A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope Part 4

_Syrrhophus macrotympanum_: Dixon, 1957:384. Gorham, 1966:165.

_Diagnosis._--Medium-sized frogs, males 17.5-26.1 mm. snout-vent, females 28.0-31.7 mm. snout-vent length; vocal slits in males; digital tips slightly expanded; first finger shorter than second; skin of dorsum pustular, that of venter areolate; snout elongate, subac.u.minate; diameter of tympanum 56.1-76.7 per cent that of eye in males, 54.3-56.8 in females; in preservative, dorsum reddish brown with numerous small black or dark brown spots (Fig. 8); venter white to cream; in life dorsum green with darker green spots, belly white; iris gold above, bronze below.

_Remarks._--Cope's (1885) original description was not sufficiently clear to enable subsequent authors to recognize this species. Taylor (1940e) described it as a _Tomodactylus_, but Dixon (1957) pointed out that _T. macrotympanum_ differed from the other species of the genus in having a poorly developed lumbo-inguinal (inguinal) gland, and placed the species in the genus _Syrrhophus_. Comparison of the holotypes of _S. verrucipes_ and _T. macrotympanum_ leaves no doubt in my mind that a single species is involved. This same species was reported by Smith and Taylor (1948) as _S. verruculatus_.

_Syrrhophus verrucipes_ bears resemblance to members of both the _leprus_ and _marnockii_ groups. In snout shape it is closer to the _leprus_ group, whereas in digital pad, the shape of the general body form, and contiguity of habitat it is most similar to the _marnockii_ group (_S. guttilatus_).

_Etymology._--Latin, meaning warty foot, probably in reference to the numerous plantar supernumerary tubercles.

_Distribution._--Moderate elevations in southeastern San Luis Potosi, Queretaro, and northwestern Hidalgo, Mexico (Fig. 7).

_Specimens examined_--(43) MeXICO, _Hidalgo_: Jacala, UMMZ 106434; 9.6 km. NE Jacala, Puerto de la Zorra, 1820 m., KU 60240-41, TCWC 11090, 11147; 8 km. S Jacala, La Placita, 1850 m., FMNH 100049 (holotype of _Tomodactylus macrotympanum_), 100791-803, 105334-35, 114287, UIMNH 15989-92, 15995-96, UMMZ 117252, USNM 137202; Tianguistengo, FMNH 113705-09, UIMNH 13328-30; near Zacualtipan, ANSP 11325 (holotype of _Syrrhophus verrucipes_). _Queretaro_: 3.5 km. S San Juan del Rio, EAL 1343. _San Luis Potosi_: 9.6 km. W Ahuacatlan, LSUMZ 4968-70.

=Syrrhophus dennisi= new species

_Syrrhophus latodactylus_: Martin, 1958:49 (in part).

_Holotype._--UMMZ 101121, adult male from a cave near El Pachon, 8 km. N Antiguo Morelos, Tamaulipas, Mexico, 250 m., collected on March 13, 1949, by Paul S. Martin.

_Paratopotypes._--(26). UMMZ 101122 (10), 101123 (2), 101126, 126993 (12).

_Diagnosis._--Medium-sized frogs, males 22.8-28.4 mm. snout-vent, females 25.9-32.0 mm. snout-vent; vocal slits in males; digital tips greatly expanded, more than twice width of digit; first finger shorter than second; skin of dorsum s.h.a.greened to pustular, that of venter weakly to moderately areolate; toes webbed basally; dorsum light brown to tan with brown vermiculations; venter white; diameter of tympanum 53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in females.

_Description and variation._--(Fig. 12). Head wider than body; head as wide or wider than long in males, sometimes longer than wide in females; snout ac.u.minate in dorsal view, elongate and rounded in lateral profile; canthus rostralis rounded but distinct; loreal region slightly concave, sloping abruptly to lip; lips not flared; eyelid about two-thirds interorbital distance; length of eye less than distance between eye and nostril; diameter of tympanum 53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in females; tympanum round and distinct in both s.e.xes; supratympanic fold moderately distinct; choanae within border of jaws, completely visible from directly below, rounded to slightly oval; dentigerous processes of prevomers and teeth absent; tongue free for posterior one-half, generally oval in outline; vocal slits present in males.

