A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope Part 2

=Syrrhophus cystignathoides campi= Stejneger, New combination

_Syrrhophus campi_ Stejneger, 1915:131-32. [Holotype.--USNM 52290, from Brownsville, Cameron Co., Texas; R. D. Camp collector, March 31, 1915]. Smith and Taylor, 1948:52. Martin, 1958:50.

_Diagnosis._--Venter smooth; usually no interorbital light and dark bars present; ground color brown in life (Fig. 4a).

_Remarks._--Martin (1958) was the first author to point out that _S. campi_ was probably a subspecies of the more southern _S. cystignathoides_. Various references in the literature might lead one to believe that the two were sympatric over much of northeastern Mexico; this error was created by the use of a single character (condition of the skin of the venter) to characterize the two populations. Specimens from southern Texas have a smooth venter, lack interorbital bars and have, in general, a brown ground color, whereas specimens from central Veracruz have an areolate venter, interorbital light and dark bars and a yellow ground color. In southern Tamaulipas and eastern San Luis Potosi, these characters vary discordantly, thereby strongly suggesting that the two populations intergrade. Both populations agree in other morphological characters; therefore, they are here treated as geographic variants.

_Etymology._--Named for the collector of the type specimens, Mr. R. D.

Camp of Brownsville, Texas.

_Distribution._--Lower Rio Grande embayment in Texas to central Nuevo Leon and Tamaulipas, Mexico. Intergrades are known from southern Tamaulipas and adjacent San Luis Potosi, Mexico (Fig. 5).

_Specimens examined._--(113) TEXAS, Cameron Co.: MCZ 10277-85, 10286 (10); Brownsville, AMNH 3215, 3218-20, 3221 (3), 5376, 62117, FMNH 105336, KU 8135-39, MCZ 3738-42, 3743 (10), TCWC 5908, 7139, TNHC 92-94, 20909, UMMZ 51760, 54031 (5), USNM 52290 (holotype); 22 mi. SE Brownsville, TNMC 14223; 8 mi. SW Brownsville, UMMZ 101127 (3); Harlingen, AMNH 62118, UMMZ 105200-205, 105206 (5), 105207 (4). _Hidalgo Co._: Bentsen-Rio Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC 7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60.

MEXICO, _Nuevo Leon_: Salto Cola de Caballo, AMNH 57953-54, FMNH 30644-45, 37169-70; Monterrey, UIMNH 13324; 40 km. SE Monterrey, UIMNH 3686. _Tamaulipas_: 80 km. Matamoros, FMNH 27150 (13).

Intergrades [_S. c. cystignathoides_ _S. c. campi_ (88)] MeXICO, _San Luis Potosi_: 5 km. E Ciudad del Maiz, UMMZ 106435; 16 km. W Naranjo, FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC 6980.

_Tamaulipas_: 5 km. W Acuna, 1060 m., UMMZ 101172, 101173 (16), 101174-76, 101177 (6); 14.5 km. NNW Chamal, 430 m., UMMZ 111337 (2); 20 km. NNW Chamal, 700 m., UMMZ 111338 (11); 8 km. N Gomez Farias, 450 m., UMMZ 101165; 8 km. NE Gomez Farias, Pano Ayuctle, UMMZ 102264, 102924 (6); 8 km. NW Gomez Farias, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 (5), 102929-32, 110124 (3); Rio Guayala, near Magiscatzin, MCZ 24138-42, 85071-81, UMMZ 88242 (2); Magiscatzin, TCWC 6981; Las Yucas, north of Aldama, MCZ 29665-68; 16 km. NE Zamorina, UMMZ 101124.

[Ill.u.s.tration: FIG. 4: _Syrrhophus cystignathoides campi_ (left, TCWC 13490) and _S. c. cystignathoides_ (right, KU 105500).

Dorsal views 2, sides of heads 3.]

=Syrrhophus cystignathoides cystignathoides= (Cope), New combination

_Phyllobates cystignathoides_ Cope, 1877:89-90 [Syntypes.--USNM 32402-32409, from Potrero, near Cordoba, Veracruz, Mexico, collected by Francis Sumichrast]. Boulenger, 1882:196.

_Syrrhophus cystignathoides_: Cope, 1879:268. Kellogg, 1932: 126-27. Taylor and Smith, 1945: 582-83. Smith and Taylor, 1948:50.

Martin, 1958:49.

_Syrrhaphus cystignathoides_: Gunther, 1900:218.

_Syrraphus cystignathoides_: Diaz de Leon, 1904:10.

_Syrrhopus cystignathoides_: Barbour and Loveridge, 1946:170.

[Ill.u.s.tration: FIG. 5: Distribution of _Syrrhophus cystignathoides campi_ (solid symbols) and the nominate subspecies (open symbols).]

