportional body measurements is questionable. Rivas (1978) noted, ‘‘there is considerable variation in characters individually, ontogenetically, seasonally, geographically, and environmentally and, therefore, they are of little or no value in distinguishing species (except relative fin position).’’ Only recently has the identification of characters, and therefore species of the genus Poecilia, been supported by molecular data (Brett and Turner, 1983; Meyer et al., 2004). Characters found to be taxonomically useful are the shape of the inner jaw dentition and the number of anal-fin rays (Schultz and Miller, 1971), the number of dorsal-fin rays, gonopodial substructures, and the presence of a dark lateral spot (Miller, 1975), possession of head pores 1 and 2a and the number of scales around the caudal peduncle (Greenfield, 1990), and the presence and position of a blotch at the caudal-fin base (Poeser, 1995). The specific and subgeneric allocations needed to identify the species for comparison to the new species are indicated in a comprehensive study (Ptacek and Breden, 1998; Breden et al., 1999) on the molecular relationships within Poecilia. Using the characters assessed here, a key is constructed that supplements the key given by Miller (1983) for the Mexican species of Poecilia. MATERIALS AND METHODS Scales counts around the caudal peduncle and ray counts of the unpaired fins follow Hubbs and Lagler (1947), wherein the first two unbranched rays were counted as one. Head pores were reported following Gosline (1949). Gonopodial rays were counted following Rosen and Gordon (1953). Special reference was made to the shape of the segments in the gonopodium, which were characterized as cubic when the sides of segments were equal in length, as normal when the length of a segment was about 1
twice as long as its width, and elongated when the segment was at least three times longer than wide. Intermediate shapes were more fully described. Segments with outgrowths were recorded as spiny when they resulted in a serrated margin, or as bumpy when frayed rather than pointed. Morphometric data were omitted in the diagnosis and descriptions because they vary more within species than between species (Rivas, 1978), and have been susceptible to environmental influences (ecomorphs, Poeser, 1992, 2003), character displacement (Rivas, 1982), or a combination of both (Poeser, 1995, 1998). Poecilia (Mollienesia) hondurensis, new species Figures 1, 2; Table 1 Holotype.—UMMZ 213664, male, 42.9 mm SL, Honduras, Olancho, confluence of Rı´o San Jose´ and Rı´o Conquire, under new cement bridge of the new San Esteba´n-Trujillo highway, 9 July 1983, G. A. Cruz and O. Melendez. Allotype.—UMMZ 248253, female, 62.8 mm SL, same data as holotype. Paratypes.—All from Honduras: Atlantida: UMMZ 213665, 19, 34.8–57.2 mm SL, same data as holotype; UMMZ 193853, 5, 44.0–62.9 mm SL, stream on Route 50, 34 km WSW of La Ceiba, 7 May 1973, A. Klee, R. Norris, R. Socolof, and H. Specht; UMMZ 203346, 1, 82.1 mm SL, Laguna Ticamaya, 1 June 1970, E. Comfort; UMMZ 213660, 2, 33– 44 mm SL, Rı´o Santuye, below bridge of Ceiba-Jutiapa highway, 8 March 1983, G. A. Cruz and M. Espinal; UMMZ 213662, 8, 43.8–63.7 mm SL, De Yoro, Rı´o Uchapa, from Santa Clara pond, near the Municipio of Olanchito, 1 April 1983, J. Valerio, G. Valerio, and M. Santos; UMMZ 213663, 31, 16–59 mm SL, Olancho, 8 July 1983, G. A. Cruz and O. Melendez; UMMZ 213666, 18, 22–45 mm SL, Olancho, Quebrada Casa Quemada, 3.5 km NW of Carbon, along the new highway to Trujillo, 9 July 1983, G. A. Cruz and O. Melendez; UMMZ 213673, 15, 29.5–42.3 mm SL, De Yoro, stream below ‘‘el puente Los Arellanos,’’ near Bellavista, 19 March 1984, G. A. Cruz; UMMZ 213686, 10, 36.8–43.2 mm SL, Olancho, Rı´o Mataderos, highway to San Esteba´n, 450 m above sea level, 8 July 1983, G. A. Cruz and O. Melendez; UMMZ 219144, 6, 30–37 mm SL, small stream on highway CA 13, 5.8 miles W of Trujillo, 28 April 1991, R. Socolof, H. Specht, and R. Wessel; UMMZ 241742, 1, 44.5 mm SL, Rı´o Aguan on highway CA 13, 44.6 mi W of Trujillo, 28 April 1991, R. Socolof, H. Specht, and R. Wessel.
