Phalloceros harpagos, Lucinda, 2008

Lucinda, Paulo Henrique Franco, 2008, Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species, Neotropical Ichthyology 6 (2), pp. 113-158 : 134-138

Phalloceros harpagos , new species

Figs. 27-29

Phalloceros sp. n. V Lucinda & Reis (2005).

Holotype. MNRJ 23610, rio São Francisco, tributary to rio Paquequer Fazenda São Francisco de Paula, approx. 21 o 59’S 42 o 54’W, Sapucaia, Rio de Janeiro, Brazil, 6Aug 1990, D.A. Halboth etal.

Paratypes. BRAZIL. Rio de Janeiro. MCP 20569, 165/10*, Teresópolis, rio Paquequer, near bridge on km 86 on road BR 116, upstream Represa Guinle, 22 º 27’18"S 42 º 57’33"W, 18 Jan 1997, J.F.P. Silva et al. MCP 30508, 6/4*; and MNRJ 14745, 59, collected with the holotype.

Non-types. Rio Paraná-Paraguay basin. Río Paraguay Drainage: PARAGUAY. Alto Paraná. ANSP 47684, 2, Puerto Bertoni. Caaguazú. NRM 42043, 1, Arroyo Guazú, where crossing Ruta 2, 25 º 28’23"S 56 º 13’34"W. Caazapá. MNHNP 1091, 6, Parque Nacional Caaguazú, Arroyo Guazú in the park border. NRM 33811, 4, 25 km SE of Asunción, Jhuguañaró. Guairá. NRM 41893, 2, stream crossing road from Mauricio J. Troche to Independencia, 5 km S of Río Tebicuary bridge, 25 º 40’56"S 56 º 18’28"W. Misiones. ANSP 173591, 40, stream crossing Ruta 1 at km A240/E125*. Ñeembucú. USNM 181784, 11, Colonial L. Vera (pond), Tebicuary drainage. Paraguarí. AMNH 99990, 50, Arroyo Mbuyapey, in Ybycuí National Park, near Ybycuí. Lower rio Paraná Drainage: PARAGUAY. Canendiyu. AMNH 47923, 5, Arroyo Guazú-Ri 10 km above Mbuty road on old route from Stroessner to Guairá. MNHNP 428, 36, Arroyo Primavera, .. [sic] km S from Arroyo Guazú-Ri by the old road from ciudad del Este to Saltos del Guairá. UMMZ 206403, 32, Río Piratiy at Balneario Municipal, 1 km S of bridge on dirt highway just W of Salto del Guairá. Itapúa. MNHNP 1082, 30 of 104, Colonia Exparadema, Compañía Tarumá, Arroyo Tarumá. MNHNP 1098, 2, Cordillera San Rafael, Colonia Alto Verá, Arroyo Tacuary. NRM 28358, 4, stream crossing Encarnación- Hohenau road ca. 7 km from Hohenau. UMMZ 206173, 30 of 170, Arroyo San Rafael 2.2 km by dirt road ca. NE of San Rafael, 26 º 38’12"S 54 º 53’W. Guairá. ANSP 175297, 2, Arroyo Perulero where it crosses road from Villarrica, 25 º 51’53"S 56 º 24’40"W. ANSP 175301, 16, stream crossing road that runs through Parque Nacional Guayaquil. Tebicuary - Paraná drainage, 25 º 41’1"S 56 º 18’31"W. Misiones. ANSP 175302, 17, stock pond by road from San Ignacio to Pilar, 26 º 52’51"S 57 º 3’5"W. ANSP 175303, 62, pool in stream crossing Asunción - Encarnación highway, 26 º 53’24"S 56 º 53’47"W. Rio Paranaíba Drainage: BRAZIL. Distrito Federal. MNRJ 19986, 9, Brasília, BR-020, N exit from Brasília, close to bridge on road to Sobradinho, near Parque Nacional de Brasília. Minas Gerais. MCP 27374, Abadia dos Dourados, rio Preto, road from Palmito to Abadia dos Dourados, 18 º 6’15"S 47 º 41’35"W. Rio Grande Drainage: BRAZIL. Minas Gerais. MCP 18134, 3, Carandaí, rio Carandaí upstream Carandaí. MCP 23498, 6, Delfim Moreira, rio Santo Antonio, affluent to rio Sapucaí. MCP 23501, 6, Cristina, rio Lambari, affluent to rio Verde. MCP 23515, 6, Poços de Caldas, ribeirão Cipó affluent to rio das Antas. MCP 23516, 6, Aiuruoca. rio Aiuruoca. MZUSP 35406, 28, Fortaleza de Minas, rio São João. São Paulo. MCP 27014, 12, Itirapina, Represa do Lobo, mouth to the rio Itaqueri. MZUSP 49231, 40 of 82, Cajuru, unnamed creek at Fazenda Santa Carlota. Rio Tietê Drainage: BRAZIL. São Paulo. MCP 20582, 35, Buritiba Mirim, first road from Biritiba Mirim to Casa Grande. MCP 20586, 22, Salesópolis, creek Paraitinguinha on road Salesópolis to Jacareí, 3 km N from Salesópolis. MCP 25526, 8, Biritiba Mirim, creek on road from Biritiba Mirim to Casa Grande, affluent to Barragem Ponte Nova at rio Tietê. MCP 27375, 1, Piracicaba, ribeirão Vermelho, road Piracicaba - Anhembi, ca. 14 km from Piracicaba, 22 º 42’59"S 47 º 43’27"W. MCP 28686, 3, Paranapiacaba, creek ca. 2 km NW from Paranapiacaba, on road from Campo Grande to Paranapiacaba, 23 º 46’21"S 46 º 18’52"W. MZUSP 35345, 20 of 70, Bofete, ribeirão Água Fria. MZUSP 45486, 11, Campinas, creek affluent to rio Atibaia, estrada da Fortuna near Rodovia D. Pedro I. MZUSP 49467, 20 of 35, Araçariguama, creek affluent to ribeirão Araçariguama, at the Mineração Morro Grande. NRM 43501, 8, road to Barbosa, 5 km from road SP-153, second creek, above bridge. Rio Paranapanema Drainage: BRAZIL. Paraná. MCP 12583, 8, Jaguariaíva, rio Capivari under bridge on road PR-92. MCP 27007, 13, Ponta Grossa, rio Verde. Rio Iguaçu Drainage: BRAZIL. Paraná. MCP 22274, 11, Paulo Frontin, arroio Barra Grande on road BR 153, 25 º 59’42"S 50 º 52’0"W. MCP 22275, 2, Cruzeiro do Iguaçu, beach at Represa Salto Caxias at Foz do Chopim, 25 º 34’5"S 53 º 7’22"W. MCP 22276, 1, Catanduvas, sanga Feijão, affluent to rio Adelaide, 25 º 15’25"S 53 º 8’30"W. MCP 22277, 4, Pérola do Oeste, creek on road BR 163, between Planalto and Pérola do Oeste, 25 º 43’44"S 53 º 47’9"W. MCP 27006, 5, Piraquara, Cavas do Piraquara. MCP 27013, 