Phalloceros caudimaculatus ( Hensel, 1868 )

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 : 121-124

Phalloceros caudimaculatus ( Hensel, 1868)

Figs. 10 , 11

Girardinus caudimaculatus Hensel, 1868: 362 . Type-locality: an der Costa da Serra bei S. Leopoldo in Brunnen und Gräben [= at the Costa da Serra at São Leopoldo in pounds and ditches].

Type-material. ZMB 7425, lectotype [by present designation] , Costa de Sierra [sic], Hensel. ZMB 7426, 5, paralectotypes [not examined], Costa da Serra at S. Leopoldo, (Hensel?). ZMB 31496 (13 of 25 females), paralectotypes, “C. da Serra,...? [illegible] bei S. Leopolds” [sic]. ZMB 33220, 1, paralectotype, collected with the lectotype .

Material Examined. Laguna dos Patos System. BRAZIL. Santa Catarina. MCP 22337, 6, Bom Jardim da Serra, rio Barrinha at Bom Jardim da Serra, way-out to Santa Bárbara, 28 º 20’15"S 49 º 37’49"W . Rio Grande do Sul. MCP 9305, 11, Pantano Grande, arroio Dom Marcos, 30 º 13’32"S 52 º 37’9"W. MCP 9539, 3, Cachoeira do Sul. MCP 10060, 1, Três Coroas, arroio Moreira, Distrito de Sander. MCP 10285, 14, Canoas, Distrito de Mato Grande, stream affluent to rio dos Sinos. MCP 10750, 5, Arroio dos Ratos, near BR 290. MCP 11356, 6, arroio Xasqueiro, near road BR 116 between Pelotas and Jaguarão. MCP 11557, 2, Gramado, arroio Linha Bonita. MCP 17249, 65, São Sepé. MCP 17385, 6, Minas do Leão, arroio Taquara under bridge on road BR 290, near Minas do Leão. MCP 17578, 29, Pelotas, arroio Totó, road to Colônia Z-3. MCP 18249, 18, Faxinal do Soturno, creek near road Faxinal do Soturno to Ivorá. MCP 19104, 2, Barra do Ribeiro. MCP 19698, 2, Encruzilhada do Sul, arroio Passo da Miséria, between Encruzilhada do Sul and Canguçu. MCP 20158, 16/4*, Sapiranga, arroio Feitoria. MCP 20837, 16, Pedro Osório, arroio Reduzinho, between Pedro Osório and Basílio, 31 º 54’25"S 52 º 55’41"W. MCP 21251, 29, Segredo, wetland at Três Vendas on road from Bela Vista to Coloninha, 29 º 15’32"S 53 º 53’34"W. MCP 21355, 68, Tavares, stream at Balneário de Bujuru, 31 º 40’18"S 51 º 24’27"W. MCP 21355, 68, Viamão, stream at Praia da Pedreira, Parque Estadual de Itapuã, 30 º 21’30"S 51 º 2’48"W. MCP 21515, 1, arroio Formoso and lateral ponds, ca. 7.8 km N from Dona Francisca, 29 º 33’49"S 53 º 20’59"W. MCP 22338, 22, Lageado Grande, arroio Lageado Grande, near road to Jaquirana, 29 º 5’34"S 50 º 37’30"W. MCP 23666, 25, Santa Maria, wetland ca. 500 m from rio Vacacaí. MCP 23883, 14, São Lourenço do Sul, stream affluent to arroio Grande between Colônia de Santana and road BR 116, 31 º 22’38"S 52 º 7’37"W. MCP 23884, 7, Tapes, arroio Teixeira and lateral pond on road BR 116, 30 º 37’26"S 51 º 32’45"W. MCP 25755, 49, Camaquã, arroio Maria Ulghim, at road from Camaquã to Vila Aurora. MCP 25784, 1, Caçapava do Sul, arroio do Banhado on road BR 153 ca. 8 km NW of Minas do Camaquã, 30 º 51’48"S 53 º 29’50"W. MCP 25874, 24, creek affluent to