Chapter 4

Syrrhophus pipilansTaylorSyrrhophus pipilansTaylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 (formerly EHT-HMS 6843), 14.6 km. S Mazatlán, Guerrero, México; collected on July 22, 1936, by Edward H. Taylor].Diagnosis.—Medium sized frogs, males 22.6-28.5 mm. snout-vent, females 21.1-29.4 mm. snout-vent length; vocal slits present in males; finger tips slightly expanded, truncate in outline; inner metatarsal tubercle less than twice the size of outer; skin of dorsum smooth to shagreened, that of venter smooth; tympanum 36.5-54.0 per cent diameter of eye; dorsum dark brown with large or small light brown, orange-brown, or yellowish spots or blotches; limbs banded; interorbital bar absent.Fig. 15:Dicegrams of ear size relative to eye diameter in the two subspecies ofSyrrhophus pipilans. N = 17 innebulosus, 18 inpipilans.Remarks.—Two subspecies were recognized by Duellman (1958). Previously both had been treated as species. The two populations were distinguished on the basis of color pattern and the size of the tympanum. Measurements of 17 males ofS. p. nebulosusfrom central Chiapas and 18 males ofS. p. pipilansfrom south-central Oaxaca and Guerrero, México, demonstrates that the supposed difference in tympanum size is not significant (Fig. 15).There is, however, a tendency for the western population ofS. pipilansto have larger tympani. Based on the present examination of 112 specimens of this species the two populations are held to be sufficiently distinct to warrant taxonomic recognition as subspecies (Fig. 16).Fig. 16:Syrrhophus pipilans nebulosus(left, KU 58908) andS. p. pipilans(right, KU 86885). ×2.7.The parotoid glands attributed to this species by Taylor (1940c:95) are merely the superficial expression of them. depressor mandibulaeand scapula. No true glands are present in the parotoid region.Syrrhophus pipilans nebulosusTaylorSyrrhophus nebulosusTaylor, 1943:353-55, pl. 27, figs. 3-5 [Holotype.—FMNH 100095 (formerly EHT-HMS 3774), near Tonolá, Chiapas, México; collected on August 27, 1935, by Hobart M. Smith and Edward H. Taylor]. Smith and Taylor, 1948:49, 51.Syrrhophus pipilans nebulosus: Duellman, 1958:2-4, 9, 12, 14. Stuart, 1963:32-33. Gorham, 1966:166-67.Diagnosis.—Diameter of tympanum 36.6-47.8 per cent that of eye; dorsum dark brown with numerous small light brown to yellowish spots.Remarks.—The distribution of this subspecies is adequately described by Duellman (1958). Fouquette (1960) described the vocalization of this frog.Etymology.—Latin,nebula, in reference to the clouded dorsal pattern.Distribution.—Low to moderate elevations along the Pacific versant of Chiapas and in the Grijalva valley of Chiapas and Guatemala (Fig. 17).Specimens examined.—(54) GUATEMALA,Huehuetenango: Jacaltenango, UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MÉXICO,Chiapas: 11.2 km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279;12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; 1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca. 2 km. NW Pueblo Nuevo Solistahuacán, KU 58897; Hda. Monserrate, 40 km. NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH 75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, TNHC 26942, Tonolá, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); 19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, KU 37809; Unión de Juarez, FMNH 105294.Syrrhophus pipilans pipilansTaylor?Syrrhopus verruculatus: Gadow, 1905:194.Syrrhophus pipilansTaylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 (formerly EHT-HMS 6843), from 14.6 km. S Mazatlán, Guerrero, México; collected on July 22, 1936, by Edward H. Taylor]. Taylor and Smith, 1945:581-82. Smith and Taylor, 1948:49, 50-51.Syrrhophus pipilans pipilans: Duellman, 1958:1-4, 8-9, 13-14, pl. 2, fig. 1. Gorham, 1966:166.Diagnosis.—Diameter of tympanum 40.6-54.0 per cent that of eye; dorsum dark brown with large light spots or blotches.Remarks.—Duellman's (1958) synopsis of this subspecies is adequate; the distribution has not been extended, but several records are now available which fill in gaps.Fig. 17:Distribution ofSyrrhophus pipilans:nebulosus(open circles) andpipilans(solid circles).Gadow's (1905) record ofS. verruculatusfrom "Buena Vista, S. Guerrero" is most likely applicable to this species. Gadow simply included the name in a list of the species he had collected during his trip in México (1902-04); nofurther comment was made on this species although references toSyrrhopus(sic) appear in several places in the paper and would appear to apply to the species he had.Etymology.—Latin,pipilo, chirping, peeping, in reference to the call of the male.Distribution.—Sea level to about 1800 meters along the Pacific versant of western México from central Guerrero to the Isthmus of Tehuantepec (Fig. 17).Specimens examined.—(62). MÉXICO,Guerrero: Acapulco, UMMZ 110125; 6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, 125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S Mazatlán, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH 126639; Xaltinanguis, FMNH 100522-24, 126640.Oaxaca: Cacahuatepec, UIMNH 52853; 8 km. NW Río Canoa, 53 km. ESE Cuajinicuilapa, UIMNH 52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; 37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 (2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ 115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2).Syrrhophus interorbitalisLangebartel and ShannonSyrrhophus interorbitalisLangebartel and Shannon, 1956: 161-65, figs. 1-2 [Holotype.—UIMNH 67061 (formerly FAS 9378), 36 mi. N Mazatlán, Sinaloa, México, collected on November 17, 1955, by E. C. Bay, J. C. Schaffner, and D. A. Langebartel]. Duellman, 1958:1-4, 10, 12, 14. Gorham, 1966:164-65.Syrrhophis interorbitalis: Campbell and Simmons, 1962:194, fig. 1.Fig. 18:Left to right.Syrrhophus interorbitalis(UIMNH 38095, ×1.5),S. nivocolimae(LACM 3203, ×1.3), andS. teretistes(KU 75263, ×1.5).Diagnosis.—Medium sized frogs, only known male 25.6 mm. snout-vent, females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in males;finger tips expanded; first finger shorter than second; outer metatarsal tubercle one-third size of inner; skin of dorsum shagreened, that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of eye in both sexes; pale yellow-brown ground color mottled with brown; limb bands broad, much wider than narrow light interspaces; interorbital bar very long, edged with dark brown to black (Fig. 18).Remarks.