Part 3

_Diagnosis._--Medium-sized frogs, males 20.6-29.0 mm. snout-vent, females 25.7-31.0 mm. snout-vent length; vocal slits in males; digital tips slightly expanded (Fig. 1); first and second fingers equal; skin of dorsum smooth to moderately pustular, that of venter smooth; snout blunt; diameter of tympanum 55.1-75.7 per cent that of eye in males, 47.6-61.7 in females; dorsum and flanks cream to gray with light brown to black flecking and vermiculations; thighs usually not banded; interorbital bar present (Fig. 8).

_Remarks._--Cope (1879) distinguished _Malachylodes_ from _Syrrhophus_ on the basis of the presence of a frontoparietal fontanelle in the holotype of _guttilatus_. The holotype is a juvenile female and as is the case in the juveniles of nearly all leptodactylids, a frontoparietal fontanelle is present. Firschein (1954) used the presence of the fontanelle to distinguish _guttilatus_ from his _petrophilus_.

As is clearly evident from the length of the synonymy, I consider a number of currently used names to be synonymous with _guttilatus_. I have seen the holotypes of all four names and am unable to recognize more than a single species. The holotype of _petrophilus_ is a male, whereas that of _smithi_ is a female. The supposed differences are a reflection of sexual dimorphism in the size of the eye (Table 5). The two holotypes, as well as those of _gaigeae_ and _Malachylodes guttilatus_ agree in color pattern.

Schmidt and Smith (1944) named _Syrrhophus gaigeae_ from the Chisos Mountains of the Big Bend region of Texas and compared it only with _S. marnockii_. Milstead, Mecham and McClintock (1950) synonymized _gaigeae_ and _marnockii_ because they were unable to verify the characters Wright and Wright (1949) used to separate them. Specimens from the Big Bend region differ from those of the Edward and Stockton Plateaus in having a vermiculate pattern, an interorbital bar, and a supratympanic stripe. In these respects they agree with specimens from northern México. Based on limited observations, the Mexican population is yellowish to brownish in life whereas the central Texas population is green in life. Lacking evidence of genetic exchange, the two are held to be specifically distinct.

Nearly every specimen examined was infested with chiggers of the genus _Hannemania_. The greatest concentrations are on the venter, in the groin, and on the thighs. Many specimens have chiggers on the digits and tarsi. The same, or a related, chigger was found on many specimens of _Syrrhophus marnockii_ and a few _S. verrucipes_, but on no other species of the genus. Mr. Willy Wrenn told me that he has seen heavy infestations of _Hannemania_ on _Syrrhophus pallidus_. Infestation by _Hannemania_ probably reflects similar ecologies rather than close relationships.

_Etymology._--Latin, _guttula_, meaning spotting or flecking, in reference to the color pattern.

_Distribution._--Moderate to intermediate elevations (600 to 2000 m.) along the Sierra Madre Oriental from the Big Bend Region of Texas to Guanajuato, México (Fig. 10).

_Specimens examined._--(32) TEXAS, _Brewster Co._: Juniper Canyon, Chisos Mts., FMNH 27361 (holotype of _S. gaigeae_), 27360, 27362-63, MCZ 15346, 27801, UMMZ 66080, 66082, 66085-91, USNM 76876; Upper Green Gulch, TCWC 15943.

MÉXICO: _Coahuila_: 8 km. S Saltillo, UIMNH 55518-21. _Guanajuato_: Guanajuato, USNM 9888 (holotype of _Malachulodes guttilatus_); 8 km. E Guanajuato, AMNH 73425; Cerro Cubilete, AMNH 73424. _Nuevo León_: 3 km. S Galeana, JDL 1215 (skeleton), UIMNH 58204; 24 km. SW Galeana. 1575 m., USNM 108594 (holotype of _Syrrhophus smithi_). _San Luis Potosí_: 5 km. SW San Luis Potosí, UIMNH 7807 (holotype of _S. petrophilus_). _Tamaulipas_: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4).

=Syrrhophus marnockii= Cope

_Syrrhophus marnockii_ Cope, 1878:253 [Syntypes.--ANSP 10765-68, from "near San Antonio," Bexar Co., Texas; collected by G. W. Marnock].

_Syrrhophus marnocki_: Yarrow, 1882:24, 193. Milstead, Mecham, and McClintock, 1950:550.

_Diagnosis._--Medium-sized frogs, males 18.4-28.9 mm. snout-vent, females 20.4-35.4 mm. snout-vent length; vocal slits in males; digital tips widened (Fig. 1); first and second fingers equal; skin of dorsum smooth to weakly pustular, that of venter smooth; snout blunt, rounded; diameter of tympanum 47.2-68.3 per cent that of eye in males, 45.8-73.3 in females; dorsum tan to light brown in preservative with rusty-brown flecks, venter white; ground color green in life; thighs banded; interorbital bar absent.

_Remarks._--Specimens from the southern edge of the Edwards Plateau and the eastern edge of the Stockton Plateau have larger flecks on the back that tend to form a vermiculate pattern like that of _S. guttilatus_. The vermiculation is never well developed (see plate 38 in Conant, 1958). Most of the specimens from the Edwards Plateau have a punctate pattern (Fig. 9).

Fossils are known from the Sangamon interglacial deposits in Foard and Knox Counties, Texas (Lynch, 1964; Tihen, 1960).

