A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope Part 3

_Diagnosis._--Small frogs, males 18.2-23.5 mm. snout-vent, females 19.0-22.5 mm. snout-vent length (small sample); vocal slits in males; digital tips scarcely expanded (Fig. 1); first finger shorter than second; outer palmar tubercle reduced in size; inner metatarsal tubercle elongate, twice the size of small, conical outer metatarsal tubercle; diameter of tympanum 35.5-46.5 per cent that of eye in both s.e.xes; dorsum brown with small pale spots (red in life); venter gray.

_Remarks._--Previous authors who treated _Syrrhophus_ placed this species in the western complex, because it occurs on the Pacific versant and has a reduced outer palmar tubercle. Duellman (1958) placed _rubrimaculatus_ apart from the other western species, because of its relatively unexpanded digital tips and coloration. The digital tips are like those in _leprus_, which _rubrimaculatus_ resembles. Except for the reduction of the outer palmar tubercle, _rubrimaculatus_ could be a member of the _leprus_ group.

_Syrrhophus rubrimaculatus_ is probably best treated as a Pacific derivative of the _leprus_ group, even though the palmar tubercles do not agree. The removal of _rubrimaculatus_ from the western complex results in a more h.o.m.ogeneous remainder and does not greatly increase the heterogeneity of the eastern complex.

_Etymology._--Latin, meaning spotted with red; in reference to the colors in life.

_Distribution._--Low to moderate elevations on the Pacific versant of southeastern Chiapas, Mexico (Fig. 7); probably extending into adjacent Guatemala.

_Specimens examined._--(48) MeXICO, _Chiapas_: Escuintla, UMMZ 88283; 6 km. NE Escuintla, UMMZ 87876-80; La Esperanza, UIMNH 13285, UMMZ 88496-97, USNM 114070 (holotype), 114054-69, 114072; Monte Cristo, UMMZ 88353; 1.3 km. N Puerto Madero, KU 58910-11; Finca San Jeronimo, 600-650 m., UIMNH 55299-312, 55313-16 (cleared and stained).

=Syrrhophus guttilatus= (Cope)

_Malachylodes guttilatus_ Cope, 1879:264 [Holotype.--USNM 9888, from Guanajuato, Guanajuato, Mexico; collected in 1877 by Alfredo Duges].

_Syrrhopus guttulatus_: Boulenger, 1888:204-06.

_Syrrhaphus guttulatus_: Gunther, 1900:317.

_Syrraphus guttulatus_: Diaz de Leon, 1904:11.

_Syrrhophus guttilatus_: Nieden, 1923:399-400. Kellogg, 1932:125, 127-28. Smith and Taylor, 1948:49, 51. Firschein, 1954:52-54.

Gorham, 1966:164.

_Syrrhophus smithi_ Taylor, 1940b:43-45, pl. 1 [Holotype.--USNM 108594, from 15 mi. SW Galeana, Nuevo Leon, Mexico, 1575 m.; collected on October 13, 1939, by Hobart M. Smith]. Smith and Taylor, 1948:49, 51. Firschein, 1954:54-55. Martin, 1958:50.

Gorham, 1966:167.

_Syrrhophus gaigeae_ Schmidt and Smith, 1944:80 [Holotype.--FMNH 27361, from the Basin, Chisos Mountains, Brewster Co., Texas; collected on July 24, 1937, by Walter L. Necker].

_Syrrhophus petrophilus_ Firschein, 1954:50-52 [Holotype.--UIMNH 7807, from 5 km. SW San Luis Potosi, San Luis Potosi, Mexico; collected on July 18, 1949, by David Langebartel]. Gorham, 1966:166.

_Syrrhophus marnocki_: Milstead, Mecham, and McClintock, 1950:548 (in part).

_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).

[Ill.u.s.tration: FIG. 9: _Syrrhophus guttilatus_ (upper left, UIMNH 55519, 1.4; lower left, UIMNH 55519, 2.3) and _S. marnockii_ (upper right, TCWC 9317, 1.4; lower right, TCWC 13510, 2.1).]

_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 s.e.xual 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 Mexico. 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. w.i.l.l.y Wrenn told me that he has seen heavy infestations of _Hannemania_ on _Syrrhophus pallidus_. Infestation by _Hannemania_ probably reflects similar ecologies rather than close relations.h.i.+ps.

[Ill.u.s.tration: FIG. 10: Distribution of _Syrrhophus guttilatus_.]

_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, Mexico (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.

MeXICO: _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 Leon_: 3 km.

S Galeana, JDL 1215 (skeleton), UIMNH 58204; 24 km. SW Galeana. 1575 m., USNM 108594 (holotype of _Syrrhophus smithi_). _San Luis Potosi_: 5 km.

SW San Luis Potosi, 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; Cla.s.sen 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).

[Ill.u.s.tration: FIG. 11: Distribution of _Syrrhophus marnockii_ (circles). Starred localities are late Pleistocene records.]

=Syrrhophus verrucipes= Cope

_Syrrhophus verrucipes_ Cope, 1885:383 [Holotype.--ANSP 11325, from near Zacualtipan, Hidalgo, Mexico (1800 feet lower in a rocky gorge of a stream near its junction with the Rio 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_: Gunther, 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, Mexico, 1850 m.; collected on July 2, 1936, by Edward H. Taylor]. Smith and Taylor, 1948:47-48.