—Duellman's (1958) measurements and proportions ofS. interorbitaliswere based exclusively on the type series, which is composed of only females; therefore hisinterorbitalisdata are not comparable with the data for the other species in his table. Campbell and Simmons (1962) collected the only known male. The type series was collected beneath rocks in a stream bed; the collectors heard calling frogs in the bushes but were unable to obtain specimens (Langebartel and Shannon, 1956). Campbell and Simmons (1962) reported that their specimen had a poorly developed interorbital bar in life; in preservative the bar compares favorably with the bar in the female (Fig. 18).Etymology.—Latin, in reference to the pale interocular band.Distribution.—Pacific lowlands of Sinaloa, México (Fig. 20).Specimens examined.—(10). MÉXICO,Sinaloa: 36 mi. N Mazatlán, UIMNH 38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlán, LACM 13773.Syrrhophus modestusTaylorSyrrhophus modestusTaylor, 1942:304-06, pl. 29 [Holotype.—FMNH 100048 (formerly EHT-HMS 3756), from Hacienda Paso del Río, Colima, México; collected on July 8, 1935, by Hobart M. Smith]. Smith and Taylor, 1948:49-50.Syrrhophus modestus modestus: Duellman, 1958:2-5, 7, 14, pl. 1, fig. 1. Gorham, 1966:166.Diagnosis.—Small frogs, males 15.8-20.1 mm. snout-vent length, single female 18.5 mm.; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed; pale cream in preservative with dark brown spots; limbs banded; bands on forearm and thigh poorly developed or absent; interorbital bar absent.Remarks.—The tympanum is concealed inS. modestus,S. nivocolimae,S. pallidus,S. teretistes, and to a lesser degree inS. interorbitalis. However, if the specimen is permitted to dry slightly, the annulus tympanicus becomes visible through the skin and a tympanum/eye ratio can be computed.One of the few cases of sympatry within the genusSyrrhophusinvolves this species;modestusandnivocolimaeare known to be sympatric at one locality in southwestern Jalisco, México.Duellman (1958) used the trinomial for this population and named a new subspecies,pallidus, from Nayarit. I considerpallidusto be specifically distinct frommodestusbecause there is no evidence of genetic exchange, and there is no overlap in the distinguishing morphological features. I do consider the two populations to be closely related but feel the inter-relationships betweenmodestus,pallidus,nivocolimae, andteretistesare more complex than would be indicated by the use of trinomials. The sympatric occurrence ofmodestusandnivocolimaeis significant; morphologically, they might otherwise be regarded as subspecies. Although allopatric, similar arguments could be advanced forthe morphologically similarpallidusandteretistes. The four are here afforded species rank since morphological similarity and allopatry are not sufficient grounds for the assumption of genetic exchange.Fig. 19:Syrrhophus modestus[left, UMMZ 115447 (WED 11155)] andS. pallidus(right, UMMZ 115453). ×2.2.Etymology.—Latin, meaning unassuming, modest, in reference to the small size of the species.Distribution.—Low elevations (up to 700 meters) in the lowlands and foothills of Colima and southwestern Jalisco, México (Fig. 20).Specimens examined.—(14). MÉXICO,Colima: Hda. Paso del Río, FMNH 100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM 139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4);Jalisco: 17.6 km. SW Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahía Tenacatita, UMMZ 84264.Syrrhophus nivocolimaeDixon and WebbSyrrhophus nivocolimaeDixon and Webb, 1966:1-4, Fig. 1 [Holotype.—LACM 3200, from Nevado de Colima (6 airline miles west of Atenquique), Jalisco, México, 7800 feet; collected on July 20, 1964, by Robert G. Webb].Diagnosis.—Small frogs, males 18.5-21.1 mm. snout-vent length, only known female 24.1 mm. snout-vent; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum warty, that of venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that of eye in males; mid-dorsal brown band from interorbital bar to anus; bands on limbs narrow, dark bandsless than one-half width of light bands, upper arm not banded; narrow interorbital light bar.Remarks.—This species is closely related toS. modestusand differs in color pattern and degree of wartiness of the skin. Dixon and Webb (1966) held thatnivocolimaehad no close relatives, but the condition of the tympanum, size, nature of the outer palmar tubercle, relative sizes of the metatarsal tubercles, and shape and size of the digital pads all point to a close relationship betweenS. modestus,S. nivocolimae, andS. pallidus.Fig. 20:Distribution of the species of themodestusgroup:interorbitalis(open circles),teretistes(solid circles),modestus(open triangles),pallidus(solid triangles) andnivocolimae(square). Arrow indicates locality of sympatry betweenmodestusandnivocolimae. Solid line about the localities forinterorbitalisis a range estimate based on call records and specimens examined.Dixon and Webb (1966) reported thatS. nivocolimaehas a large tympanum (50.0-59.0 per cent diameter of eye). However, my examination of the type series and several other specimens from Jalisco reveals that thelargest tympanum/eye ratio is 39.3 per cent. Therefore, the tympanum/eye ratio inS. nivocolimaeis in agreement with those forS. modestus,S. pallidus, andS. teretistes(Table 6).Etymology.—niv, Latin, and Colima (Nevado de), meaning high on the volcano, in reference to the higher distribution of this species (around 2000 meters) than other members of the group.Distribution.—Known from southwestern Jalisco, México, at moderate to high elevations (600-2400 meters).Specimens examined.—(48) MÉXICO,Jalisco: 17.6 km. SW Autlán, 606 m., KU 102626, 102631; 6.4 km. W Atenquique, 2060 m., KU 102628-30, 102632; 8 km. W Atenquique, 1970 m., LACM 3210-12; 9.6 km. W Atenquique, 2360 m., LACM 3200 (holotype), 3201-09; 14.5 km. W Atenquique, 2000 m., LACM 25424-36, 25439-41, 25446; 15 km. W Atenquique, LACM 37044-46, 37244-47; 16 km. W Atenquique, 2105 m., LACM 25443-45; 17 km. W Atenquique, 2180 m., LACM 25442.Syrrhophus pallidusDuellman, New combinationSyrrhophus modestus: Davis and Dixon, 1957:146.Syrrhophus modestus pallidusDuellman, 1958:2-3, 5-7, 14, pl. 3 [Holotype.—UMMZ 115452, from San Blas, Nayarit, México, sea level; collected on August 13, 1956, by William E. and Ann S. Duellman]. Zweifel, 1960:86-88, 91, 93-94, 118, 120-22. Gorham, 1966:166.Syrrhophis modestus pallidus: Campbell and Simmons, 1962:194.Diagnosis.—Small frogs, males 17.9-19.3 mm. snout-vent length; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed, its diameter 27.0-35.6 per cent of eye in males; ground color cream vermiculated with brown, upper arm and thigh lacking, or with few, indistinct, bands; interorbital bar absent.Remarks.