Department of Ichthyology, Zoo ¨ logisch Museum Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands; E-mail:
[email protected]. Submitted: 15 June 2009. Accepted: 19 June 2011. Associate Editor: C. J. Ferraris. DOI: 10.1643/CI-09-111 F 2011 by the American Society of Ichthyologists and Herpetologists
Poeser—A new species of Poecilia
Fig. 1. Poecilia hondurensis, new species, holotype, male, UMMZ 213664, confluence of Rı´o San Jose´ and Rı´o Conquire, bridge on San Esteba´n-Trujillo highway, Olancho, Honduras.
Diagnosis.—Poecilia hondurensis is distinguished from all other species of Poecilia that have an inner row of tricuspid teeth in each jaw by the absence of a spinal hook on gonopodial ray 5 and an extremely reduced membranous hook on gonopodial ray 3. Description.—Holotype, adult male (Fig. 1), with distinct blotch at caudal-fin base. Tricuspid inner jaw dentition. Supra-orbital head pores 1 and 2a visible. Dorsal fin with nine or ten rays, anal fin with nine rays. Usually 16 scales around caudal peduncle (one female specimen, 82.1 mm SL, UMMZ 203346, with 18 scales). Lateral scales 27 or 28. Shape and outline of gonopodial rays (Fig. 3) normal for Mollienesia (Miller, 1975), except G4p is much thicker than G4a. Gonopodial ray 3 with hood nearly reaching tip. Terminal segments of ray 3 cubic, last two seemingly fused. Next five segments with weak ventral extensions, followed by segments with ventral and dorsal extrusions. Dorsal extrusions extend to ray 4a. No obvious membranous hook at tip. Segments of ray 4a elongate, four times longer than high. Segments of ray 4p much thicker than those of ray 4a, but equally long. Terminal eight segments of ray 4p more or less cubic, followed by eight segments with dorsal serrae. Rays 5a and 5p with unmodified terminal segments, without terminal hook or extruding spine. Allotype, adult female (Fig. 2), with two or three rows of spots in dorsal fin, posterior upper margin of dorsal fin has spots. Dark blotch at caudal-fin base not reaching upper and lower margins of caudal fin, body with few spots. Nine anal and dorsal-fin rays. Comparisons.—Poecilia hondurensis is most similar to P. marcellinoi, which also possesses tricuspid inner teeth and a blotch on the caudal base. However, the blotch in P. hondurensis is located on the caudal fin (vs. on the scales of the caudal peduncle in P. marcellinoi). Poecilia hondurensis further differs from P. marcellinoi in having no extruding spines at the gonopodial tip (vs. well developed in P. marcellinoi). Poecilia hondurensis differs from P. latipinna, P. velifera, and P. kykesis in its short dorsal fin (9–10 vs. 12–18 dorsal-fin rays). Poecilia hondurensis has a tricuspid inner jaw dentition, distinguishing it from the unicuspid teeth of P. mexicana, P. thermalis, P. gillii, P. petenensis, P. butleri, P. latipunctata, P. nelsoni, P. sulphuraria, P. orri, P. teresae, and P. rositae from Central America, and from P. mechthildae, P. koperi, P. boesemani, and P. vandepolli from South America. Poecilia hondurensis differs from P. sphenops by having fewer scales around the caudal peduncle (16 vs. 18). Poecilia hondurensis differs from P. chica by having more anal-fin rays (consistently 9 vs. 8). Poecilia hondurensis is different from
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Fig. 2. Poecilia hondurensis, new species, allotype, female, UMMZ 248253, confluence of Rı´o San Jose´ and Rı´o Conquire, bridge on San Esteba´n-Trujillo highway, Olancho, Honduras.