arroio Correio, between Caçapava do Sul and Lavras do Sul, 30 º 43’15"S 53 º 40’53"W. MCP 25899, 2, Lavras do Sul, stream affluent to arroio Camaquã Chico, 30 º 59’6"S 53 º 58’20"W. MCP 26051, 2, Dois Irmãos, arroio Feitoria at Picada Verão, 29 º 34’37"S 51 º 1’39"W. MCP 26126, 24, Santa Maria Herval, arroio Cadeia, affluent to rio Caí, at Prainha do Jacaré. MCP 27298, 2, Candiota, arroio Candiota, Passo do Cerro, near confluence with rio Jaguarão, 31 º 49’57"S 53 º 51’19"W. MCP 29254, 61, São Francisco de Paula, arroio Lava-pés, affluent to rio Caí ca. 20 km from road RS 235. MCP 30889, 9, Canela, arroio Caçador, tributary to rio Caí, 29 º 18’50"S 50 º 47’15"W. UFRGS 2728, 98, Rio Grande, km 115 of road BR 471. Estaçäo Ecológica do Taim. URUGUAY. Rocha. ZVC-P 3209, 9, Ruta 14, km 498, bañado de los Índios, Laguna Negra - Laguna Merín drainage. ZVC-P 3229, 2, Ruta 14, km 457, Río San Luis - Laguna Merín. ZVC-P 4006, 3, Ruta 15, km 179, Laguna Merín. Rio Ibicuí drainage : BRAZIL. Rio Grande do Sul. MCP 9490, 6, Cacequi, stream near road from Cacequi to São Francisco de Assis, Vila Itapevi. MCP 14136, 10, Dom Pedrito. MCP 23081, 2, São Francisco de Assis, rio Inhacundá at São Francisco de Assis exit to Manuel Viana, 29 º 32’51"S 55 º 8’11"W. MCP 26746, 9, Santana do Livramento, arroio Ibicuí da Faxina near BR 158, 30 º 47’31"S 55 º 12’35"W. Rio Negro drainage : BRAZIL. Rio Grande do Sul. MCP 22880, 27, rio Paraizinho on road from Bagé to BR 293, ca. 1 km S of BR 293 . URUGUAY. Cerro Largo. MCP 10394, 10,Arreria, Río Negro, hacienda Arreria. Colonia. ZVC-P 676, 5, Carmelo, Río Uruguay. Durazno. ZVC-P 3324, 3, lateral laguna, Arroyo Cordobés, affluent to Río Yí, affluent to Río Negro. ZVC-P 3400, 1, 3 km NW from Estación de AFE, paraje 329, nearby mouth of Río Tacuarembó on Río Negro. Rio Mampituba drainage: BRAZIL. Santa Catarina. MCP 29267, 90, Praia Grande, arroio Moia Coco at Vila Rosa ca. 5 km NW from Praia Grande, 29 º 10’27"S 49 º 58’26"W . Rio Grande do Sul. MCP 14840, 34, Três Cachoeiras, rio das Pacas, near Morro Azul. MCP 23731, 4, Mampituba,rio Panela. Rio Tramandaí drainage : BRAZIL. Rio Grande do Sul. MCP 10801, 5, Osório, rio Maquiné. MCP 20291, 27, Torres, creek on dunes. MCP 25294, 30, Terra de Areia, rio Três Pinheiros, under bridge on road to Vila Itati, ca. 7 km N from road BR 101. MCP 25385, 7, Maquiné, arroio Água Parada. MCP 29283, 5, Itati, stream on road between “Rota do Sol” ( RS 486) and Vila Itati, 29 o 29’22"S 50 o 6’20"W. UFRGS 4456, 20, Tramandaí. USNM 297952, 1, rio Sanga Funda, crossing BR 101 road just S of Terra de Areia. Laguna Castillos drainage : URUGUAY. Rocha. ZVC-P 3237, 8, bridge at Arroyo Valizas, Laguna Castillos. ZVC-P 3248, 21, pond alongside secondary road between Balneario Aguas Dulces and Ruta 10, Laguna Castillos. Argentinean coastal drainage: Buenos Aires. AMNH 93354, 54, Punta Lara, just N of La Plata, in roadside ditches.

