Chapter 3

Syrrhophus cystignathoides campiStejneger, New combinationSyrrhophus campiStejneger, 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 thatS. campiwas probably a subspecies of the more southernS. cystignathoides. Various references in the literature might lead one to believe that the two were sympatric over much of northeastern México; 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 Potosí, 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 Río Grande embayment in Texas to central Nuevo León and Tamaulipas, México. Intergrades are known from southern Tamaulipas and adjacent San Luis Potosí, México (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-Río Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC 7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60.MEXICO,Nuevo León: 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).Fig. 4:Syrrhophus cystignathoides campi(left, TCWC 13490) andS. c. cystignathoides(right, KU 105500). Dorsal views ×2, sides of heads ×3.Intergrades [S. c. cystignathoides×S. c. campi(88)] MÉXICO,San Luis Potosí: 5 km. ECiudaddel Maiz, UMMZ 106435; 16 km. W Naranjo, FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC 6980.Tamaulipas: 5 km. W Acuña, 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 Gómez Farías, 450 m., UMMZ 101165; 8 km. NE Gómez Farías, Pano Ayuctle, UMMZ 102264, 102924 (6); 8 km. NW Gómez Farías, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 (5), 102929-32, 110124 (3); Río 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.Syrrhophus cystignathoides cystignathoides(Cope), New combinationPhyllobates cystignathoidesCope, 1877:89-90 [Syntypes.—USNM 32402-32409, from Potrero, near Córdoba, Veracruz, México, 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: Günther, 1900:218.Syrraphus cystignathoides: Díaz de León, 1904:10.Syrrhopus cystignathoides: Barbour and Loveridge, 1946-170[** 1946:170].Fig. 5:Distribution ofSyrrhophus 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 verruculatusand noted that if it was aSyrrhophusit would probably be referrable toS. cystignathoides. Peters' (1871) original description corresponds well withS. cystignathoides, and the type-locality ("Huanusco" = Huatusco) is within the range of that species. Firschein (1954) expresseddoubt thatverruculatuswas aSyrrhophus, because Peters placed it in another genus. However, Peters describedverruculatusa decade before Cope diagnosed the genus Syrrhophus. Most frogs now calledSyrrhophus, plus a number of lower Central American frogs now placed in a variety of genera were placed inPhyllobatesby Boulenger, Cope, and Peters.The types ofPhyllobates verruculatuswere destroyed during World War II (Günther Peters,in litt.); the specimens subsequently assigned to the taxon by Kellogg (1932) areSyrrhophus cystignathoides. Because the type specimens are lost and because the name antedates the more established name,cystignathoides, I favor retainingPhyllobates verruculatusPeters as anomen dubium.Smith and Taylor (1948) reportedS. verruculatusfrom Tianguistengo, Hidalgo, México. These specimens are examples ofverrucipes. Smith (1947) reported a specimen ofverruculatusfrom San Lorenzo, Veracruz. Firschein (1954) referred it tocystignathoides, and Duellman (1960) concluded that both authors were in error and that the specimen (USNM 123530) was aleprus.Etymology.—The trivial name is the diminutive ofCystignathus, 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 Potosí to Central Veracruz, México (Fig. 5).Specimens examined.—(130), MÉXICO,Puebla: Necaxa, UMMZ 69519-20.San Luis Potosí: 5 km. W Aguismón, LSUMZ 4962-63; along Río 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 Córdoba, FMNH 104588, UIMNH 13321; Cuautlapam, 1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortín de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortín de las Flores, UIMNH 42799-808, UMMZ 105389; 3.2 km. N Fortín de las Flores, UIMNH 26633-35; 4.8 km. N Fortín de las Flores, UIMNH 71967-68; 3.2 km. W Fortín 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 Passa, UIMNH 49293, 49297; 1 km. E Plan del Río, 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 Tezuitlán (Puebla), UMMZ 105388; Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, 70441-42; 9.6 km. NW Tihuatlán, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU 23966; 26 km. NW Tuxpan, UMMZ 126419.Syrrhophus leprusCopeSyrrhophus leprusCope, 1879:268-69 [Holotype.—USNM 10040, from Santa Efigena, Oaxaca, México, 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: Günther, 1900:217.Syrrhophus leprus leprus: Neill, 1965:85-86.Syrrhophus leprus cholorumNeill, 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].