—Considerable debate has been waged relative to the value of subspecies and to the reasons for recognizing distinct disjunct populations as species versus subspecies. Lacking evidence of genetic exchange, I prefer to retain disjunct populations that are distinctive as species.All known specimens ofpalliduscan be separated from those ofmodestusby color pattern. The two nominal species exhibit overlap in proportions but the same can be said about nearly every species ofSyrrhophus; therefore, overlap in proportions can be disregarded in assessing specific versus subspecific rank. Until contrary evidence is forthcoming, I consider the disjunct populations heretofore held to be subspecies ofmodestusto be specifically distinct. The specimens of the disjunct population ofpalliduson the Tres Marias do not differ from the mainland population in Nayarit. This evidence, though perhaps secondary, supports my contention that two species should be recognized.Etymology.—Latin, in reference to the pale ground color in comparison with that ofS. modestus.Distribution.—Low elevations in coastal Nayarit and on Islas Tres Marias (Fig. 20).Specimens examined.—(12) MÉXICO,Nayarit: 18.8 mi. NW Ahuacatlán, UIMNH 7808; San Blas, UMMZ 115452 (holotype), 115453-57; 17 km. NE San Blas, 150 m., MSU 5085; 12.8 km. E San Blas, UIMNH 71979; 31 km. E San Blas, UIMNH 71978; 13.5 km. N Tepic, UIMNH 71980-81.Syrrhophus teretistesDuellmanSyrrhophus teretistesDuellman, 1958:2-3, 10-14, pl. 2, fig. 2 [Holotype.—UMMZ 115451, from 4.8 km. NW Tepic, Nayarit, México, 840 m.; collected on August 12, 1956, by William E. Duellman]. Gorham, 1966:167.Diagnosis.—Medium-sized frogs, males 19.2-23.2 mm. snout-vent length, single known female 24.8 mm. snout-vent; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum partially concealed, its diameter 28.6-43.8 per cent of eye in males; ground color brown vermiculated with dark brown to nearly black; upper arm and thigh banded; interorbital light bar absent.Remarks.—S. teretistesappears to be most closely related toS. pallidus; I consider it to be an upland derivative ofpallidus. Morphologically, the differences between the two are few, but lacking evidence of genetic exchange they are retained as species.Etymology.—Greek, in reference to the whistle-like nature of the call.Distribution.—Moderate elevations (840-1200 meters) in the Sierra Occidental of Nayarit, Sinaloa, and Durango, México (Fig. 20).Specimens examined.—(13) MÉXICO,Nayarit: 4.8 km. NW Tepic, 840 m., UMMZ 115451 (holotype).Sinaloa: Santa Lucía, 1090 m., KU 75263-72; 1 km. NE Santa Lucía, 1156 m., KU 78257; 2.2 km. NE Santa Lucía, 1156 m., KU 78258.DiscussionThere are relatively few clear-cut morphological differences among the fourteen species now assigned toSyrrhophus. The majority of the species are allopatric and differ primarily in color patterns. Sympatric occurrence serves as an indicator of specific distinctness and is one of the more practical tests of species validity when cross-breeding experiments are not possible. Two cases of sympatric occurrence are known for the species ofSyrrhophusin western México:modestusandnivocolimaeare sympatric in southern Jalisco andpipilans nebulosusandrubrimaculatusare sympatric in southeastern Chiapas. In eastern México,longipesandverrucipesare sympatric in southern Hidalgo, andlongipesis sympatric withcystignathoides,dennisi, andguttilatusin southern Tamaulipas.Syrrhophus cystignathoidesandleprusare apparently sympatric in central Veracruz.Subspecific assignments have been made only when there is evidence of intergradation. The sympatric occurrence of morphologically similar species in this genus has led me to adopt a conservative approach to the degree of difference philosophy. I have therefore recognized all morphologically distinct allopatric populations as species.Fig. 21:Generic distributions ofSyrrhophus(stipple) andTomodactylus(hatching). Black areas are zones of intergeneric sympatry.Fig. 22:Altitudinal distributions ofSyrrhophusandTomodactylus. Widths of the columns are proportional to the numbers of species at a given altitude; narrowest width equals one species.Syrrhophusis closely allied to another Mexican leptodactylid genus,Tomodactylus, which was revised by Dixon (1957), who along with numerous other authors noted the close relationship between the two genera. There is an almost complete lack of sympatry between the two genera; in very few places in México do they coexist (Fig. 21).Tomodactylushas its greatest diversity in the Cordillera Volcánica and Sierra Madre del Sur, whereasSyrrhophusreaches its greatest diversity in the Sierra Madre Oriental and eastern foothills. The species of both genera are about the same size and presumably have similar requirements insofar as food, breeding sites, and habitat selection.Four cases of intergeneric sympatry are known for the two genera: 1) the Chilpancingo region of Guerrero, 2) the lowlands of Colima and the mountains just inland in Jalisco, 3) the lowlands of central Nayarit, and 4) the Sierra Madre Occidental on the Durango-Sinaloan border. The apparent sympatry in the Chilpancingo region involves four species:S. pipilans,T. albolabris,T. dilatus, andT. nitidus. Of the four,T. dilatusappears to be completely allopatric in that it occurs at higher altitudes (above 2000 meters), whereas the other three occur below 1800 meters in the region (Davis and Dixon, 1965). In the Colima-Jalisco region,Tomodactylustendsto occur higher (Dixon and Webb, 1966) than some of theSyrrhophus, but one subspecies ofTomodactylus nitidusis a lowland frog, occurring sympatrically with the lowlandSyrrhophus modestus. A similar situation is observed in Nayarit; the lowlandTomodactylusoccurs sympatrically with the smallSyrrhophus pallidus. In both cases theSyrrhophusis smaller than theTomodactylus.Frogs of the genusSyrrhophustend to occur at lower elevations than do their close relatives of the genusTomodactylus(Fig. 22). This generalization is complicated by the occurrence in the Sierra Madre Oriental in relatively high altitudeSyrrhophus(up to 2000 m.) and the occurrence in Michoacán of low altitudeTomodactylus(to sea level). There are noTomodactylusin the Sierra Madre Oriental, whereas the genusSyrrhophusis represented in the lowlands of western México (modestusgroup).SyrrhophusandTomodactylusexhibit essentially parapatric distributions. The two genera as now composed can be characterized as low to moderate elevation frogs (Syrrhophus) and moderate to intermediate elevation frogs (Tomodactylus).Literature CitedBaird, S. F.1859. Reptiles of the Boundary. United States and Mexican Boundary Survey, pp. 1-35, pls. 1-41.Barbour, T.1923. The reappearance of Batrachyla longipes. Proc. New England Zool. Club, 8:81-83.Barbour, T., andA. Loveridge1946. Typical reptiles and amphibians; supplement. Bull. Mus. Comp. Zool., 96:59-214.Boulenger, G. A.1882. Catalogue of the Batrachia Salientia ... British Museum., 2nd ed.1888. Note on the classification of the Ranidae. Proc. Zool. Soc. London, 1888, pt. 2:204-06.Campbell, H. W., andR. S. Simmons1962. Notes on some reptiles and amphibians from western Mexico. Bull. So. California Acad. Sci., 61:193-203.Conant, R.1958. A field guide to reptiles and amphibians. Houghton-Mifflin Co. Boston. 