P. catemaconis, P. maylandi, and P. marcellinoi in lacking extruding spines (vs. present) on the gonopodium. Poecilia hondurensis further differs from P. formosa for being a bisexual species (vs. an all-female species). Poecilia hondurensis is most similar to P. marcellinoi, with which it shares the caudal blotch. However, the caudal spot is a fused area of caudal-fin spots (versus caudal blotch located on the caudal-peduncle scales in P. marcellinoi). These two species also differ in gonopodial characters. Poecilia hondurensis shares gonopodial characters with P. orri, e.g., missing external spines or hooks, but it differs in body pigmentation (blotch present at caudal-fin base vs. absent) and inner jaw dentition (tricuspid vs. unicuspid). Color in alcohol.—Ground coloration of body brown, some specimens with vertical bands. Other paratypes with scattered specks on the body and caudal peduncle. Specimens with up to eight pigment specks on the body show onset to a melanistic pattern. Dorsal fin with row of spots, no blotch. Color in life.—Males light yellow on dorsal and caudal fins. Dorsal fin with thin black edge. Males and females with black line along both sides of dorsal-fin base, also on dorsal side of body. Distribution.—Poecilia hondurensis is known only from the Caribbean drainage basins of Honduras. Remarks.—Files at the University of Michigan (UMMZ) show that the species described herein was under investigation by R. R. Miller since 1953. Due to the lack of sufficient material, Miller did not feel comfortable publishing at the time. His greatest concern was that well developed males were lacking. The species was independently discovered by Gustavo Cruz, a scientist from Honduras who provided extra material to Miller and added valuable data on localities for the present description. Miller and Cruz were working on a joint publication when Miller passed away. While studying the fish at the UMMZ, I discovered the documents relating to the new species and made arrangements to describe the species with the approval of Cruz. A sample of 11 of the 26 species of Mollienesia mentioned in the present paper was investigated in two molecular studies (Ptacek and Breden, 1998; Breden et al., 1999). These 11 species formed a monophyletic clade, separating them from P. vivipara, P. caucana, and P. reticulata. These results supported the recognition of the subgenus Mollienesia sensu Miller (1975). The species of Mollienesia have been subdivided into three groups based on the size of the dorsal fin (Miller, 1983): the sailfin mollies; the shortfin mollies; and a hybrid between these groups, P. formosa. Only four shortfin
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Copeia 2011, No. 3
Table 1. List of Species of the Subgenus Mollienesia Used in the Comparison with P. hondurensis, New Species, Using Rosen and Bailey (1963) as a Benchmark.
Species Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia
boesemani butleri catemaconis formosa gillii koperi kykesis latipinna latipunctata marcellinoi maylandi
Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia Poecilia
mechthildae mexicana nelsoni orri petenensis rositae salvatoris sphenops sulphuraria teresae thermalis vandepolli velifera
Remarks Described in Poeser (2003); Shortfin molly group Revalidated by Schultz and Miller (1971); Shortfin molly group Described in Miller (1975); Shortfin molly group Hybrid between shortfin and sailfin molly groups (Miller, 1983) Revalidated by Bussing (1987); Shortfin molly group Described in Poeser (2003); Shortfin molly group Replacement name for Mollienesia petenensis (cf. Poeser, 2002); Sailfin molly group Sailfin molly group Shortfin molly group Described in Poeser (1995); Shortfin molly group Synonym of P. sphenops in Rosen and Bailey (1963) as P. sphenops pallida (5unidentifiable species [Miller et al., 2005:236]); Renamed and revalidated by Meyer (1983) Described in Meyer et al. (2002); Shortfin molly group Revalidated by Schultz and Miller (1971); Shortfin molly group Revalidated by Poeser (2003); Shortfin molly group Revalidated by Miller (1983); Shortfin molly group Revalidated by Poeser (2002); Shortfin molly group Described in Meyer et al. (2004) Revalidated by Miller (1994); Shortfin molly group Shortfin molly group Shortfin molly group Described in Greenfield (1990) Revalidated by Poeser (2003); Shortfin molly group Revalidated by Feltkamp and Kristensen (1969); Shortfin molly group Sailfin molly group
mollies, including the wide-ranging P. sphenops, were recognized as valid species in Rosen and Bailey (1963), whereas ten species have been resurrected more recently from synonymy of their P. sphenops. In addition, eight species of shortfin mollies have been described as new since 1963 (Table 1). The new species is a shortfin molly and is also a member of the P. sphenops complex within Mollienesia. Etymology.—This species was named ‘‘hondurensis’’ in the manuscript notes of Miller and Cruz after carefully delimitation of its range. I have retained this name in honor of both discoverers.