[Pg 19]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 subacuminate; 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.Fig. 6:Dorsal views ofSyrrhophus leprusshowing variation in dorsal pattern (left, UMMZ 121244, ×2; right, KU 26106, ×1.7). Side of head (UIMNH 42726, ×7).Fig. 7:Distribution of three species of eastern complexSyrrhophus:leprus(circles),rubrimaculatus(triangles), andverrucipes(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, México.Duellman (1958, 1960) regardedS. leprusas 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, México, 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: Chinajá, KU 55961-62.El Petén: 15 km. NW Chinajá, KU 55963; Piedras Negras, USNM 114085-92; Tikal, UMMZ 117035; Uaxactún, AMNH 55121-22.MÉXICO,Oaxaca: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La Gloria, USNM 114093; 30.5 km. N Matías 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; Fortín 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 Andrés 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, 1.6km. S Catemaco, UMMZ 118222 (2); Volcán San Martín, south slope, UMMZ 118223; Volcán San Martín, Rancho El Tular, UIMNH 35399-400, 40340-41.Syrrhophus rubrimaculatusTaylor and SmithSyrrhophus rubrimaculatusTaylor and Smith, 1945:583-85 [Holotype.—USNM 114070, from La Esperanza, near Escuintla, Chiapas, México, 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.Fig. 8:Syrrhophus rubrimaculatus(upper right, KU 58911, ×1.6; lower right, KU 58910, ×4) andS. verrucipes(upper left, UIMNH 15995, ×1.6; lower left, UIMNH 15989, ×3.7).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 ofeye in both sexes; dorsum brown with small pale spots (red in life); venter gray.Remarks.—Previous authors who treatedSyrrhophusplaced this species in the western complex, because it occurs on the Pacific versant and has a reduced outer palmar tubercle. Duellman (1958) placedrubrimaculatusapart from the other western species, because of its relatively unexpanded digital tips and coloration. The digital tips are like those inleprus, whichrubrimaculatusresembles. Except for the reduction of the outer palmar tubercle,rubrimaculatuscould be a member of theleprusgroup.Syrrhophus rubrimaculatusis probably best treated as a Pacific derivative of theleprusgroup, even though the palmar tubercles do not agree. The removal ofrubrimaculatusfrom the western complex results in a more homogeneous 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, México (Fig. 7); probably extending into adjacent Guatemala.Specimens examined.—(48) MÉXICO,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 Jerónimo, 600-650 m., UIMNH 55299-312, 55313-16 (cleared and stained).Syrrhophus guttilatus(Cope)Malachylodes guttilatusCope, 1879:264 [Holotype.—USNM 9888, from Guanajuato, Guanajuato, México; collected in 1877 by Alfredo Duges].Syrrhopus guttulatus: Boulenger, 1888:204-06.Syrrhaphus guttulatus: Günther, 1900:317.Syrraphus guttulatus: Díaz de León, 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 smithiTaylor, 1940b:43-45, pl. 1 [Holotype.—USNM 108594, from 15 mi. SW Galeana, Nuevo León, México, 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 gaigeaeSchmidt 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 petrophilusFirschein, 1954:50-52 [Holotype.—UIMNH 7807, from 5 km. SW San Luis Potosí, San Luis Potosí, México; 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).Fig. 9:Syrrhophus guttilatus(upper left, UIMNH 55519, ×1.4; lower left, UIMNH 55519, ×2.3) andS. marnockii(upper right, TCWC 9317, ×1.4; lower right, TCWC 13510, ×2.1).Remarks.—Cope (1879) distinguishedMalachylodesfromSyrrhophuson the basis of the presence of a frontoparietal fontanelle in the holotype ofguttilatus. 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 distinguishguttilatusfrom hispetrophilus.As is clearly evident from the length of the synonymy, I consider a number of currently used names to be synonymous withguttilatus. I have seen the holotypes of all four names and am unable to recognize more than a single species. The holotype ofpetrophilusis a male, whereas that ofsmithiis 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 ofgaigeaeandMalachylodes guttilatusagree in color pattern.Schmidt and Smith (1944) namedSyrrhophus gaigeaefrom the Chisos Mountains of the Big Bend region of Texas and compared it only withS. marnockii. Milstead, Mecham and McClintock (1950) synonymizedgaigeaeandmarnockiibecause 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 vermiculatepattern, 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.Fig. 10:Distribution ofSyrrhophus guttilatus.Nearly every specimen examined was infested with chiggers of the genusHannemania. 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 ofSyrrhophus marnockiiand a fewS. verrucipes, but on no other species of the genus. Mr. Willy Wrenn told me that he has seen heavy infestations ofHannemaniaonSyrrhophus pallidus. Infestation byHannemaniaprobably 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 ofS. 