366 pp.Cope, E. D.1866. On the structures and distribution of the genera of the arciferous Anura. J. Acad. Nat. Sci. Philadelphia, n. ser., 6:67-112.1877. Tenth contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc., 17:85-98.1878. New genus of Cystignathidae from Texas. Amer. Nat., 12:252-53.1879. Eleventh contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc., 18:261-77.1885. A contribution to the herpetology of Mexico.Ibid., 22:379-404.Davis, W. B., andJ. R. Dixon1957. Notes on Mexican amphibians, with description of a newMicrobatrachylus. Herpetologica, 13:145-47.1965. Amphibians of the Chilpancingo Region, Mexico.Ibid., 20:225-33.Díaz de León, J.1904. Indice de los Batracios que se enquentran en la Republica Méxicana. Imprenta de Ricardo Rodriquez Romo. Aguascalientes. 40 pp.Dixon, J. R.1957. Geographic variation and distribution of the genus Tomodactylus in Mexico. Texas J. Sci., 9:379-409.Dixon, J. R., andR. G. Webb1966. A newSyrrhophusfrom Mexico (Amphibia: Leptodactylidae). Cont. Sc., Los Angeles Co. Mus., 102:1-5.Duellman, W. E.1958. A review of the frogs of the genusSyrrhophusin western Mexico. Occ. Pap. Mus. Zool. Univ. Michigan, 594:1-15.1960. A distributional study of the amphibians of the Isthmus of Tehuantepec, Mexico. Univ. Kansas Publs. Mus. Nat. Hist., 13:19-72.Firschein, I. L.1954. Definition of some little-understood members of the leptodactylid genusSyrrhophus, with a description of a new species. Copeia, (1):48-58.Fouquette, M. J.1960. Call structure in frogs of the family Leptodactylidae. Texas J. Sci., 12:201-15.Gadow, H.1905. The distribution of Mexican amphibians and reptiles. Proc. Zool. Soc. London, 1905, pt. 2:191-244.Gorham, S. W.1966. Liste der rezenten Amphibien und Reptilien.... Das Tierreich. Lief, 85:1-222.Günther, A. C. L. G.1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. 326 pp., 76 pls. Syrrhophus section dated 1900.Kellogg, R.1932. Mexican tailless amphibians in the United States National Museum. Bull. U.S. Natl. Mus., 160.Langebartel, D. A., andF. A. Shannon1956. A new frog (Syrrhophus) from the Sinoloan lowlands of Mexico. Herpetologica, 12:161-65.Lynch, J. D.1963. The status ofEleutherodactylus longipes(Baird) of Mexico (Amphibia: Leptodactylidae). Copeia, (3):580-81.1964. Additional hylid and leptodactylid remains from the Pleistocene of Texas and Florida Herpetologica. 20:141-42.1967.EpirhexisCope, 1866 (Amphibia: Salientia): request for suppression under the plenary powers. I. N. (S). Bull. Zool. Nomencl., 24:313-15.1968. Genera of leptodactylid frogs in México. Univ. Kansas Publs., Mus. Nat. Hist., 17:503-15.Martin, P. S.1958. A biogeography of reptiles and amphibians in the Gomez Farias region, Tamaulipas, Mexico. Misc. Publs. Mus. Zool. Univ. Michigan, 101:1-102.Milstead, W. M.,J. S. Mecham, andH. McClintock1950. The amphibians and reptiles of the Stockton Plateau in northern Terrell County, Texas. Texas J. Sci., 2:543-62.Neill, W. T.1965. New and noteworthy amphibians and reptiles from British Honduras. Bull. Florida State Mus., 9:77-130.Nieden, F.1923. Anura I ... Das Tierreich. Lief., 46:1-584.Peters, W.1871. Über neue Amphibien ... des Konigl. Zoologischen Museums. Monatsb. k. k. Preuss. Akad. Wiss. Berlin, 1870:641-52.Schmidt, K. P., andT. F. Smith1944. Amphibians and reptiles of the Big Bend Region of Texas. Field Mus. Nat. Hist., zool. ser., 29:75-96.Smith, H. M.1947. Notes on Mexican amphibians and reptiles. J. Washington Acad. Sci., 37:408-12.Smith, H. M., andE. H. Taylor1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U.S. Natl. Mus., 194:1-118.Stejneger, L.1915. A new species of tailless batrachian from North America. Proc. Biol. Soc. Washington, 28:131-32.Taylor, E. H.1940a. A new eleutherodactylid frog from Mexico. Proc. New England Zool. Club, 18:13-16.1940b. Two new anuran amphibians from Mexico. Proc. U.S. Natl. Mus., 89:43-47, 1 pl.1940c. A new Syrrhophus from Guerrero, Mexico. Proc. Biol. Soc. Washington, 53:95-98, 1 pl.1940d. New species of Mexican Anura. Univ. Kansas Sci. Bull., 26:385-405.1940e. Herpetological miscellany no. I.Ibid., 26:489-571.1942. New Caudata and Salientia from México.Ibid., 28:295-323.1943. Herpetological novelties from Mexico.Ibid., 29:343-61.1952. A review of the frogs and toads of Costa Rica.Ibid., 35:577-942.Taylor, E. H., andH. M. Smith1945. Summary of the collections of amphibians made in Mexico under the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. Natl. Mus., 95:521-613.Tihen, J. A.1960. Notes on Late Cenozoic hylid and leptodactylid frogs from Kansas, Oklahoma and Texas. Southwest. Nat., 5:66-70.Wright, A. H., andA. A. Wright1949. Handbook of frogs and toads. 3rd ed. Comstock. 640 pp.Yarrow, H. C.1882. Checklist of North American Reptilia and Batrachia, with catalogue of specimens in U.S. National Museum. Bull. U.S. Natl. Mus., 24:1-249.Zweifel, R. G.1960. Results of the Puritan-American Museum of Natural History Expedition to Western Mexico. 9. Herpetology of the Tres Marias Islands. Bull. Amer. Mus. Nat. Hist., 119:81-128.Transcriber's NotesAlthoughSyrrhophus marnockiandSyrrhophus marnockiiboth appear in this text, a literature search shows that both spellings have been used and the two instances where there is only one "i" at the end are in reference to priviously published names. Therefore, they were left as is. With the exception of the list below and a number of silent corrections, the text presented is that of the original printed version. The original cover was modified to include graphics from the article.Typographical CorrectionsPageCorrection3otherwse => otherwise5poltypic => polytypic12interorbtal => interorbital14neublosus => nebulosus16Cuidad => Ciudad161946-170 => 1946:17022rubrimacultaus => rubrimaculatus27resemblence => resemblance