_Diagnosis._--Venter areolate; interorbital light and dark bars present; ground color yellow to brownish-yellow in life (Fig. 4b).

_Remarks._--Firschein (1954) briefly considered the status of Peters'

(1871) _Phyllobates verruculatus_ and noted that if it was a _Syrrhophus_ it would probably be referrable to _S. cystignathoides_.

Peters' (1871) original description corresponds well with _S. cystignathoides_, and the type-locality ("Hua.n.u.sco" = Huatusco) is within the range of that species. Firschein (1954) expressed doubt that _verruculatus_ was a _Syrrhophus_, because Peters placed it in another genus. However, Peters described _verruculatus_ a decade before Cope diagnosed the genus Syrrhophus. Most frogs now called _Syrrhophus_, plus a number of lower Central American frogs now placed in a variety of genera were placed in _Phyllobates_ by Boulenger, Cope, and Peters.

The types of _Phyllobates verruculatus_ were destroyed during World War II (Gunther Peters, _in litt._); the specimens subsequently a.s.signed to the taxon by Kellogg (1932) are _Syrrhophus cystignathoides_. Because the type specimens are lost and because the name antedates the more established name, _cystignathoides_, I favor retaining _Phyllobates verruculatus_ Peters as a _nomen dubium_.

Smith and Taylor (1948) reported _S. verruculatus_ from Tianguistengo, Hidalgo, Mexico. These specimens are examples of _verrucipes_. Smith (1947) reported a specimen of _verruculatus_ from San Lorenzo, Veracruz.

Firschein (1954) referred it to _cystignathoides_, and Duellman (1960) concluded that both authors were in error and that the specimen (USNM 123530) was a _leprus_.

_Etymology._--The trivial name is the diminutive of _Cystignathus_, a once-used generic name for several leptodactylid frogs.

_Distribution._--Low and moderate elevations in the foothills along the Sierra Madre Oriental from eastern San Luis Potosi to Central Veracruz, Mexico (Fig. 5).

_Specimens examined._--(130), MeXICO, _Puebla_: Necaxa, UMMZ 69519-20.

_San Luis Potosi_: 5 km. W Aguismon, LSUMZ 4962-63; along Rio Axtla, road to Xilitla, UMMZ 105500; Tamazunchale, UIMNH 3199; 6.5 km. N Tamazunchale, UMMZ 104039; 8 km. N Tamazunchale, UMMZ 119490.

_Veracruz_: Coatepec, 1210 m., FMNH 704966-67; 11 km. SE Coatepec, 850 m., FMNH 70468-70; below Cordoba, FMNH 104588, UIMNH 13321; Cuautlapam, 1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortin de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortin de las Flores, UIMNH 42799-808, UMMZ 105389; 3.2 km. N Fortin de las Flores, UIMNH 26633-35; 4.8 km. N Fortin de las Flores, UIMNH 71967-68; 3.2 km. W Fortin de las Flores (Barranca Metlac), 910 m., UIMNH 49294-95, UMMZ 115444-46, 118221, 119893 (2); Huatusco, KU 100363; Jalapa, 1400 m., FMNH 70440, 70443-51, 70454-65; 16 km. NE Jalapa, 1300 m., FMNH 70452-53; 8 km. E Jalapa, UIMNH 13338; 9.5 km. S Jalapa, UMMZ 122083 (2); Mirador, KU 23967; Paraja Nuevo, El Suchil, UMMZ 85490 (7), 85491 (2), 90315; La Pa.s.sa, UIMNH 49293, 49297; 1 km. E Plan del Rio, 240 m., UMMZ 102067 (2); Potrero Viejo, FMNH 104583, 104586, 105326-27, KU 26789, 100357-62, UIMNH 13323, 13340-43; USNM 32402 (lectotype), 32403-04, 32406-09; 9.6 km. S Santa Rosa, TCWC 12785; 24 km. NE Tezuitlan (Puebla), UMMZ 105388; Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, 70441-42; 9.6 km. NW Tihuatlan, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU 23966; 26 km. NW Tuxpan, UMMZ 126419.

=Syrrhophus leprus= Cope

_Syrrhophus leprus_ Cope, 1879:268-69 [Holotype.--USNM 10040, from Santa Efigena, Oaxaca, Mexico, Francis Sumichrast collector].

Kellogg, 1932:124-5, 128. Taylor and Smith, 1945:582. Smith and Taylor, 1948:50-51. Duellman, 1958:8, pl. 1, Fig. 2; 1960:56-57.

Gorham, 1966:165.

_Syrrhaphus leprus_: Gunther, 1900:217.

_Syrrhophus leprus leprus_: Neill, 1965:85-86.

_Syrrhophus leprus cholorum_ Neill, 1965:85-86 [Holotype.--Wilfred T. Neill collection 1525, from 3.9 mi. N San Antonio, Toledo District, British Honduras, collected October 28, 1959, by R. A. Allen, T. C. Allen, and W. T. Neill].