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 ofMalachulodes 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 ofSyrrhophus smithi).San Luis Potosí: 5 km. SW San Luis Potosí, UIMNH 7807 (holotype ofS. petrophilus).Tamaulipas: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4).Syrrhophus marnockiiCopeSyrrhophus marnockiiCope, 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 ofS. 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 verificationby 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).Fig. 11:Distribution ofSyrrhophus marnockii(circles). Starred localities are late Pleistocene records.Syrrhophus verrucipesCopeSyrrhophus verrucipesCope, 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 macrotympanumTaylor, 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 aTomodactylus, but Dixon (1957) pointed out thatT. macrotympanumdiffered from the other species of the genus in having a poorly developed lumbo-inguinal (inguinal) gland, and placed the species in the genusSyrrhophus. Comparison of the holotypes ofS. verrucipesandT. macrotympanumleaves no doubt in my mind that a single species is involved. This same species was reported by Smith and Taylor (1948) asS. verruculatus.Syrrhophus verrucipesbearsresemblanceto members of both theleprusandmarnockiigroups. In snout shape it is closer to theleprusgroup, whereas in digital pad, the shape of the general body form, and contiguity of habitat it is most similar to themarnockiigroup (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 ofTomodactylus 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 ofSyrrhophus verrucipes).Queretaro: 3.5 km. S San Juan del Río, EAL 1343.San Luis Potosí: 9.6 km. W Ahuacatlán, LSUMZ 4968-70.Syrrhophus dennisinew speciesSyrrhophus 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 inTable 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 inS. longipes(=S. latodactylusof Taylor and Martin).Fig. 12:Syrrhophus dennisisp. 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 combinationBatrachyla longipesBaird, 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 latodactylusTaylor, 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'sBatrachyla longipesto the frog Taylor (1940d) calledSyrrhophus latodactylusbecause the color pattern (Fig. 13) predominant in the southern part of the range agrees with that described (figured) forBatrachyla 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 nameBatrachyla longipesis currently that of anomen dubium(Lynch, 1963). At that time, I was unaware of the geographic variation in color pattern inSyrrhophus latodactylus.The exact type-locality ofBatrachyla longipesis 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 placeBatrachyla longipesinEleutherodactylusinstead ofSyrrhophus. Barbour (1923) and Kellogg (1932) associated another species (E. batrachylus) withlongipes. Taylor (1940a) noted this as a case of misidentification and corrected the error but leftlongipesin the genusEleutherodactylus. Lynch (1963) noted several points of morphological agreement betweenSyrrhophusandB. longipesbut did not placelongipesinSyrrhophus.Baird's (1859) figures of the holotype do not illustrate prevomerine teeth, but according to Cope (1866) they were present in the holotype. The digitaltips of the frog in the figure are somewhat narrower than those typically seen inS. latodactylus. If the specimen was slightly desiccated, as possibly was the case, the digits would appear narrower. There is no evidence contrary to placingSyrrhophus latodactylusin the synonymy ofBatrachyla longipes.Fig. 13:Dorsal views ofSyrrhophus longipesillustrating geographic variation in pattern (left, TCWC 12179, ×1.5; right, KU 92572, ×1.8); side of head (TCWC 10966, ×6).Application of Baird's nameBatrachyla longipesto the species of frog heretofore calledSyrrhophus latodactylusposes one serious problem.Batrachylalongipesis the type-species (by original designation) of the genusEpirhexisCope, 1866, which has priority overSyrrhophusCope, 1878. IfBatrachyla longipesis left in the status of anomen dubium,Epirhexiscan be forgotten, for the two names are tied together. However, since it seems almost certain thatBatrachyla longipesandSyrrhophus latodactylusare conspecific, the former name should not be left as anomen dubium.Epirhexisnever came into general usage (Cope cited the name four times, but no one else has used it), whereasSyrrhophusis well established in the zoological literature. It would serve only to confuse the literature to adhere strictly to the Law of Priority and replaceSyrrhophuswithEpirhexis. Therefore,Syrrhophusis used in this paper, even thoughEpirhexishas priority. A request for the suppression ofEpirhexisCope, 1866, has been submitted to the International Commission of Zoological Nomenclature (Lynch, 1967).Etymology.—Latin, meaning long-footed; Taylor'slatodactylusrefers to the wide digital pads.Fig. 14:Distribution ofSyrrhophus dennisi(triangle) andS. longipes(circles).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 ofS. 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 cystignathoides campiStejneger, New combination