Syrrhophus pipilansTaylor

Syrrhophus pipilansTaylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 (formerly EHT-HMS 6843), 14.6 km. S Mazatlán, Guerrero, México; collected on July 22, 1936, by Edward H. Taylor].

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

Fig. 15:Dicegrams of ear size relative to eye diameter in the two subspecies ofSyrrhophus pipilans. N = 17 innebulosus, 18 inpipilans.

Remarks.—Two subspecies were recognized by Duellman (1958). Previously both had been treated as species. The two populations were distinguished on the basis of color pattern and the size of the tympanum. Measurements of 17 males ofS. p. nebulosusfrom central Chiapas and 18 males ofS. p. pipilansfrom south-central Oaxaca and Guerrero, México, demonstrates that the supposed difference in tympanum size is not significant (Fig. 15).There is, however, a tendency for the western population ofS. pipilansto have larger tympani. Based on the present examination of 112 specimens of this species the two populations are held to be sufficiently distinct to warrant taxonomic recognition as subspecies (Fig. 16).

Fig. 16:Syrrhophus pipilans nebulosus(left, KU 58908) andS. p. pipilans(right, KU 86885). ×2.7.

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

Syrrhophus pipilans nebulosusTaylor

Syrrhophus nebulosusTaylor, 1943:353-55, pl. 27, figs. 3-5 [Holotype.—FMNH 100095 (formerly EHT-HMS 3774), near Tonolá, Chiapas, México; collected on August 27, 1935, by Hobart M. Smith and Edward H. Taylor]. Smith and Taylor, 1948:49, 51.

Syrrhophus pipilans nebulosus: Duellman, 1958:2-4, 9, 12, 14. Stuart, 1963:32-33. Gorham, 1966:166-67.

Diagnosis.—Diameter of tympanum 36.6-47.8 per cent that of eye; dorsum dark brown with numerous small light brown to yellowish spots.

Remarks.—The distribution of this subspecies is adequately described by Duellman (1958). Fouquette (1960) described the vocalization of this frog.

Etymology.—Latin,nebula, in reference to the clouded dorsal pattern.

Distribution.—Low to moderate elevations along the Pacific versant of Chiapas and in the Grijalva valley of Chiapas and Guatemala (Fig. 17).

Specimens examined.—(54) GUATEMALA,Huehuetenango: Jacaltenango, UMMZ 117036; 35 km. SE La Mesilla, TNHC 29652. MÉXICO,Chiapas: 11.2 km. N Arriaga, 300 m., UMMZ 125891; 11.8 km. N Arriaga, UMMZ 117279;12.8 km. N Arriaga, UMMZ 117280; 17.5 km. S Arriaga, UIMNH 57108-109; 1.5 km. S Bochil, 1250 m., KU 58898-908; Cerro Hueco, 7 km. S Tuxtla Gutierrez, UMMZ 123007; 3.2 km. S Ixtapa, UMMZ 124000; Linda Vista, ca. 2 km. NW Pueblo Nuevo Solistahuacán, KU 58897; Hda. Monserrate, 40 km. NW Arriaga, UMMZ 102258; near San Ricardo, FMNH 100720; Tapachula, FMNH 75792, 103242, 100695-96, UIMNH 13292; 56 km. E Tapanatepec, Oaxaca, TNHC 26942, Tonolá, FMNH 100095 (holotype), 100686-92, UIMNH 13293-95; Tuxtla Gutierrez, FMNH 100693-94, UIMNH 13297; 19 km. N Tuxtla Gutierrez, TNHC 25229-30; 15.5 km. NE Tuxtla Gutierrez, UMMZ 119892 (3); 19 km. NE Tuxtla Gutierrez, UMMZ 119891 (3); 8 km. NNW Tuxtla Gutierrez, KU 37809; Unión de Juarez, FMNH 105294.

