A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope Part 5

=Syrrhophus pipilans nebulosus= Taylor

_Syrrhophus nebulosus_ Taylor, 1943:353-55, pl. 27, figs. 3-5 [Holotype.--FMNH 100095 (formerly EHT-HMS 3774), near Tonola, Chiapas, Mexico; collected on August 27, 1935, by Hobart M.

Smith and Edward H. Taylor]. Smith and Taylor, 1948:49, 51.

_Syrrhophus pipilans nebulosus_: Duellman, 1958:2-4, 9, 12, 14.

Stuart, 1963:32-33. Gorham, 1966:166-67.

_Diagnosis._--Diameter of tympanum 36.6-47.8 per cent that of eye; dorsum dark brown with numerous small light brown to yellowish spots.

_Remarks._--The distribution of this subspecies is adequately described by Duellman (1958). Fouquette (1960) described the vocalization of this frog.

_Etymology._--Latin, _nebula_, in reference to the clouded dorsal pattern.

_Distribution._--Low to moderate elevations along the Pacific versant of Chiapas and in the Grijalva valley of Chiapas and Guatemala (Fig. 17).

_Specimens examined._--(54) GUATEMALA, _Huehuetenango_: Jacaltenango, UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MeXICO, _Chiapas_: 11.2 km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279; 12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; 1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca.

2 km. NW Pueblo Nuevo Solistahuacan, KU 58897; Hda. Monserrate, 40 km.

NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH 75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, TNHC 26942, Tonola, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); 19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, KU 37809; Union de Juarez, FMNH 105294.

=Syrrhophus pipilans pipilans= Taylor

_?Syrrhopus verruculatus_: Gadow, 1905:194.

_Syrrhophus pipilans_ Taylor, 1940c:95-97, pl. 1 [Holotype.--FMNH 100072 (formerly EHT-HMS 6843), from 14.6 km. S Mazatlan, Guerrero, Mexico; collected on July 22, 1936, by Edward H. Taylor].

Taylor and Smith, 1945:581-82. Smith and Taylor, 1948:49, 50-51.

_Syrrhophus pipilans pipilans_: Duellman, 1958:1-4, 8-9, 13-14, pl. 2, fig. 1. Gorham, 1966:166.

_Diagnosis._--Diameter of tympanum 40.6-54.0 per cent that of eye; dorsum dark brown with large light spots or blotches.

_Remarks._--Duellman's (1958) synopsis of this subspecies is adequate; the distribution has not been extended, but several records are now available which fill in gaps.

[Ill.u.s.tration: FIG. 17: Distribution of _Syrrhophus pipilans_: _nebulosus_ (open circles) and _pipilans_ (solid circles).]

Gadow's (1905) record of _S. verruculatus_ from "Buena Vista, S.

Guerrero" is most likely applicable to this species. Gadow simply included the name in a list of the species he had collected during his trip in Mexico (1902-04); no further comment was made on this species although references to _Syrrhopus_ (sic) appear in several places in the paper and would appear to apply to the species he had.

_Etymology._--Latin, _pipilo_, chirping, peeping, in reference to the call of the male.

_Distribution._--Sea level to about 1800 meters along the Pacific versant of western Mexico from central Guerrero to the Isthmus of Tehuantepec (Fig. 17).

_Specimens examined._--(62). MeXICO, _Guerrero_: Acapulco, UMMZ 110125; 6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, 125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S Mazatlan, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH 126639; Xaltinanguis, FMNH 100522-24, 126640. _Oaxaca_: Cacahuatepec, UIMNH 52853; 8 km. NW Rio Canoa, 53 km. ESE Cuajinicuilapa, UIMNH 52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; 37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 (2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ 115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2).

=Syrrhophus interorbitalis= Langebartel and Shannon

_Syrrhophus interorbitalis_ Langebartel and Shannon, 1956: 161-65, figs. 1-2 [Holotype.--UIMNH 67061 (formerly FAS 9378), 36 mi. N Mazatlan, Sinaloa, Mexico, collected on November 17, 1955, by E. C. Bay, J. C. Schaffner, and D. A. Langebartel]. Duellman, 1958:1-4, 10, 12, 14. Gorham, 1966:164-65.

_Syrrhophis interorbitalis_: Campbell and Simmons, 1962:194, fig. 1.

[Ill.u.s.tration: FIG. 18: Left to right. _Syrrhophus interorbitalis_ (UIMNH 38095, 1.5), _S. nivocolimae_ (LACM 3203, 1.3), and _S. teretistes_ (KU 75263, 1.5).]

_Diagnosis._--Medium sized frogs, only known male 25.6 mm. snout-vent, females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in males; finger tips expanded; first finger shorter than second; outer metatarsal tubercle one-third size of inner; skin of dorsum s.h.a.greened, that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of eye in both s.e.xes; pale yellow-brown ground color mottled with brown; limb bands broad, much wider than narrow light inters.p.a.ces; interorbital bar very long, edged with dark brown to black (Fig. 18).