Syrrhophus campiStejneger, 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 thatS. campiwas probably a subspecies of the more southernS. cystignathoides. Various references in the literature might lead one to believe that the two were sympatric over much of northeastern México; 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 Potosí, 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 Río Grande embayment in Texas to central Nuevo León and Tamaulipas, México. Intergrades are known from southern Tamaulipas and adjacent San Luis Potosí, México (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-Río Grande State Park, UMMZ 114378; 6 mi. S McAllen, TNHC 7136-39; Santa Ana Refuge, TCWC 13495-96; Weslaco, TCWC 17658-60.

MEXICO,Nuevo León: 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).

Fig. 4:Syrrhophus cystignathoides campi(left, TCWC 13490) andS. c. cystignathoides(right, KU 105500). Dorsal views ×2, sides of heads ×3.

Intergrades [S. c. cystignathoides×S. c. campi(88)] MÉXICO,San Luis Potosí: 5 km. ECiudaddel Maiz, UMMZ 106435; 16 km. W Naranjo, FMNH 104584; Salto de Agua, 34 km. WSW Antigua Morelos, TCWC 6980.Tamaulipas: 5 km. W Acuña, 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 Gómez Farías, 450 m., UMMZ 101165; 8 km. NE Gómez Farías, Pano Ayuctle, UMMZ 102264, 102924 (6); 8 km. NW Gómez Farías, 1060 m., LSUMZ 11084, UMMZ 101199, 102928 (5), 102929-32, 110124 (3); Río 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.

Syrrhophus cystignathoides cystignathoides(Cope), New combination

Phyllobates cystignathoidesCope, 1877:89-90 [Syntypes.—USNM 32402-32409, from Potrero, near Córdoba, Veracruz, México, 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: Günther, 1900:218.

Syrraphus cystignathoides: Díaz de León, 1904:10.

Syrrhopus cystignathoides: Barbour and Loveridge, 1946-170[** 1946:170].

Fig. 5:Distribution ofSyrrhophus 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 verruculatusand noted that if it was aSyrrhophusit would probably be referrable toS. cystignathoides. Peters' (1871) original description corresponds well withS. cystignathoides, and the type-locality ("Huanusco" = Huatusco) is within the range of that species. Firschein (1954) expresseddoubt thatverruculatuswas aSyrrhophus, because Peters placed it in another genus. However, Peters describedverruculatusa decade before Cope diagnosed the genus Syrrhophus. Most frogs now calledSyrrhophus, plus a number of lower Central American frogs now placed in a variety of genera were placed inPhyllobatesby Boulenger, Cope, and Peters.