_Diagnosis._--Medium-sized frogs, males 20.5-26.5 mm. in snout-vent, females 22.0-29.3 mm. in snout-vent length; vocal slits present in males; tips of fingers dilated slightly; first finger longer than second; inner metatarsal tubercle twice size of small, conical outer metatarsal tubercle; skin of dorsum pustular, that of venter smooth; snout subac.u.minate; diameter of tympanum 47.5-62.5 per cent of eye in males, 38.6-57.9 per cent in females; dorsum yellowish-green with chocolate brown blotches or spots forming reticulations in most specimens; venter white to gray; flanks brown, spotted with white or not; limbs banded; interorbital bar obscured by dorsal pattern.

[Ill.u.s.tration: FIG. 6: Dorsal views of _Syrrhophus leprus_ showing variation in dorsal pattern (left, UMMZ 121244, 2; right, KU 26106, 1.7). Side of head (UIMNH 42726, 7).]

[Ill.u.s.tration: FIG. 7: Distribution of three species of eastern complex _Syrrhophus_: _leprus_ (circles), _rubrimaculatus_ (triangles), and _verrucipes_ (squares).]

_Remarks._--My distribution map (Fig. 7) differs somewhat from that of Duellman (1958), who was unaware of specimens reported by Taylor and Smith (1945) from central Veracruz, Mexico.

Duellman (1958, 1960) regarded _S. leprus_ as having a gray venter.

Neill (1965) characterized his new subspecies on the basis of white venter and spots on the dorsum. Some specimens from throughout the range have only small round spots, instead of vermiculations (Fig. 6). The gray ventral coloration is largely restricted to the population in Los Tuxtlas, Veracruz, but only about 80 per cent of the specimens from the Los Tuxtlas have gray venters, whereas specimens from Guatemala, Oaxaca, Tabasco, and central Veracruz, Mexico, have white venters (rarely gray).

Since the specimens from British Honduras are not distinct from specimens throughout most of the range, there is no reason to recognize them as a subspecies.

_Etymology._--Greek, _lepra_, leprosy, in reference to the mottled color pattern.

_Distribution._--Discontinuous; central Veracruz to British Honduras to low elevations in the foothills of the Sierra Madre Oriental, Los Tuxtlas, Sierra Madre de Chiapas (Isthmus of Tehuantepec (Fig. 7)).

_Specimens examined._--(84). GUATEMALA, _Alta Verapaz_: Chinaja, KU 55961-62. _El Peten_: 15 km. NW Chinaja, KU 55963; Piedras Negras, USNM 114085-92; Tikal, UMMZ 117035; Uaxactun, AMNH 55121-22.

MeXICO, _Oaxaca_: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La Gloria, USNM 114093; 30.5 km. N Matias Romero, UIMNH 39459, 71969; Santa Efigenia, USNM 10040 (holotype). _Tabasco_: Teapa, UMMZ 113799-800; 13.5 km. W Teapa, UMMZ 120253. _Veracruz_: 27.5 km. N Acayucan, UIMNH 42726; Atoyac, UIMNH 13331, 49296; 3.2 km. N Catemaco, UIMNH 71976-77; Coyame, UIMNH 38995, 38998, 40342; Dos Amates, TCWC 21211; Fortin de Las Flores, FMNH 113751, 113753; Paraja Nuevo, El Suchil, UMMZ 90315; Potrero Viejo, FMNH 113743-50, 126114-18, KU 26104-06, UIMNH 13332-37, UMMZ 88837; San Andres Tuxtla, UIMNH 27123-31, 28611, 71975, UMMZ 115450 (5); San Lorenzo, USNM 123530; 4.5 km. NW Santiago Tuxtla, JDL 992 (skeleton), UIMNH 27122; 32 km. S Sayula, EAL 1696; Tepalapan, 1.6 km. S Catemaco, UMMZ 118222 (2); Volcan San Martin, south slope, UMMZ 118223; Volcan San Martin, Rancho El Tular, UIMNH 35399-400, 40340-41.

=Syrrhophus rubrimaculatus= Taylor and Smith

_Syrrhophus rubrimaculatus_ Taylor and Smith, 1945:583-85 [Holotype.--USNM 114070, from La Esperanza, near Escuintla, Chiapas, Mexico, collected May 13, 1940, by H. M. and R. Smith].

Duellman, 1958:1-4, 7, 12, 14. Gorham, 1966:167.

_Syrrhophus rubrimaculata_: Smith and Taylor, 1948:48-49.

[Ill.u.s.tration: FIG. 8: _Syrrhophus rubrimaculatus_ (upper right, KU 58911, 1.6; lower right, KU 58910, 4) and _S. verrucipes_ (upper left, UIMNH 15995, 1.6; lower left, UIMNH 15989, 3.7).]