KEY TO THE CENTRAL AMERICAN SPECIES OF POECILIA, SOUTH OF MEXICO 1a. 1b. 2a.
2b. 3a.
3b. 4a. 4b. 5a. 5b. 6a.
Inner jaw dentition unicuspid 2 Inner jaw dentition tricuspid 10 12 or more dorsal-fin rays P. kykesis Tributary of Rı´o Usumacinta and nearby lakes, Pete´n, Guatemala to southeastern part of Mexico, and basal part of Yucatan Peninsula, Quintana Roo and Belize Fewer than 12 dorsal-fin rays 3 18 scales around caudal peduncle P. mexicana Atlantic coast of Central America, from Texas border, through Yucatan into Guatemala 16 scales around caudal peduncle 4 8 anal-fin rays 5 9 anal-fin rays 6 Head pores 1, 2a present. C. 18–22; LLS 28 P. rositae Guatemala and Belize Head pores 1, 2a absent. C. 16–18; LLS 26–28 P. nelsoni From Cabo Corrientes, Jalisco, Mexico to Guatemala Slender species, torpedo body shape. 8 dorsal-fin rays P. teresae Upstream habitats in Belize Deep bodied species, sides compressed. 9 or 10 dorsal-fin rays 7 Gonopodium without terminal hook or spine P. orri Eastern part of Yucatan Peninsula, Quintana Roo, Northern Honduras, and offshore islands Gonopodium with terminal hook or spine 8 Dorsal-fin rays 10–11 (10 modally). C. 18; LLS 28–29 P. petenensis Endemic to Lake Pete´n __________________________________________
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Fig. 3. Gonopodium of Poecilia hondurensis, new species. Note the absence of an extruding spine on ray 5p, the very reduced hook at ray 3, and that ray 4p is much thicker than ray 4a.
7b. 8a.
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Poeser—A new species of Poecilia
8b. 9a. 9b.
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10b. 11a.
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Dorsal-fin rays 8–9 (9 modally). C. 18–22; LLS 26–27 9 P. salvatoris Body with vertical stripes, never yellow belly El Salvador, Guatemala, and Honduras Body without vertical stripes, yellow belly P. gillii Occurs from Atlantic coastal areas of Colombia, northward along both oceanic coasts of Costa Rica and Panama, to Pacific side of Guatemala, and Atlantic side of Honduras 18 scales around caudal peduncle P. sphenops From the Atlantic slopes of Mexico, Rı´o de la Palma Sola, to the Pacific side of Guatemala 16 scales around caudal peduncle 11 Gonopodium with distinct terminal hook and spine. Blotch at caudal-fin base on scales of caudal P. marcellinoi peduncle. LLS 26–27 Pacific side of Guatemala, Honduras, Nicaragua, and El Salvador Gonopodium with reduced terminal hook on ray 3, no spine on ray 5. Blotch at caudal-fin base on membrane of caudal-fin. LLS 27–28 P. hondurensis, new species Atlantic slopes of Honduras ___
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MATERIAL EXAMINED Poecilia gillii: Costa Rica, Turrialba: FMNH 6027, holotype of Platypoecilus tropicus, female, 46.1 mm SL; FMNH 6028, holotype of Poecilia tenuis, Tirribi; FMNH 6029, 2 paratypes of P. tenuis, same data as holotype; FMNH 6360, holotype of P. caudata, female, 68.5 mm SL, Turrubares; FMNH 96297, allotype of P. caudata, male, 64.6 mm SL, same data as holotype; FMNH 6258, 7 paratypes of P. caudata, largest female 60.8 mm SL, Patarra, San Jose´; FMNH 59535, 2 paratypes of Platypoecilus tropicus. Poecilia kykesis: ZMB 6082, syntype of Mollienesia petenensis, Guatemala, Lake Pete´n; AMNH 37659, 5, Guatemala, Pete´n, Flores, Lake Pete´n, Arroyo San Helena, opposite Flores South, 28 March 1954, M. S. Gordon and Pinelo; AMNH 37660, 2, Guatemala, Pete´n, Flores, Lake Pete´n, Arroyo de