The types ofPhyllobates verruculatuswere destroyed during World War II (Günther Peters,in litt.); the specimens subsequently assigned to the taxon by Kellogg (1932) areSyrrhophus cystignathoides. Because the type specimens are lost and because the name antedates the more established name,cystignathoides, I favor retainingPhyllobates verruculatusPeters as anomen dubium.

Smith and Taylor (1948) reportedS. verruculatusfrom Tianguistengo, Hidalgo, México. These specimens are examples ofverrucipes. Smith (1947) reported a specimen ofverruculatusfrom San Lorenzo, Veracruz. Firschein (1954) referred it tocystignathoides, and Duellman (1960) concluded that both authors were in error and that the specimen (USNM 123530) was aleprus.

Etymology.—The trivial name is the diminutive ofCystignathus, 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 Potosí to Central Veracruz, México (Fig. 5).

Specimens examined.—(130), MÉXICO,Puebla: Necaxa, UMMZ 69519-20.San Luis Potosí: 5 km. W Aguismón, LSUMZ 4962-63; along Río 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 Córdoba, FMNH 104588, UIMNH 13321; Cuautlapam, 1000 m., FMNH 106477-80, KU 100364, UIMNH 58200-03, UMMZ 105392; Fortín de las Flores, UIMNH 13322, 13339; 1.6 km. N Fortín de las Flores, UIMNH 42799-808, UMMZ 105389; 3.2 km. N Fortín de las Flores, UIMNH 26633-35; 4.8 km. N Fortín de las Flores, UIMNH 71967-68; 3.2 km. W Fortín 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 Passa, UIMNH 49293, 49297; 1 km. E Plan del Río, 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 Tezuitlán (Puebla), UMMZ 105388; Teocelo, FMNH 70437-38, KU 26080, 26790; 3.2 km. N Teocelo, FMNH 70439, 70441-42; 9.6 km. NW Tihuatlán, UIMNH 3684-85; 15 km. ENE Tlacotepec, KU 23966; 26 km. NW Tuxpan, UMMZ 126419.

Syrrhophus leprusCope

Syrrhophus leprusCope, 1879:268-69 [Holotype.—USNM 10040, from Santa Efigena, Oaxaca, México, 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: Günther, 1900:217.

Syrrhophus leprus leprus: Neill, 1965:85-86.

Syrrhophus leprus cholorumNeill, 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].

[Pg 19]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 subacuminate; 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.

Fig. 6:Dorsal views ofSyrrhophus leprusshowing variation in dorsal pattern (left, UMMZ 121244, ×2; right, KU 26106, ×1.7). Side of head (UIMNH 42726, ×7).

Fig. 7:Distribution of three species of eastern complexSyrrhophus:leprus(circles),rubrimaculatus(triangles), andverrucipes(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, México.

Duellman (1958, 1960) regardedS. leprusas 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, México, 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: Chinajá, KU 55961-62.El Petén: 15 km. NW Chinajá, KU 55963; Piedras Negras, USNM 114085-92; Tikal, UMMZ 117035; Uaxactún, AMNH 55121-22.

MÉXICO,Oaxaca: Cerro San Pedro del Isthmo, UIMNH 35510; Finca La Gloria, USNM 114093; 30.5 km. N Matías 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; Fortín 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 Andrés 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, 1.6km. S Catemaco, UMMZ 118222 (2); Volcán San Martín, south slope, UMMZ 118223; Volcán San Martín, Rancho El Tular, UIMNH 35399-400, 40340-41.

Syrrhophus rubrimaculatusTaylor and Smith

Syrrhophus rubrimaculatusTaylor and Smith, 1945:583-85 [Holotype.—USNM 114070, from La Esperanza, near Escuintla, Chiapas, México, 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.