Syrrhophus pipilans pipilansTaylor

?Syrrhopus verruculatus: Gadow, 1905:194.

Syrrhophus pipilansTaylor, 1940c:95-97, pl. 1 [Holotype.—FMNH 100072 (formerly EHT-HMS 6843), from 14.6 km. S Mazatlán, Guerrero, México; collected on July 22, 1936, by Edward H. Taylor]. Taylor and Smith, 1945:581-82. Smith and Taylor, 1948:49, 50-51.

Syrrhophus pipilans pipilans: Duellman, 1958:1-4, 8-9, 13-14, pl. 2, fig. 1. Gorham, 1966:166.

Diagnosis.—Diameter of tympanum 40.6-54.0 per cent that of eye; dorsum dark brown with large light spots or blotches.

Remarks.—Duellman's (1958) synopsis of this subspecies is adequate; the distribution has not been extended, but several records are now available which fill in gaps.

Fig. 17:Distribution ofSyrrhophus pipilans:nebulosus(open circles) andpipilans(solid circles).

Gadow's (1905) record ofS. verruculatusfrom "Buena Vista, S. Guerrero" is most likely applicable to this species. Gadow simply included the name in a list of the species he had collected during his trip in México (1902-04); nofurther comment was made on this species although references toSyrrhopus(sic) appear in several places in the paper and would appear to apply to the species he had.

Etymology.—Latin,pipilo, chirping, peeping, in reference to the call of the male.

Distribution.—Sea level to about 1800 meters along the Pacific versant of western México from central Guerrero to the Isthmus of Tehuantepec (Fig. 17).

Specimens examined.—(62). MÉXICO,Guerrero: Acapulco, UMMZ 110125; 6.4 km. N Acapulco, FMNH 100389, 100525; Agua del Obispo, 980-1000 m., FMNH 75791, 100518-21, 100526, KU 86884-86, UIMNH 13315, UMMZ 119152, 125890 (4); 13.3 km. NW Coyuca, UIMNH 38367, 71982-83; 14.5 km. S Mazatlán, FMNH 100072 (holotype), 100408, 100511-17, UIMNH 13302-309; Tierra Colorado, 300 m., KU 67961, UIMNH 13313-14; near El Treinte, FMNH 126639; Xaltinanguis, FMNH 100522-24, 126640.Oaxaca: Cacahuatepec, UIMNH 52853; 8 km. NW Río Canoa, 53 km. ESE Cuajinicuilapa, UIMNH 52852; 6.4 km. N El Candelaria, UIMNH 9501; 11.2 km. S El Candelaria, UIMNH 9502; 17 km. NE Juchatengo, 1600 m., KU 86887; 31.5 km. N Pochutla, UMMZ 123999 (2); 32.9 km. N Pochutla, 850 m., UMMZ 123996; 37.1 km. N Pochutla, UMMZ 123998 (2); 41.4 km. N Pochutla, UMMZ 123997 (2); Cerro Quiengola, FMNH 105653; 3.8 km. N Santiago Chivela, UMMZ 115449; 14.5 km. W Tehuantepec, UMMZ 115448 (2).

Syrrhophus interorbitalisLangebartel and Shannon

Syrrhophus interorbitalisLangebartel and Shannon, 1956: 161-65, figs. 1-2 [Holotype.—UIMNH 67061 (formerly FAS 9378), 36 mi. N Mazatlán, Sinaloa, México, collected on November 17, 1955, by E. C. Bay, J. C. Schaffner, and D. A. Langebartel]. Duellman, 1958:1-4, 10, 12, 14. Gorham, 1966:164-65.

Syrrhophis interorbitalis: Campbell and Simmons, 1962:194, fig. 1.

Fig. 18:Left to right.Syrrhophus interorbitalis(UIMNH 38095, ×1.5),S. nivocolimae(LACM 3203, ×1.3), andS. teretistes(KU 75263, ×1.5).

Diagnosis.—Medium sized frogs, only known male 25.6 mm. snout-vent, females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in males;finger tips expanded; first finger shorter than second; outer metatarsal tubercle one-third size of inner; skin of dorsum shagreened, that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of eye in both sexes; pale yellow-brown ground color mottled with brown; limb bands broad, much wider than narrow light interspaces; interorbital bar very long, edged with dark brown to black (Fig. 18).

Remarks.—Duellman's (1958) measurements and proportions ofS. interorbitaliswere based exclusively on the type series, which is composed of only females; therefore hisinterorbitalisdata are not comparable with the data for the other species in his table. Campbell and Simmons (1962) collected the only known male. The type series was collected beneath rocks in a stream bed; the collectors heard calling frogs in the bushes but were unable to obtain specimens (Langebartel and Shannon, 1956). Campbell and Simmons (1962) reported that their specimen had a poorly developed interorbital bar in life; in preservative the bar compares favorably with the bar in the female (Fig. 18).

Etymology.—Latin, in reference to the pale interocular band.

Distribution.—Pacific lowlands of Sinaloa, México (Fig. 20).

Specimens examined.—(10). MÉXICO,Sinaloa: 36 mi. N Mazatlán, UIMNH 38094-96, 67061 (holotype), 71970-74; 65 mi. N Mazatlán, LACM 13773.

Syrrhophus modestusTaylor

Syrrhophus modestusTaylor, 1942:304-06, pl. 29 [Holotype.—FMNH 100048 (formerly EHT-HMS 3756), from Hacienda Paso del Río, Colima, México; collected on July 8, 1935, by Hobart M. Smith]. Smith and Taylor, 1948:49-50.