Many scattered pustules on dorsum; flanks areolate; skin of venter areolate or not (variability may be due to differences in preservation); ventral disc distinct on chest and lower abdomen; inguinal gland present or not, when present varying from very large and distinct to poorly defined; axillary gland absent.

First finger shorter than second; all fingers bearing truncate tips with pads, each pad having a terminal groove; fingers fringed; fingers three and four having dilated pads two to three times width of digit; subarticular tubercles large, conical, rounded, simple; supernumerary tubercles numerous on thenar surface, none on digits; three palmar tubercles, outer slightly smaller than largest supernumerary tubercles; row of tubercles on outer edge of forearm variable, weak to very distinct; tips of toes wider than digits, rounded to truncate at tips, each pad having terminal groove; toes having lateral fringes, bases of toes united by web, web not extending to basal subarticular tubercle; subarticular tubercles smaller than those of hand, round, conical, simple; supernumerary tubercles numerous on plantar surfaces, extending between metatarsal tubercles, present on toes between basal two subarticular tubercles in some specimens; outer metatarsal tubercle round, conical, one-half as large as ovoid, non-compressed inner metatarsal tubercle; tarsal tubercles or folds absent.

Ground color pale reddish-brown to tan dorsally, creamy on flanks; dorsal pattern consisting of reddish-brown to brown vermiculations extending onto flanks; distinct interorbital light bar present; loreal region darker than snout, reddish-brown compared to tan or pale reddish-brown; arms colored like dorsum; thighs banded, unicolor brown on posterior surfaces; shanks and tarsi banded; venter white to cream punctated with brown in some specimens.

The variation in proportions is summarized in Table 5.

_Remarks._--Martin (1958) expressed some doubt that this series of 26 specimens was identical with "_S. latodactylus_." My study indicates that the specimens from El Pachon represent a distinctive but allied species. Males of the two species can be readily separated by the relative sizes of the tympani, presence or absence of vocal slits, and color pattern. Females of the two species can be separated by color pattern. Within the type-series, the pattern varies from weakly to strongly vermiculate but is always recognizable as vermiculate rather than spotted as in _S. longipes_ (= _S. latodactylus_ of Taylor and Martin).

[Ill.u.s.tration: FIG. 12: _Syrrhophus dennisi_ sp. nov., holotype, UMMZ 101121 (dorsum 1.8, side of head 6.1).]

_Etymology._--The specific name is a patronym for David M. Dennis, whose drawings greatly enhance the worth of this paper.

_Distribution._--Known only from the type series.

=Syrrhophus longipes= (Baird), New combination

_Batrachyla longipes_ Baird, 1859:35, pl. 37, fig. 1-3 [Holotype.--apparently USNM 3237 (cited as 3207 by Cope, 1887:16), now lost, from 40 Leagues from (probably north) Mexico City; collected by John Potts]. Kellogg, 1932:107.

_Epirhexis longipes_: Cope, 1866:96.

_Eleutherodactylus longipes_: Kellogg, 1932:107 (part). Smith and Taylor, 1948:61. Lynch, 1963:580-581. Gorham, 1966:82.

_Syrrhophus latodactylus_ Taylor, 1940d:396-401, pl. 43, figs. A-F, text fig. 7 [Holotype.--FMNH 100063 (formerly EHT-HMS 6807), from Huasteca Canyon, 15 km. W Monterrey, Nuevo Leon, Mexico, 680 m.; collected on June 20, 1936, by Edward H. Taylor]. Smith and Taylor, 1948:50-52. Martin, 1958:48-50. Gorham, 1966:165.