Fig. 8:Syrrhophus rubrimaculatus(upper right, KU 58911, ×1.6; lower right, KU 58910, ×4) andS. verrucipes(upper left, UIMNH 15995, ×1.6; lower left, UIMNH 15989, ×3.7).

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 ofeye in both sexes; dorsum brown with small pale spots (red in life); venter gray.

Remarks.—Previous authors who treatedSyrrhophusplaced this species in the western complex, because it occurs on the Pacific versant and has a reduced outer palmar tubercle. Duellman (1958) placedrubrimaculatusapart from the other western species, because of its relatively unexpanded digital tips and coloration. The digital tips are like those inleprus, whichrubrimaculatusresembles. Except for the reduction of the outer palmar tubercle,rubrimaculatuscould be a member of theleprusgroup.

Syrrhophus rubrimaculatusis probably best treated as a Pacific derivative of theleprusgroup, even though the palmar tubercles do not agree. The removal ofrubrimaculatusfrom the western complex results in a more homogeneous 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, México (Fig. 7); probably extending into adjacent Guatemala.

Specimens examined.—(48) MÉXICO,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 Jerónimo, 600-650 m., UIMNH 55299-312, 55313-16 (cleared and stained).

Syrrhophus guttilatus(Cope)

Malachylodes guttilatusCope, 1879:264 [Holotype.—USNM 9888, from Guanajuato, Guanajuato, México; collected in 1877 by Alfredo Duges].

Syrrhopus guttulatus: Boulenger, 1888:204-06.

Syrrhaphus guttulatus: Günther, 1900:317.

Syrraphus guttulatus: Díaz de León, 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 smithiTaylor, 1940b:43-45, pl. 1 [Holotype.—USNM 108594, from 15 mi. SW Galeana, Nuevo León, México, 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 gaigeaeSchmidt 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 petrophilusFirschein, 1954:50-52 [Holotype.—UIMNH 7807, from 5 km. SW San Luis Potosí, San Luis Potosí, México; 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).

Fig. 9:Syrrhophus guttilatus(upper left, UIMNH 55519, ×1.4; lower left, UIMNH 55519, ×2.3) andS. marnockii(upper right, TCWC 9317, ×1.4; lower right, TCWC 13510, ×2.1).

Remarks.—Cope (1879) distinguishedMalachylodesfromSyrrhophuson the basis of the presence of a frontoparietal fontanelle in the holotype ofguttilatus. 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 distinguishguttilatusfrom hispetrophilus.

As is clearly evident from the length of the synonymy, I consider a number of currently used names to be synonymous withguttilatus. I have seen the holotypes of all four names and am unable to recognize more than a single species. The holotype ofpetrophilusis a male, whereas that ofsmithiis 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 ofgaigeaeandMalachylodes guttilatusagree in color pattern.

Schmidt and Smith (1944) namedSyrrhophus gaigeaefrom the Chisos Mountains of the Big Bend region of Texas and compared it only withS. marnockii. Milstead, Mecham and McClintock (1950) synonymizedgaigeaeandmarnockiibecause 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 vermiculatepattern, 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.

Fig. 10:Distribution ofSyrrhophus guttilatus.

Nearly every specimen examined was infested with chiggers of the genusHannemania. 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 ofSyrrhophus marnockiiand a fewS. verrucipes, but on no other species of the genus. Mr. Willy Wrenn told me that he has seen heavy infestations ofHannemaniaonSyrrhophus pallidus. Infestation byHannemaniaprobably 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 ofS. 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 ofMalachulodes 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 ofSyrrhophus smithi).San Luis Potosí: 5 km. SW San Luis Potosí, UIMNH 7807 (holotype ofS. petrophilus).Tamaulipas: 1.6 km. NW La Joya de Salas, 1530 m., UMMZ 110736 (4).

Syrrhophus marnockiiCope

Syrrhophus marnockiiCope, 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 ofS. 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 verificationby 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).

Fig. 11:Distribution ofSyrrhophus marnockii(circles). Starred localities are late Pleistocene records.