Cuxa. Poecilia mexicana: NMW 61228, 7 syntypes, 29.7–40.4 mm SL, Mexico, Orizaba; UMMZ 95516, holotype of Mollienesia sphenops macrura, female, 96 mm SL, Guatemala, Pete´n, Rı´o San Pedro de Martir, El Pasa do Caballo; UMMZ 97874, holotype of Mollienesia sphenops vantynei, male, 72 mm SL, Guatemala, Pete´n, Aguada at Uaxactun; UMMZ 102127, holotype of Mollienesia sphenops altissima, male, 53 mm SL, Mexico, Yucatan, Miramar Spring, 1 km SW of Talcha, N of Motul on coast. Poecilia orri: Honduras, Bay Islands Dept.: ANSP 70158, holotype, 28.6 mm SL, Bonacca Island, harbor front and wharves; AMNH 29835, paratype of P. orri, same data as holotype; ANSP 71752, 5 paratypes of Poecilia vetiprovidentiae, up to 62.8 mm SL, Old Providence island, J mi S of Ironwood Hill in Huffington’s Creek at Watering Place and above, E side of island. ACKNOWLEDGMENTS I am thankful for the friendly reception and kind assistance that I have received from the following persons, while visiting the collections under their care: P. Bartsch (ZMB, Berlin), B. Brown (AMNH, New York), G. Duhamel (MNHN, Paris), W. Fink (UMMZ, Ann Arbor), D. Nelson (UMMZ, Ann Arbor), P. Pruvost (MNHN, Paris), M. Rogers (FMNH, Chicago), M. Stiassny (AMNH, New York), K. Swagel (FMNH, Chicago), and C. Latour (ZMB, Berlin). A very special thank
you to G. Cruz (MHN, University of Honduras), who kindly provided firsthand information on the species that he had planned to describe with R. Miller. I also appreciate greatly the support from J. Me´rido (MHN, University of Honduras), who provided many freshly caught specimens from nearly all Honduran species of Poecilia for comparison. I am also grateful to W. Matamoros (Ichthyology Lab., University of Southern Mississippi), who provided valuable information on the species of Poecilia and their habitats from Honduras, and for his kind donation of material now catalogued at ZMA. I also thank the management of the Amsterdam International Community School for their kind support offering me the time to complete this study. LITERATURE CITED Breden, F., M. B. Ptacek, M. Rashed, D. Taphorn, and C. A. Figueiredo. 1999. Molecular phylogeny of the livebearing fish genus Poecilia (Cyprinodontiformes: Poeciliidae). Molecular Phylogeny and Evolution 12:95–104. Brett, B. L. H., and B. J. Turner. 1983. Genetic divergence in the Poecilia sphenops complex in Middle America. Biochemical Systematics and Ecology 11:127–137. Bussing, W. A. 1987. Peces de las aquas continentales de Costa Rica. Editorial Universidad de Costa Rica, San Jose´, Costa Rica. Feltkamp, C. A., and I. Kristensen. 1969. Ecological and morphological characters of different populations of Poecilia sphenops vandepolli (Cyprinodontidae). Studies of the Fauna of Curac¸ao 31:102–130. Gosline, W. A. 1949. The sensory canals of the head in some cyprinodont fishes, with particular reference to the genus Fundulus. Occasional Papers, Museum of Zoology, University of Michigan 519:1–17. Greenfield, D. W. 1990. Poecilia teresae, a new species of poeciliid fish from Belize, Central America. Copeia 1990:449–454. Hubbs, C. L. 1924. Studies of the fishes of the order Cyprinodontes. Miscellaneous Publications of the Museum of Zoology, University of Michigan 13:1–31. Hubbs, C. L., and K. F. Lagler. 