_Diagnosis._--Large frogs, males 22.1-33.2 mm. snout-vent, females 26.8-39.6 mm. snout-vent length; vocal slits lacking in males; digital tips greatly expanded (more than twice the width of digit); first finger shorter than second; skin of dorsum pustular, that of venter smooth; diameter of tympanum in males 61.1-87.2 per cent that of eye, 49.5-72.1 per cent in females; dorsum tan with large or small spots and blotches; limbs banded; interorbital bar or triangle present.

_Remarks._--I have applied Baird's _Batrachyla longipes_ to the frog Taylor (1940d) called _Syrrhophus latodactylus_ because the color pattern (Fig. 13) predominant in the southern part of the range agrees with that described (figured) for _Batrachyla longipes_.

The color pattern of individuals in the southern part of the range of this species consists of large spots or blotches, whereas in the northwestern part the pattern is made up of smaller spots. In the northeastern part of the range, the pattern is more reduced and tends to consist of heavy flecking. The interorbital bar is narrower in specimens from Nuevo Leon and Tamaulipas and is triangular in specimens from Hidalgo and Queretaro.

The status of the name _Batrachyla longipes_ is currently that of a _nomen dubium_ (Lynch, 1963). At that time, I was unaware of the geographic variation in color pattern in _Syrrhophus latodactylus_.

The exact type-locality of _Batrachyla longipes_ is not known. If it is 40 Leagues north of Mexico City, the locality would be in an area where the species has a blotched instead of a flecked or spotted pattern. No justifiable evidence was presented to place _Batrachyla longipes_ in _Eleutherodactylus_ instead of _Syrrhophus_. Barbour (1923) and Kellogg (1932) a.s.sociated another species (_E. batrachylus_) with _longipes_.

Taylor (1940a) noted this as a case of misidentification and corrected the error but left _longipes_ in the genus _Eleutherodactylus_. Lynch (1963) noted several points of morphological agreement between _Syrrhophus_ and _B. longipes_ but did not place _longipes_ in _Syrrhophus_.

Baird's (1859) figures of the holotype do not ill.u.s.trate prevomerine teeth, but according to Cope (1866) they were present in the holotype.

The digital tips of the frog in the figure are somewhat narrower than those typically seen in _S. latodactylus_. If the specimen was slightly desiccated, as possibly was the case, the digits would appear narrower.

There is no evidence contrary to placing _Syrrhophus latodactylus_ in the synonymy of _Batrachyla longipes_.

[Ill.u.s.tration: FIG. 13: Dorsal views of _Syrrhophus longipes_ ill.u.s.trating geographic variation in pattern (left, TCWC 12179, 1.5; right, KU 92572, 1.8); side of head (TCWC 10966, 6).]

Application of Baird's name _Batrachyla longipes_ to the species of frog heretofore called _Syrrhophus latodactylus_ poses one serious problem.

_Batrachyla longipes_ is the type-species (by original designation) of the genus _Epirhexis_ Cope, 1866, which has priority over _Syrrhophus_ Cope, 1878. If _Batrachyla longipes_ is left in the status of a _nomen dubium_, _Epirhexis_ can be forgotten, for the two names are tied together. However, since it seems almost certain that _Batrachyla longipes_ and _Syrrhophus latodactylus_ are conspecific, the former name should not be left as a _nomen dubium_. _Epirhexis_ never came into general usage (Cope cited the name four times, but no one else has used it), whereas _Syrrhophus_ is well established in the zoological literature. It would serve only to confuse the literature to adhere strictly to the Law of Priority and replace _Syrrhophus_ with _Epirhexis_. Therefore, _Syrrhophus_ is used in this paper, even though _Epirhexis_ has priority. A request for the suppression of _Epirhexis_ Cope, 1866, has been submitted to the International Commission of Zoological Nomenclature (Lynch, 1967).

_Etymology._--Latin, meaning long-footed; Taylor's _latodactylus_ refers to the wide digital pads.