Syrrhophus modestus modestus: Duellman, 1958:2-5, 7, 14, pl. 1, fig. 1. Gorham, 1966:166.

Diagnosis.—Small frogs, males 15.8-20.1 mm. snout-vent length, single female 18.5 mm.; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed; pale cream in preservative with dark brown spots; limbs banded; bands on forearm and thigh poorly developed or absent; interorbital bar absent.

Remarks.—The tympanum is concealed inS. modestus,S. nivocolimae,S. pallidus,S. teretistes, and to a lesser degree inS. interorbitalis. However, if the specimen is permitted to dry slightly, the annulus tympanicus becomes visible through the skin and a tympanum/eye ratio can be computed.

One of the few cases of sympatry within the genusSyrrhophusinvolves this species;modestusandnivocolimaeare known to be sympatric at one locality in southwestern Jalisco, México.

Duellman (1958) used the trinomial for this population and named a new subspecies,pallidus, from Nayarit. I considerpallidusto be specifically distinct frommodestusbecause there is no evidence of genetic exchange, and there is no overlap in the distinguishing morphological features. I do consider the two populations to be closely related but feel the inter-relationships betweenmodestus,pallidus,nivocolimae, andteretistesare more complex than would be indicated by the use of trinomials. The sympatric occurrence ofmodestusandnivocolimaeis significant; morphologically, they might otherwise be regarded as subspecies. Although allopatric, similar arguments could be advanced forthe morphologically similarpallidusandteretistes. The four are here afforded species rank since morphological similarity and allopatry are not sufficient grounds for the assumption of genetic exchange.

Fig. 19:Syrrhophus modestus[left, UMMZ 115447 (WED 11155)] andS. pallidus(right, UMMZ 115453). ×2.2.

Etymology.—Latin, meaning unassuming, modest, in reference to the small size of the species.

Distribution.—Low elevations (up to 700 meters) in the lowlands and foothills of Colima and southwestern Jalisco, México (Fig. 20).

Specimens examined.—(14). MÉXICO,Colima: Hda. Paso del Río, FMNH 100048 (holotype), 100167, 100299, UIMNH 13300, UMMZ 110877 (2), USNM 139729; 7.2 km. SW Tecolapa, UMMZ 115477 (4);Jalisco: 17.6 km. SW Autlan, 606 m., KU 102627; 3.2 km. N La Resolana, UMMZ 102100; Bahía Tenacatita, UMMZ 84264.

Syrrhophus nivocolimaeDixon and Webb

Syrrhophus nivocolimaeDixon and Webb, 1966:1-4, Fig. 1 [Holotype.—LACM 3200, from Nevado de Colima (6 airline miles west of Atenquique), Jalisco, México, 7800 feet; collected on July 20, 1964, by Robert G. Webb].

Diagnosis.—Small frogs, males 18.5-21.1 mm. snout-vent length, only known female 24.1 mm. snout-vent; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum warty, that of venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that of eye in males; mid-dorsal brown band from interorbital bar to anus; bands on limbs narrow, dark bandsless than one-half width of light bands, upper arm not banded; narrow interorbital light bar.

Remarks.—This species is closely related toS. modestusand differs in color pattern and degree of wartiness of the skin. Dixon and Webb (1966) held thatnivocolimaehad no close relatives, but the condition of the tympanum, size, nature of the outer palmar tubercle, relative sizes of the metatarsal tubercles, and shape and size of the digital pads all point to a close relationship betweenS. modestus,S. nivocolimae, andS. pallidus.

Fig. 20:Distribution of the species of themodestusgroup:interorbitalis(open circles),teretistes(solid circles),modestus(open triangles),pallidus(solid triangles) andnivocolimae(square). Arrow indicates locality of sympatry betweenmodestusandnivocolimae. Solid line about the localities forinterorbitalisis a range estimate based on call records and specimens examined.

Dixon and Webb (1966) reported thatS. nivocolimaehas a large tympanum (50.0-59.0 per cent diameter of eye). However, my examination of the type series and several other specimens from Jalisco reveals that thelargest tympanum/eye ratio is 39.3 per cent. Therefore, the tympanum/eye ratio inS. nivocolimaeis in agreement with those forS. modestus,S. pallidus, andS. teretistes(Table 6).

Etymology.—niv, Latin, and Colima (Nevado de), meaning high on the volcano, in reference to the higher distribution of this species (around 2000 meters) than other members of the group.

Distribution.—Known from southwestern Jalisco, México, at moderate to high elevations (600-2400 meters).

Specimens examined.—(48) MÉXICO,Jalisco: 17.6 km. SW Autlán, 606 m., KU 102626, 102631; 6.4 km. W Atenquique, 2060 m., KU 102628-30, 102632; 8 km. W Atenquique, 1970 m., LACM 3210-12; 9.6 km. W Atenquique, 2360 m., LACM 3200 (holotype), 3201-09; 14.5 km. W Atenquique, 2000 m., LACM 25424-36, 25439-41, 25446; 15 km. W Atenquique, LACM 37044-46, 37244-47; 16 km. W Atenquique, 2105 m., LACM 25443-45; 17 km. W Atenquique, 2180 m., LACM 25442.

Syrrhophus pallidusDuellman, New combination

Syrrhophus modestus: Davis and Dixon, 1957:146.

Syrrhophus modestus pallidusDuellman, 1958:2-3, 5-7, 14, pl. 3 [Holotype.—UMMZ 115452, from San Blas, Nayarit, México, sea level; collected on August 13, 1956, by William E. and Ann S. Duellman]. Zweifel, 1960:86-88, 91, 93-94, 118, 120-22. Gorham, 1966:166.

Syrrhophis modestus pallidus: Campbell and Simmons, 1962:194.

Diagnosis.—Small frogs, males 17.9-19.3 mm. snout-vent length; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed, its diameter 27.0-35.6 per cent of eye in males; ground color cream vermiculated with brown, upper arm and thigh lacking, or with few, indistinct, bands; interorbital bar absent.

Remarks.—Considerable debate has been waged relative to the value of subspecies and to the reasons for recognizing distinct disjunct populations as species versus subspecies. Lacking evidence of genetic exchange, I prefer to retain disjunct populations that are distinctive as species.