_Etymology._--A patronym for the collector of the type specimens.

_Distribution._--The Edwards Plateau and the extreme eastern edge of the Stockton Plateau in Texas (Fig. 11). The fossil records lie some 200 miles to the north. Two specimens (FMNH 103216-17) from Brownsville, Cameron Co., Texas, were formerly in the EHT-HMS collection (nos. 31348-49). Data given in Taylor's field catalogue (housed in the Division of Reptiles, Field Museum) are "Brownsville, A. J. Kirn collector, April 15, 1934." Until verification by recently collected material is available, this record must be disregarded.

_Specimens examined._--(103) TEXAS, _Bandera Co._: 10 mi. SW Medina, TCWC 13508-10; 8 mi. W Medina, KU 60243; 13 mi. W Medina, KU 60242, TCWC 13506-07. _Bexar Co._: UIMNH 34694; Classen ranch, near San Antonio. UMMZ 98891; Helotes, EAL 1560, MCZ 11837 (2), UMMZ 64045, USNM 13635; 2 mi. N Helotes, TCWC 9234-35; 3.5 mi. N Helotes, LSUMZ 10363; 8 mi. N Helotes, TCWC 1549, 4364; San Antonio, FMNH 15553-56, TCWC 13497-99. _Blanco Co._: 8 mi. NE Blanco, TCWC 4782. _Comal Co._: New Braunfels, TCWC 13500-05; 5 mi. NE New Braunfels, UMMZ 71016 (10). _Hays Co._: San Marcos, AMNH 22661-64, 32700, FMNH 15245-46, 26250, 26253-57, 37617, 37665, MCZ 15649-50, 23268-69; 6 mi. SW San Marcos, TCWC 5070-71, 7140, 9232-33, 9236, 9316-17, 9320. _Kendall Co._: 11 mi. E Boerne, AMNH 54660-61, 54662 (2); 10 mi. W Boerne, KU 18441; Kendalia, UIMNH 21434. _Kerr Co._: Kerr W. M. Area, TCWC 15859; 40 mi. NW Kerrville, TCWC 6555. _Medina Co._: UIMNH 13287-88; 12 mi. N Castroville, UIMNH 21423; 14 mi. N Castroville, UIMNH 21424-25; 16 mi. N Castroville, UIMNH 21421-22; 17 mi. N Castroville, UIMNH 21428-29; 18 mi. N Castroville, UIMNH 21426-27, 21430-33; 6.5 mi. NW Rio Medina, KU 18440. _Real Co._: Rio Frio, FMNH 55156-57. _Travis Co._: Austin, AMNH 44221-22; Mount Bonnell, 5 mi. S Austin, UMMZ 101453 (10). _Uvalde Co._: 13 mi. from Uvalde, UIMNH 62322. _Val-Verde Co._: 40 mi. N Del Rio, JDL 214 (skeleton).

=Syrrhophus verrucipes= Cope

_Syrrhophus verrucipes_ Cope, 1885:383 [Holotype.--ANSP 11325, from near Zacualtipán, Hidalgo, México (1800 feet lower in a rocky gorge of a stream near its junction with the Río San Miguel), collected by Dr. Santiago Bernard]. Kellogg, 1932:126-29. Smith and Taylor, 1948:52-53. Firschein, 1954:55-57. Gorham, 1966:167.

_Syrrhaphus verrucipes_: Günther, 1900:216-17.

_Tomodactylus macrotympanum_ Taylor, 1940e:496-99, pl. 55, figs. 2a-b. [Holotype.--FMNH 100049 (formerly EHT-HMS 6838), from La Placita, 8 km. S Jacala, Hidalgo, México, 1850 m.; collected on July 2, 1936, by Edward H. Taylor]. Smith and Taylor, 1948:47-48.

_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, subacuminate; 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 Potosí, Queretaro, and northwestern Hidalgo, México (Fig. 7).

_Specimens examined_--(43) MÉXICO, _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 Zacualtipán, ANSP 11325 (holotype of _Syrrhophus verrucipes_). _Queretaro_: 3.5 km. S San Juan del Río, EAL 1343. _San Luis Potosí_: 9.6 km. W Ahuacatlán, LSUMZ 4968-70.

=Syrrhophus dennisi= new species

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

_Holotype._--UMMZ 101121, adult male from a cave near El Pachón, 8 km. N Antiguo Morelos, Tamaulipas, México, 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 shagreened 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 acuminate 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 sexes; 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 Pachón 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).

_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) México 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 León, México, 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 León 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 México 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) associated 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 illustrate 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_.

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.

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

_Specimens examined._--(122) MÉXICO, _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 León_: 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 Potosí_: 13 km. E Santa Barberita, LSUMZ 2295; second camp, San Luis Potosí 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 Gómez Farías, 1060 m., UMMZ 101166; 11 km. WNW Gómez Farías, 1800 m., UMMZ 108507 (3); 8 km. NW Gómez Farías, 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 Mazatlán, Guerrero, México; 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 shagreened, 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.

_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, México, 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).

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.

=Syrrhophus pipilans nebulosus= Taylor

_Syrrhophus nebulosus_ Taylor, 1943:353-55, pl. 27, figs. 3-5 [Holotype.--FMNH 100095 (formerly EHT-HMS 3774), near Tonolá, Chiapas, México; 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.