17, Curitiba, Fazenda Canguiri. MCP 27018, 89, Faxinal do Céu, creek affluent to rio Iguaçu. MHNCI 9065, 6, Araucária, Guajuvira, Chácara Kotowski, PR 423. USNM 297981, 9, headwaters of rio Iguaçu, near Serrinha (road BR 476 between Lapa and Contenda). USNM 304995, 8, ribeirão Amola-Faca, tributary to rio Iguaçu, near Balsa N. in Fazenda Lara Maria. Santa Catarina. USNM 320323, 3, rio Negrinho, under bridge on road BR 280, rio Negrinho. Coastal drainages (from north ro south). Rio Itaboapana Drainage. BRAZIL. Espírito Santo. MCP 17839, 2, rio São José das Torres near road BR-101, between São José das Torres and Travessão, 21 º 4’43"S 41 º 14’2"W. Rio Paraíba do Sul Drainage: BRAZIL. Rio de Janeiro. MNRJ 14424, 34, Teresópolis, ribeirão Santa Rita (tributary to the left margin of rio Paquequer), near Fazenda Santo Afonso. MNRJ 15312, 7, Carmo, córrego Astréia near Astréia. MNRJ 15539, 7, Sumidouro, rio Paquequer, upstream Sumidouro. USNM 257030, 72, rio dos Coitinhos at km 55 on road RJ 155, headwater tributary to rio Piraí, near road RJ 155 between Angra dos Reis and Rio Claro, 22 º 51’20"S 44 º 14’30"W. São Paulo. MCP 18132, 6, Piquete, creek affluent to rio Piquete at Benfica. 22 º 35’46"S 45 º 13’6"W. MCP 20570, 3, Arapeí, small tributary to rio Bananal, ca. 6 km NE from Arapeí, on road from Arapeí to Santana do Bom Sucesso ( SP 66), 22 º 40’32"S 44 º 24’36"W. MCP 20575, 3, São José do Barreiro, rio Água Santa, 22 º 38’40"S 44 º 34’31"W. MCP 20584, 7, Taubaté, córrego Itaí on road Taubaté - São Luís Paraitinga, ca. 5 km S from Taubaté 23 º 4’17"S 45 º 33’20"W. MCP 20587, 17, São Luís Paraitinga, ribeirão do Pinga tributary to ribeirão do Chapéu at Catuçaba, 23 º 14’37"S 45 º 12’32"W. MCP 20588, 27, Silveiras, córrego Macaquinho, tributary to rio Paraitinga, ca. 5 km NW from Bairro dos Macacos, 22 º 50’47"S 44 º 50’30"W. MCP 27015, 9, Paraibuna, rio Paraíba do Sul. MZUSP 47189, 27, Cunha, Serra dos Macacos, riacho do Macaquinho, near Sítio do Sr. Jair Romão. MZUSP 47193, 21 of 30, Mogi das Cruzes, Cachoeirinha do Angico, Sítio do Sr. Manuel Figueiredo, Bairro de Itapeti. MZUSP 47202, 27, Santa Isabel, headwaters of rio Piläes, Sítio Bem Viver. MZUSP 47582, 9, Pindamonhangaba, ribeirão Grande, talhão 1, near Sede of Fazenda São Sebastião do Ribeirão Grande. USNM 309782, 56, stream about halfway between Santa Branca and Salesópolis. Rio Macaé Drainage: BRAZIL. Rio de Janeiro. MCP 18133, 2, rio Macaé on road from Casemiro de Abreu to Lumiar, ca. 5 km E from Lumiar. MCP 20566, 7, Macaé, rio dos Frades at Frades, 22 º 14’34"S 42 º 6’18"W. Rio Macacu Drainage: BRAZIL. Rio de Janeiro. USNM 309773, 3, rio Macacu near Cachoeiras de Macacu, small tributary ca. 1-2 km from town, highway bridge over stream, collected near and under bridge. Lagoa de Saquarema Drainage: BRAZIL. Rio de Janeiro. USNM 257028, 33, rio Seco, tributary to lagoa de Saquarema along road Amaral Peixoto between Sampaio Correia and Bacaxa on map. locality is at bridge. USNM 257029, 15, rio do Mato Grosso, tributary to rio Roncador which flows into lago de Saquarema. Rio São João Drainage: BRAZIL. Rio de Janeiro. MCP 20565, 9, Silva Jardim, rio Iguapé near road BR 101, ca. 14 Km SE from Casemiro de Abreu, 22 o 30’58"S 42 o 19’20"W. Baía de Guanabara System: BRAZIL. Rio de Janeiro. MZUSP 43471, 20 of 32, Rio de Janeiro, Floresta da Tijuca. USNM 309784, 21, km 27 on road Amaral - Peixoto, about 1 km from Itapeba in small tributary to rio Fundo, near Maricá. MNRJ 15133, 1, Magé, road to Santo Aleixo, rio Santo Aleixo tributary to rio Roncador near Santo Aleixo, 22 o 35’ 23"S 43 o 2’45"W. Rio Mato Grosso-Roncador System. UNT 6768, 10, Saquarema, rio Mato Grosso, 22°53’12"S 42°39’54"W. Rio Tavares Drainage: BRAZIL. São Paulo. USNM 257037, 51, southern branch of rio Tavares, Ubatuba system at nursery of “flora Brazilia ”. Rio Escuro Drainage: BRAZIL. São Paulo. USNM 257027, 17, rio Escuro. Rio Juqueí Drainage: BRAZIL. São Paulo. MZUSP 49936, 11, São Sebastião, Praia de Juqueí, stream affluent to rio Juqueí. Ilha Bela: MCP 31131, 317, Ilha Bela, stream in the road to Praia dos Castelhanos, 1.5 km from Ilha Bela, 23 º 49’30"S 45 º 21’48"W. Rio Cabuçu Drainage: BRAZIL. São Paulo. NRM 43502, 4, Baixada Santista, unpaved from road Santos to Rio de Janeiro, tributary creek of the rio Cabuçu. Other small coastal drainages in São Paulo State: MCP 30999, 58, Barra do Turvo, rio Bonito, affluent to rio Pardo, Bairro Rio Bonito, 24 º 50’13"S 48 º 29’27"W. MCP 31128, 148, São Sebastião, córrego da Barra, 23 º 47’22"S 45 º 33’7"W. MCP 31144, 64, Peruíbe, stream Cachoeira da Lagoa in Guaraú, 24 º 22’5"S 47 º 3’17"W. MCP 31186, 11, Mongaguá, lateral pond ca. 3 km from road SP 55, 24 º 7’1"S 46 º 43’58"W. MCP 31187, 10, Itanhaém, affluent stream to rio Preto ou Branco ca. 2 km from state airport 24 º 8’37"S 46 º 46’34"W. MCP 31622, 155, Caraguatatuba, creek on Praia de Massaguaçú, ca. 700 m W of road SP 55, 23 º 34’23"S 45 º 19’38"W. MCP 31623, 185, Santos, creek at Carnaíra ca. 100 m from road SP 55, at border Bertioga - Santos, 23 º 52’56"S 46 º 11’29"W .