Diagnosis. Phalloceros caudimaculatus 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 caudimaculatus 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) absence of hook in gonopodial appendix (vs. hook present). Phalloceros caudimaculatus can be distinguished from P. enneaktinos , P. titthos , P. elachistos , and P. harpagos by the absence of hooks in both halves of the gonopodial appendix (vs. hook present in both halves). Phalloceros . caudimaculatus is readily distinguished from P. heptaktinos by the absence of sexual dimorphism on number of dorsal-fin rays: males with 8 rays (very exceptionally seven) and females with seven rays (very exceptionally 8). Phalloceros caudimaculatus can be distinguished from P. ocellatus , P. mikrommatos , and P. leticiae by the lateral spot: P. caudimaculatus possesses a non-ocellated elliptical and vertically elongate lateral spot (rarely absent), whereas P. ocellatus , P. mikrommatos , and P. leticiae possess roundish to rounded and well-defined ocellated lateral spot.

Description. Morphometric data in Tables 5 and 6 . Range of SL: 18.9 to 45.0 mm (females), 14.3 to 28.8 mm (males). Dorsal-fin rays: 7 [3], 8 [266*], 9 [6]. Branched pectoral-fin rays: 5 [2], 6 [76], 7 [121], 8 [5] (types total number: 10 [6], 11 [5]). Pelvic-fin rays: 4 [1], 5 [83] (males), 4 [1], 5 [185] (females). Anal-fin rays of females: 10 [23], 11* [156].Anal-fin rays of males: 8 [3], 9 [91], 10 [7]. Branched caudal-fin rays: 10 [6], 11* [68], 12 [135], 13 [30], 14 [11]. Predorsal scales:12 [3], 13 [23],14 *[159], 15* [6]. Longitudinal series of scales: 27 [28], 28 [141], 29* [104], 30 [12], 31 [2]. Series of scales around caudal peduncle: 16* [296]. Transverse series of scales: 6 [1], 7* [297]. Serrae on R4p:8 [1],9 [9],10 [30], 11 [31],12 [23], 13*[16],15 [2]. Epipleural ribs: 10 [1], 12 [2], 13 [1]. Pleural ribs: 14 [4]. Vertebrae: 33 [4]. Female urogenital papilla straight along midline, located along midventral line and between anus and base of first anal-fin ray. Hooks of gonopodial appendix absent ( Fig. 11 ).

Color in alcohol. Eye black with greenish brown pupil. Ground color pale brown, darker in upper half. Border of scales and subjacent skin replete with brown chromatophores, more concentrated at short distance from scale border, forming v-band 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 on 14 th, 15 th, and 16 th or 16 th, 17 th, and 18 th scale of longitudinal series. Lateral spot covering approximately one and half scale on horizontal direction, and four or five scales on vertical, sometimes reaching dorsal and ventral profiles. Dorsal-fin membrane bearing dark band of chromatophores on distal border of dorsal fin. Dark band of chromatophores near dorsal-fin base, more intense in posterior rays. Pectoral, pelvic, and caudal fins hyaline. Dark brown line along R3. Anal fin of females hyaline, except for patch of dark chromatophores on first three rays.

Geographical variation. Color pattern of P. caudimaculatus varies considerably. The lateral spot is very frequently present, however, it may be absent. When present the lateral spot cover one to six transverse series of scales and less than one to three scales of the longitudinal series. Similarly lateral dark bars varies from inconspicuous to visible. It was not possible to correlate the color pattern to any geographical population for it varies among the specimens of the same population.

Distribution. Laguna dos Patos system, lower portions of rio Uruguai, rio Tramandaí drainage, rio Mampituba drainage and coastal drainages of Uruguay and Argentina ( Fig. 12 ).