Syrrhophus verrucipesCope

Syrrhophus verrucipesCope, 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 macrotympanumTaylor, 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 aTomodactylus, but Dixon (1957) pointed out thatT. macrotympanumdiffered from the other species of the genus in having a poorly developed lumbo-inguinal (inguinal) gland, and placed the species in the genusSyrrhophus. Comparison of the holotypes ofS. verrucipesandT. macrotympanumleaves no doubt in my mind that a single species is involved. This same species was reported by Smith and Taylor (1948) asS. verruculatus.

Syrrhophus verrucipesbearsresemblanceto members of both theleprusandmarnockiigroups. In snout shape it is closer to theleprusgroup, whereas in digital pad, the shape of the general body form, and contiguity of habitat it is most similar to themarnockiigroup (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 ofTomodactylus 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 ofSyrrhophus verrucipes).Queretaro: 3.5 km. S San Juan del Río, EAL 1343.San Luis Potosí: 9.6 km. W Ahuacatlán, LSUMZ 4968-70.

Syrrhophus dennisinew 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 inTable 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 inS. longipes(=S. latodactylusof Taylor and Martin).

Fig. 12:Syrrhophus dennisisp. 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 longipesBaird, 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 latodactylusTaylor, 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'sBatrachyla longipesto the frog Taylor (1940d) calledSyrrhophus latodactylusbecause the color pattern (Fig. 13) predominant in the southern part of the range agrees with that described (figured) forBatrachyla 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 nameBatrachyla longipesis currently that of anomen dubium(Lynch, 1963). At that time, I was unaware of the geographic variation in color pattern inSyrrhophus latodactylus.

The exact type-locality ofBatrachyla longipesis 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 placeBatrachyla longipesinEleutherodactylusinstead ofSyrrhophus. Barbour (1923) and Kellogg (1932) associated another species (E. batrachylus) withlongipes. Taylor (1940a) noted this as a case of misidentification and corrected the error but leftlongipesin the genusEleutherodactylus. Lynch (1963) noted several points of morphological agreement betweenSyrrhophusandB. longipesbut did not placelongipesinSyrrhophus.

Baird's (1859) figures of the holotype do not illustrate prevomerine teeth, but according to Cope (1866) they were present in the holotype. The digitaltips of the frog in the figure are somewhat narrower than those typically seen inS. latodactylus. If the specimen was slightly desiccated, as possibly was the case, the digits would appear narrower. There is no evidence contrary to placingSyrrhophus latodactylusin the synonymy ofBatrachyla longipes.

Fig. 13:Dorsal views ofSyrrhophus longipesillustrating geographic variation in pattern (left, TCWC 12179, ×1.5; right, KU 92572, ×1.8); side of head (TCWC 10966, ×6).

Application of Baird's nameBatrachyla longipesto the species of frog heretofore calledSyrrhophus latodactylusposes one serious problem.Batrachylalongipesis the type-species (by original designation) of the genusEpirhexisCope, 1866, which has priority overSyrrhophusCope, 1878. IfBatrachyla longipesis left in the status of anomen dubium,Epirhexiscan be forgotten, for the two names are tied together. However, since it seems almost certain thatBatrachyla longipesandSyrrhophus latodactylusare conspecific, the former name should not be left as anomen dubium.Epirhexisnever came into general usage (Cope cited the name four times, but no one else has used it), whereasSyrrhophusis well established in the zoological literature. It would serve only to confuse the literature to adhere strictly to the Law of Priority and replaceSyrrhophuswithEpirhexis. Therefore,Syrrhophusis used in this paper, even thoughEpirhexishas priority. A request for the suppression ofEpirhexisCope, 1866, has been submitted to the International Commission of Zoological Nomenclature (Lynch, 1967).

Etymology.—Latin, meaning long-footed; Taylor'slatodactylusrefers to the wide digital pads.

Fig. 14:Distribution ofSyrrhophus dennisi(triangle) andS. longipes(circles).

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 ofS. 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.

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