Diagnosis. Phalloceros harpagos can be distinguished from P. anisophallos , P. megapolos , P. spiloura , P. reisi , P. buckupi , P. alessandrae , P. lucenorum , P. uai , P. pellos , and P. malabarbai by the female urogenital papilla straight along midline and located between the anus and the base of first anal-fin ray (vs. curved to the right, located laterally; border of the anal aperture in contact with the first anal-fin ray). Phalloceros harpagos can be distinguished from P. tupinamba , P. leptokeras , and P. aspilos by (1) the female urogenital papilla straight along midline (vs. slightly left turned); (2) absence of a lateral ramus of the female urogenital papilla (vs. lateral ramus present); and (3) small and simple hook in gonopodial appendix (vs. large sickle like hook). Phalloceros harpagos can be distinguished from P. caudimaculatus , P. heptaktinos , P. ocellatus , P. mikrommatos , and P. leticiae by the possession of a hook in the gonopodial appendix (vs. hook absent). Phalloceros harpagos can be distinguished from P. titthos by the absence of symphyseal papillae on large females (vs. symphyseal papillae present); and from P. enneaktinos by the possession of seven or eight dorsal-fin rays (vs. nine) (see remarks). Phalloceros harpagos can be distinguished from P. elachistos by the hook of gonopodial appendix not forming a lateral protuberance (vs. very small in adults, forming a lateral minute protuberance – see remarks). In juveniles hook present and located on the appendix corner (vs. absent or (rarely) minuscule, when present in juveniles hook never situated on the appendix corner). Skin present covering the tip of appendix on juveniles (vs. skin absent). Distal half of appendix narrower than proximal half (vs. approximately as wide as or slightly narrower than proximal half).