Remarks. Among the original type series ( Hensel, 1868: 362- 364) only the seven specimens from lots ZMB 7425 and ZMB 7426 are surely syntypes of Girardinus caudimaculatus . It is not absolutely sure, whether the 25 specimens of lot ZMB 31496 belongs to the type series, since this number is secondarily based on an old label from the anatomical collection: “No, 25202, “ Girardinus caudimaculata, C. da Serra “, - the remainder of the label is illegible ( Paepke & Seegers, 1986). This is the collection of the former “Institute of Anatomy of the Humboldt-University” in the 19th century, which used to belong to the Medical Faculty. This is due to the fact that human-anatomists were traditionally doing much research work on comparative anatomy of vertebrates. Thus, collected material was often divided between Zoological (Philosophical) Faculty with the Zoological Collection and the Medical Faculty. Only from approximately 1890 onwards the collections were reunited. But, apparently also R. Virchow, the famous pathologist working at the Charity Hospital at Humboldt-University, took over some of the original anatomical collection for study (Peter Bartsch in litt., 2000). As there is no apparent evidence that Hensel has not examined specimens from lot ZMB 31496, it should be more advisable to label them as syntypes. Whether oncoming evidences demonstrate that these specimens have not been syntypes, they will lose their type status.

Reinhold F. Hensel was a German geographer and naturalist who lived in Brazil from 1863 to 1866. He collected in Rio Grande do Sul, especially in Porto Alegre and the German colonies to the north of that city ( Papavero, 1973). Further, when Hensel returned to Germany, he published the results of his works in Brazil. In 1867 he published a large geographical report of the Província of São Pedro do Rio Grande do Sul (presently the Rio Grande do Sul State) including a map of the region ( Hensel, 1867). Later, he dealt with descriptions of new vertebrate taxa, including Girardinus caudimaculatus ( Hensel, 1868; 1870). The type-locality of G. caudimaculatus is mentioned as “an der Costa da Serra bei S. Leopoldo in Brunnen und Gräben” [= at the Costa da Serra at São Leopoldo in pounds and ditches]. The 1867 map shows that Costa da Serra corresponds to the area located in the south margin of the rio Cadeia and arroio Feitoria nearby the Encosta da Serra, and is delimited to the south by the rio dos Sinos drainage. This region currently corresponds to portions of the counties of São Sebastião do Caí, Estância Velha, and Ivoti. Although it is not possible to precisely locate the type-locality it seems very probable that Hensel would have collected his syntypes in the drainage of the rio Cadeia.

One of 26 specimens of USNM 309769, is not P. caudimaculatus (is P. harpagos ). The overall coloration of this individual is different from the remaining and P. harpagos is not sympatric with P. caudimaculatus . This specimen likely came from another lot by mistake.

Figures

Fig. 10. Phalloceros caudimaculatus, MCP 20158, arroio Feitoria, Sapiranga, Rio Grande do Sul, Brazil.(A) Male, 24.5 mm SL; (B) female,36.8 mmSL.

Fig. 11. Gonopodium tip of Phalloceros caudimaculatus, MCP 20158. R3, R4a, R4p, R5a, R5p indicate gonopodial rays. Scale bar 1 mm.

Fig. 12. Collection localities of Phalloceros ocellatus (circle, 1), P. tupinamba (triangle, 2), P. lucenorum (five-pointed star, 3), P. buckupi (eight-pointed star, 4), P. titthos (inverted triangle, 5), and P. caudimaculatus (square, 6). Some symbols represent more than one lot or locality. Numbers represent type localities. The inset is the range of the genus.

Tables

Table 5. Descriptive morphometrics of female specimens of Phalloceros caudimaculatus, P. anisophallos, P. alessandrae, and P. aspilos. Measurements 1-10 are percents of standard length and measurements 11-13 are percents of head length.