[Ill.u.s.tration: FIG. 14: Distribution of _Syrrhophus dennisi_ (triangle) and _S. longipes_ (circles).]

_Distribution._--Moderate elevations (650 to 2000 meters) along the Sierra Madre Oriental from central Nuevo Leon to northern Hidalgo, Mexico (Fig. 14).

_Specimens examined._--(122) MeXICO, _Hidalgo_: 3 km. NE Jacala, AMNH 52977; 9.6 km. NE Jacala, 1800 m., TCWC 10966-70, 12179; 8 km. S Jacala, La Placita, 1850 m., FMNH 100266-68, 103244, UIMNH 13291, 13327. _Nuevo Leon_: Salto Cola de Caballo, KU 92572; Huasteca Canyon, 15 km. W Monterrey, 680 m., FMNH 100063 (holotype of _S. latodactylus_), UIMNH 13290; 6.5 km. N Pablillo, EAL 1319; Sabinas Hidalgo, USNM 139728.

_Queretaro_: Cueva de los Riscos, 8 km. SW Jalpan, KU 106300. _San Luis Potosi_: 13 km. E Santa Barberita, LSUMZ 2295; second camp, San Luis Potosi road, UIMNH 13326; Xilitla, Cueva sin nombre, UMMZ 125892.

_Tamaulipas_: 4 km. W El Carrizo, 500 m., UMMZ 111343 (31); 8 km. N Chamal, Bee Cave, KU 106299; 14.5 km. NNW Chamal, 420 m., UMMZ 111339-40, 111342 (4), 111344 (11); 19 km. NNW Chamal, 700 m., UMMZ 111341 (3); El Chihue, 1880 m., UMMZ 111289 (4); 11 km. N Gomez Farias, 1060 m., UMMZ 101166; 11 km. WNW Gomez Farias, 1800 m., UMMZ 108507 (3); 8 km. NW Gomez Farias, 1060-1400 m., LSUMZ 11085, UMMZ 101167 (3), 101168 (4), 101169 (2), 101170 (3), 101171 (2), 101360-61, 102860, 102933 (4), 102934 (2), 102935-38, 102939 (2), 102940-43, 108800 (3), 110735, 111345-46.

=Syrrhophus pipilans= Taylor

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

_Diagnosis._--Medium sized frogs, males 22.6-28.5 mm. snout-vent, females 21.1-29.4 mm. snout-vent length; vocal slits present in males; finger tips slightly expanded, truncate in outline; inner metatarsal tubercle less than twice the size of outer; skin of dorsum smooth to s.h.a.greened, that of venter smooth; tympanum 36.5-54.0 per cent diameter of eye; dorsum dark brown with large or small light brown, orange-brown, or yellowish spots or blotches; limbs banded; interorbital bar absent.

[Ill.u.s.tration: FIG. 15: Dicegrams of ear size relative to eye diameter in the two subspecies of _Syrrhophus pipilans_. N = 17 in _nebulosus_, 18 in _pipilans_.]

_Remarks._--Two subspecies were recognized by Duellman (1958).

Previously both had been treated as species. The two populations were distinguished on the basis of color pattern and the size of the tympanum. Measurements of 17 males of _S. p. nebulosus_ from central Chiapas and 18 males of _S. p. pipilans_ from southcentral Oaxaca and Guerrero, Mexico, demonstrates that the supposed difference in tympanum size is not significant (Fig. 15). There is, however, a tendency for the western population of _S. pipilans_ to have larger tympani. Based on the present examination of 112 specimens of this species the two populations are held to be sufficiently distinct to warrant taxonomic recognition as subspecies (Fig. 16).

[Ill.u.s.tration: FIG. 16: _Syrrhophus pipilans nebulosus_ (left, KU 58908) and _S. p. pipilans_ (right, KU 86885). 2.7.]

The parotoid glands attributed to this species by Taylor (1940c:95) are merely the superficial expression of the _m. depressor mandibulae_ and scapula. No true glands are present in the parotoid region.