Description. Morphometric data in Tables 9 and 10. Range of SL: 18.4 to 46.8 mm (females), 14.8 to 34.1 mm (males). Dorsal-fin rays: 7 [15], 8* [639], 9 [21]. Branched pectoral-fin rays: 5 [3], 6 [245], 7* [273], 8 [5]. Pelvic-fin rays: 5* [203] (males), 4 [2], 5 [450] (females).Anal-fin rays of females:10 [85], 11 [349], 12 [1, incomplete]. Anal-fin rays of males: 8 [4], 9* [221], 10 [21]. Branched caudal-fin rays: 8 [1], 10 [13], 11 [118], 12* [375], 13 [89], 14 [17], 15 [1], 16 [1]. Predorsal scales: 12 [3], 13 [49], 14* [368], 15 [32]. Longitudinal series of scales: 27 [22], 28 [196], 29* [444], 30 [35]. Series of scales around caudal peduncle: 14 [5], 15 [2], 16* [688].Transverse series of scales: 7* [562]. Serrae on R4p: 8 [8], 9 [27], 10 [76], 11 [42], 12 [36], 13 [31], 14* [11], 15 [5], 16 [1]. Epipleural ribs: 11 [2], 13 [2]. Pleural ribs: 14 [2], 15 [2]. Vertebrae: 31 [1], 32 [2], 33 [1]. Female urogenital papilla straight, located along midventral line and between anus and base of first anal-fin ray. Gonopodial appendix hook present ( Fig. 28). Hook present in juveniles and located on appendix corner. Juveniles bearing skin covering tip of appendix. Distal half of appendix narrower than proximal half ( Fig. 28).