	Phalloceros caudimaculatus		Phalloceros anisophallos		Phalloceros alessandrae		Phalloceros aspilos
Character	n = 158		n = 28		n = 2		n = 13
	Range	Mean	Range	Mean	Range	Mean	Range	Mean
Standard length (mm)	18.9-45.0	28.2	21.9-45.1	34.0	22.1-24.7	23.4	25.8-38.3	34.2
1-Head length	16.8-24.8	21.5	18.1-26.2	21.6	22.1-25.2	23.6	17.6-20.8	19.0
2-Snout-occipital distance	18.9-26.2	22.4	16.2-22.6	18.6	19.6-19.9	19.8	18.0-22.1	20.1
3-Predorsal distance	58.0-66.3	62.1	60.7-65.2	63.2	63.6-64.3	63.9	58.4-63.2	60.8
4-Dorsal-fin base length	7.8-13.9	10.3	8.8-11.9	10.2	9.6-10.8	10.2	9.3-12.8	10.6
5-Anal-fin base length	6.7-14.4	9.6	6.7-10.7	9.3	7.7-10.2	8.9	8.4-11.0	10.0
6-Body depth	21.3-28.9	25.6	21.9-28.2	25.8	21.4-24.8	23.1	23.0-26.1	24.9
7-Pre-pelvic length	40.3-50.0	43.5	42.6-47.9	45.4	44.9-45.8	45.4	40.1-44.5	42.3
8-Preanal length	54.6-64.7	58.0	56.7-60.1	58.8	57.2-58.3	57.8	55.0-59.7	56.9
9-Postanal length	31.0-39.9	36.0	33.2-38.9	35.7	35.5-35.8	35.6	35.0-39.3	37.0
10-Caudal peduncle depth	13.7-17.6	15.4	14.2-16.9	15.8	14.1-14.4	14.3	13.0-14.8	14.0
11-Snout length	14.6-27.7	21.3	20.1-31.5	25.3	25.1-28.3	26.7	17.6-25.9	21.3
12-Orbital diameter	32.7-47.0	39.0	31.5-43.6	37.0	38.0-40.2	39.1	34.0-45.2	39.5
13-Postorbital length	38.7-56.3	47.2	33.4-47.2	40.1	35.3-40.9	38.1	42.7-52.2	46.0

Table 6. Descriptive morphometrics of male specimens of Phalloceros Phalloceros caudimaculatus, P. anisophallos, P. alessandrae, and P. aspilos. Measurements 1-10 are percents of standard length and measurements 11-13 are percents of head length. H = holotype.

	Phalloceros caudimaculatus		Phalloceros anisophallos			Phalloceros alessandrae	Phalloceros aspilos
Character	n = 64			n = 10		n =1		n = 3
	Range	Mean	H	Range	Mean	H	H	Range	Mean
Standard length (mm)	14.3-28.8	19.5	28.2	16.2-28.2	22.7	17.5	23.6	22.2-23.8	23.2
1-Head length	16.6-24.4	21.5	23.3	20.6-25.6	22.3	23.2	20.5	19.3-20.5	19.8
2-Snout-occipital distance	16.4-25.4	22.3	17.9	15.7 -21.4	18.8	20.4	21.8	20.1-21.8	21.2
3-Predorsal distance	55.2-62.1	58.7	60.2	57.9 -60.3	58.9	60.9	55.7	54.8-56.3	55.6
4-Dorsal-fin base length	7.5-14.5	10.8	11.1	8.1 -12.3	10.6	9.1	11.4	11.4-13.1	12.1
5-Anal-fin base length	5.8-10.8	8.0	7.9	6.5 -8.7	7.7	8.0	6.9	6.6-8.0	7.2
6-Body depth	30.9-37.4	33.9	36.7	30.6 -36.7	33.5	33.0	31.8	30.8-32.8	31.8
7-Pre-pelvic length	27.3-34.9	30.4	30.0	29.9 -32.5	30.7	30.9	27.6	27.3-27.7	27.5
8-Preanal length	37.2-45.2	40.8	39.1	38.3 -42.8	40.0	40.4	39.3	38.0-39.7	39.0
9-Postanal length	52.8-60.9	56.6	58.6	54.6 -59.3	57.5	55.9	59.0	58.5-59.5	59.0
10-Caudal peduncle depth	13.9-18.5	16.1	17.8	14.6 -18.8	16.4	15.0	15.5	14.8-15.5	15.2
11-Snout length	16.3-30.2	22.7	28.3	25.1 -31.3	27.9	29.0	22.8	21.1-25.1	23.0
12-Orbital diameter	36.2-50.5	41.9	32.7	32.7 -48.1	39.2	50.1	42.4	42.4-46.3	44.0
13-Postorbital length	33.6-50.0	43.0	42.8	29.5 -42.8	37.3	23.8	41.6	37.3-42.8	40.6