_Remarks._--Duellman's (1958) measurements and proportions of _S. interorbitalis_ were based exclusively on the type series, which is composed of only females; therefore his _interorbitalis_ data are not comparable with the data for the other species in his table. Campbell and Simmons (1962) collected the only known male. The type series was collected beneath rocks in a stream bed; the collectors heard calling frogs in the bushes but were unable to obtain specimens (Langebartel and Shannon, 1956). Campbell and Simmons (1962) reported that their specimen had a poorly developed interorbital bar in life; in preservative the bar compares favorably with the bar in the female (Fig. 18).

_Etymology._--Latin, in reference to the pale interocular band.

_Distribution._--Pacific lowlands of Sinaloa, Mexico (Fig. 20).

_Specimens examined._--(10). MeXICO, _Sinaloa_: 36 mi. N Mazatlan, UIMNH 38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlan, LACM 13773.

=Syrrhophus modestus= Taylor

_Syrrhophus modestus_ Taylor, 1942:304-06, pl. 29 [Holotype.--FMNH 100048 (formerly EHT-HMS 3756), from Hacienda Paso del Rio, Colima, Mexico; collected on July 8, 1935, by Hobart M. Smith].

Smith and Taylor, 1948:49-50.

_Syrrhophus modestus modestus_: Duellman, 1958:2-5, 7, 14, pl. 1, fig. 1. Gorham, 1966:166.

_Diagnosis._--Small frogs, males 15.8-20.1 mm. snout-vent length, single female 18.5 mm.; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum s.h.a.greened, that of venter smooth; tympanum concealed; pale cream in preservative with dark brown spots; limbs banded; bands on forearm and thigh poorly developed or absent; interorbital bar absent.

_Remarks._--The tympanum is concealed in _S. modestus_, _S. nivocolimae_, _S. pallidus_, _S. teretistes_, and to a lesser degree in _S. interorbitalis_. However, if the specimen is permitted to dry slightly, the annulus tympanicus becomes visible through the skin and a tympanum/eye ratio can be computed.

One of the few cases of sympatry within the genus _Syrrhophus_ involves this species; _modestus_ and _nivocolimae_ are known to be sympatric at one locality in southwestern Jalisco, Mexico.

Duellman (1958) used the trinomial for this population and named a new subspecies, _pallidus_, from Nayarit. I consider _pallidus_ to be specifically distinct from _modestus_ because there is no evidence of genetic exchange, and there is no overlap in the distinguishing morphological features. I do consider the two populations to be closely related but feel the interrelationships between _modestus_, _pallidus_, _nivocolimae_, and _teretistes_ are more complex than would be indicated by the use of trinomials. The sympatric occurrence of _modestus_ and _nivocolimae_ is significant; morphologically, they might otherwise be regarded as subspecies. Although allopatric, similar arguments could be advanced for the morphologically similar _pallidus_ and _teretistes_.

The four are here afforded species rank since morphological similarity and allopatry are not sufficient grounds for the a.s.sumption of genetic exchange.

[Ill.u.s.tration: FIG. 19: _Syrrhophus modestus_ [left, UMMZ 115447 (WED 11155)] and _S. pallidus_ (right, UMMZ 115453). 2.2.]

_Etymology._--Latin, meaning una.s.suming, modest, in reference to the small size of the species.

_Distribution._--Low elevations (up to 700 meters) in the lowlands and foothills of Colima and southwestern Jalisco, Mexico (Fig. 20).

_Specimens examined._--(14). MeXICO, _Colima_: Hda. Paso del Rio, FMNH 100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM 139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4); _Jalisco_: 17.6 km. SW Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahia Tenacat.i.ta, UMMZ 84264.

=Syrrhophus nivocolimae= Dixon and Webb

_Syrrhophus nivocolimae_ Dixon and Webb, 1966:1-4, Fig. 1 [Holotype.--LACM 3200, from Nevado de Colima (6 airline miles west of Atenquique), Jalisco, Mexico, 7800 feet; collected on July 20, 1964, by Robert G. Webb].

_Diagnosis._--Small frogs, males 18.5-21.1 mm. snout-vent length, only known female 24.1 mm. snout-vent; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum warty, that of venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that of eye in males; mid-dorsal brown band from interorbital bar to a.n.u.s; bands on limbs narrow, dark bands less than one-half width of light bands, upper arm not banded; narrow interorbital light bar.

_Remarks._--This species is closely related to _S. modestus_ and differs in color pattern and degree of wartiness of the skin. Dixon and Webb (1966) held that _nivocolimae_ had no close relatives, but the condition of the tympanum, size, nature of the outer palmar tubercle, relative sizes of the metatarsal tubercles, and shape and size of the digital pads all point to a close relationship between _S. modestus_, _S. nivocolimae_, and _S. pallidus_.

[Ill.u.s.tration: FIG. 20: Distribution of the species of the _modestus_ group: _interorbitalis_ (open circles), _teretistes_ (solid circles), _modestus_ (open triangles), _pallidus_ (solid triangles) and _nivocolimae_ (square). Arrow indicates locality of sympatry between _modestus_ and _nivocolimae_. Solid line about the localities for _interorbitalis_ is a range estimate based on call records and specimens examined.]