Color in alcohol. Eye black with greenish brown pupil. Ground Color brown, darker in upper half. Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, conferring reticulate pattern to body sides, mainly on upper half. Brown chromatophores scattered through whole body, more concentrated on dorsal portion, mainly on head, snout, opercle, and ventral surface of mandible. Lateral dark brown spot located between 16 th and 19 th scale of longitudinal series. Dorsal-fin membrane hyaline bearing dark band of chromatophores on distal border of dorsal fin and another near dorsal-fin base. Pectoral, pelvic, and caudal fins hyaline. Dark brown line along R3. First three anal-fin rays of females covered of brown chromatophores forming patch of dark pigmentation.

Intraspecific Variability. Variation in some morphological features (e. g. color pattern, hook on gonopodial appendix, measurements) has been found among distinct populations of P. harpagos during this revisionary study. Although differences may be found between some population pairs, it has not been possible to assign any of them as diagnostically distinct from the remaining populations. So, these populations are referred to as the same species.

Degree of development of spots varies enormously, ranging from large and wide dark patch through small and narrow spot to no apparent mark. Variation on color cannot be associated as exclusive of any local population, for all variants are generally present in any of the populations. Alternatively the frequencies of each of the color “ types ” can be correlated to populations.

Specimens from coastal drainages of Rio de Janeiro and São Paulo usually possess a roundish and short lateral spot, whereas southern populations in Santa Catarina usually show an elongate and wide lateral spot reaching dorsal and ventral profiles. Generally, specimens from the rio Paraná drainage possess a narrow and vertically elongate spot not reaching dorsal and ventral profiles. Individuals from some populations, e. g. rio Ribeira do Iguape, rio Iguaçu, present a very narrow and short lateral spot.

Particularly, the population in the rio Ribeira de Iguape is an interesting case. This river is formed by the junction of rio Ribeira and rio Juquiá. Specimens from the rio Ribeira have a very narrow and short lateral spot, while specimens from the rio Juquiá possess a large and wide lateral spot. They are quite different from each other. Despite this, none of them can be distinguished from the bulk of P. harpagos , i. e. each can not be diagnosed or recognized as a distinctive species.

Similarly, some specimens of the lower portions of rio Paraná differ from specimens from rio Paraguay. The former show narrow and vertically elongate lateral spot, whereas the later show large and roundish lateral spot. The two forms occur in the lower rio Paraná. This difference is congruent with differences in the depth of body and caudal peduncle of males. Again, both are clearly different from each other but cannot be distinguished from populations along the range of P. harpagos as a whole.

Degree of development of the hook in the gonopodial appendix may also vary from a large hook to lateral expansion, especially in specimens from coastal drainages in southern Santa Catarina, which usually present a shorter hook on gonopodial appendix.

At the moment, it is advisable to treat these differences as intraspecific variation, unless new evidences become available and allow the recognition of new diagnosable taxonomic entities.

Etymology. From the Greek ’αρπαγος [ = harpagos ], n. m. nom. sg., meaning hook, alluding the presence of a hook in the gonopodial appendix. An noun in apposition.

Distribution. Rio Paraná-Paraguay basin and coastal drainages from rio Itaboapana (ES) to rio Araranguá (SC) ( Fig. 29).