1947. Fishes of the Great Lakes region. Cranbrook Institute of Science 26:1–186. Meyer, M. K. 1983. Une nouvelle espe`ce de Poecilia du Guerrero, Mexique (Pisces: Poeciliidae). Revue Franc¸aise Aquariologique 10:55–58. Meyer, M. K., V. Etzel, and D. Bork. 2002. A new species of Poecilia, subgenus Mollienesia, from Venezuela, with comments on Poecilia cuneata Garman, 1895 and other related taxa (Teleostei: Cyprinodontiformes: Poeciliidae). Zoologische Abhandlungen (Dresden) 52:49–55. Meyer, M. K., K. Schneider, A. C. Radda, B. Wilde, and M. Schartl. 2004. A new species of Poecilia, subgenus Mollienesia, from upper rı´o Cahabo´n system, Guatemala, with remarks on the nomenclature of Mollienesia petenensis Gu ¨ nther, 1866 (Teleostei: Cyprinodontiformes: Poeciliidae). Zoologische Abhandlungen (Dresden) 54:145–154. Miller, R. R. 1975. Five new species of Mexican poeciliid fishes of the genera Poecilia, Gambusia, and Poeciliopsis. Occasional Papers of the Museum of Zoology, University of Michigan 672:1–44. Miller, R. R. 1983. Checklist and key to the mollies of Mexico (Pisces: Poeciliidae: Poecilia, subgenus Mollienesia). Copeia 1983:817–822. Miller, R. R. 1994. Origin and classification of the Liberty Molly. Tropical Fish Hobbyist 6:104, 106, 108.
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Miller, R. R., W. L. Minckley, and S. M. Norris. 2005. Freshwater Fishes of Mexico. University of Chicago Press, Chicago. Poeser, F. N. 1992. Re-establishment and redescription of Poecilia vandepolli Van Lidth de Jeude, 1887 (Pisces: Poeciliinae), with comments on related species. Studies of the Natural History of the Caribbean Region 71:79–98. Poeser, F. N. 1995. Nonrandom variation in Poecilia marcellinoi n. sp. and P. salvatoris Regan, 1907 in El Salvador (Pisces, Poeciliidae). Bijdrage tot de Dierkunde 64:239–252. Poeser, F. N. 1998. The role of character displacement in the speciation of Central American members of the genus Poecilia (Poeciliidae). Italian Journal of Zoology 65(Supplement):145–147. Poeser, F. N. 2002. Poecilia kykesis nom. nov., a new name for Mollienesia petenensis Gu ¨ nther, 1866, with resurrection, redescription and designation of a lectotype for Poecilia petenensis Gu ¨ nther, 1866 (Pisces: Poeciliidae). Contributions to Zoology 70:243–246. Poeser, F. N. 2003. From the Amazon river to the Amazon molly and back again. Unpubl. Ph.D. diss., University of Amsterdam, Amsterdam. Poeser, F. N., M. Kempkes, and I. J. H. Isbru ¨ cker. 2005. Description of Poecilia (Acanthophacelus) wingei n. sp. from the Parı´a Peninsula, Venezuela, including notes on
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Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology 74:97–113. Ptacek, M. B., and F. Breden. 1998. Phylogenetic relationships among mollies (Poeciliidae: Poecilia: Mollienesia group) based on mitochondrial DNA sequences. Journal of Fish Biology 53(Suppl. A):64–81. Rivas, L. R. 1978. A new species of poeciliid fish of the genus Poecilia from Hispaniola, with reinstatement and redescription of P. dominicensis (Evermann and Clark). Northeast Gulf Science 2:98–112. Rivas, L. R. 1982. Character displacement and coexistence in two poeciliid fishes of the genus Poecilia (Mollienesia) from Hispaniola. Northeastern Gulf Science 5: 1–24. Rosen, D. E., and R. M. Bailey. 1963. The poeciliid fishes (Cyprinodontiformes), their structure, zoogeography, and systematics. Bulletin of the American Museum of Natural History 126:1–176. Rosen, D. E., and M. Gordon. 1953. Functional anatomy and evolution of male genitalia in poeciliid fishes. Zoologica 38:1–47. Schultz, R. J., and R. R. Miller. 1971. Species of the Poecilia sphenops group (Pisces: Poeciliidae) in Mexico. Copeia 1971:282–290.