All known specimens ofpalliduscan be separated from those ofmodestusby color pattern. The two nominal species exhibit overlap in proportions but the same can be said about nearly every species ofSyrrhophus; therefore, overlap in proportions can be disregarded in assessing specific versus subspecific rank. Until contrary evidence is forthcoming, I consider the disjunct populations heretofore held to be subspecies ofmodestusto be specifically distinct. The specimens of the disjunct population ofpalliduson the Tres Marias do not differ from the mainland population in Nayarit. This evidence, though perhaps secondary, supports my contention that two species should be recognized.

Etymology.—Latin, in reference to the pale ground color in comparison with that ofS. modestus.

Distribution.—Low elevations in coastal Nayarit and on Islas Tres Marias (Fig. 20).

Specimens examined.—(12) MÉXICO,Nayarit: 18.8 mi. NW Ahuacatlán, UIMNH 7808; San Blas, UMMZ 115452 (holotype), 115453-57; 17 km. NE San Blas, 150 m., MSU 5085; 12.8 km. E San Blas, UIMNH 71979; 31 km. E San Blas, UIMNH 71978; 13.5 km. N Tepic, UIMNH 71980-81.

Syrrhophus teretistesDuellman

Syrrhophus teretistesDuellman, 1958:2-3, 10-14, pl. 2, fig. 2 [Holotype.—UMMZ 115451, from 4.8 km. NW Tepic, Nayarit, México, 840 m.; collected on August 12, 1956, by William E. Duellman]. Gorham, 1966:167.

Diagnosis.—Medium-sized frogs, males 19.2-23.2 mm. snout-vent length, single known female 24.8 mm. snout-vent; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum partially concealed, its diameter 28.6-43.8 per cent of eye in males; ground color brown vermiculated with dark brown to nearly black; upper arm and thigh banded; interorbital light bar absent.

Remarks.—S. teretistesappears to be most closely related toS. pallidus; I consider it to be an upland derivative ofpallidus. Morphologically, the differences between the two are few, but lacking evidence of genetic exchange they are retained as species.

Etymology.—Greek, in reference to the whistle-like nature of the call.

Distribution.—Moderate elevations (840-1200 meters) in the Sierra Occidental of Nayarit, Sinaloa, and Durango, México (Fig. 20).

Specimens examined.—(13) MÉXICO,Nayarit: 4.8 km. NW Tepic, 840 m., UMMZ 115451 (holotype).Sinaloa: Santa Lucía, 1090 m., KU 75263-72; 1 km. NE Santa Lucía, 1156 m., KU 78257; 2.2 km. NE Santa Lucía, 1156 m., KU 78258.

Discussion

There are relatively few clear-cut morphological differences among the fourteen species now assigned toSyrrhophus. The majority of the species are allopatric and differ primarily in color patterns. Sympatric occurrence serves as an indicator of specific distinctness and is one of the more practical tests of species validity when cross-breeding experiments are not possible. Two cases of sympatric occurrence are known for the species ofSyrrhophusin western México:modestusandnivocolimaeare sympatric in southern Jalisco andpipilans nebulosusandrubrimaculatusare sympatric in southeastern Chiapas. In eastern México,longipesandverrucipesare sympatric in southern Hidalgo, andlongipesis sympatric withcystignathoides,dennisi, andguttilatusin southern Tamaulipas.Syrrhophus cystignathoidesandleprusare apparently sympatric in central Veracruz.

Subspecific assignments have been made only when there is evidence of intergradation. The sympatric occurrence of morphologically similar species in this genus has led me to adopt a conservative approach to the degree of difference philosophy. I have therefore recognized all morphologically distinct allopatric populations as species.

Fig. 21:Generic distributions ofSyrrhophus(stipple) andTomodactylus(hatching). Black areas are zones of intergeneric sympatry.

Fig. 22:Altitudinal distributions ofSyrrhophusandTomodactylus. Widths of the columns are proportional to the numbers of species at a given altitude; narrowest width equals one species.

Syrrhophusis closely allied to another Mexican leptodactylid genus,Tomodactylus, which was revised by Dixon (1957), who along with numerous other authors noted the close relationship between the two genera. There is an almost complete lack of sympatry between the two genera; in very few places in México do they coexist (Fig. 21).Tomodactylushas its greatest diversity in the Cordillera Volcánica and Sierra Madre del Sur, whereasSyrrhophusreaches its greatest diversity in the Sierra Madre Oriental and eastern foothills. The species of both genera are about the same size and presumably have similar requirements insofar as food, breeding sites, and habitat selection.

Four cases of intergeneric sympatry are known for the two genera: 1) the Chilpancingo region of Guerrero, 2) the lowlands of Colima and the mountains just inland in Jalisco, 3) the lowlands of central Nayarit, and 4) the Sierra Madre Occidental on the Durango-Sinaloan border. The apparent sympatry in the Chilpancingo region involves four species:S. pipilans,T. albolabris,T. dilatus, andT. nitidus. Of the four,T. dilatusappears to be completely allopatric in that it occurs at higher altitudes (above 2000 meters), whereas the other three occur below 1800 meters in the region (Davis and Dixon, 1965). In the Colima-Jalisco region,Tomodactylustendsto occur higher (Dixon and Webb, 1966) than some of theSyrrhophus, but one subspecies ofTomodactylus nitidusis a lowland frog, occurring sympatrically with the lowlandSyrrhophus modestus. A similar situation is observed in Nayarit; the lowlandTomodactylusoccurs sympatrically with the smallSyrrhophus pallidus. In both cases theSyrrhophusis smaller than theTomodactylus.

Frogs of the genusSyrrhophustend to occur at lower elevations than do their close relatives of the genusTomodactylus(Fig. 22). This generalization is complicated by the occurrence in the Sierra Madre Oriental in relatively high altitudeSyrrhophus(up to 2000 m.) and the occurrence in Michoacán of low altitudeTomodactylus(to sea level). There are noTomodactylusin the Sierra Madre Oriental, whereas the genusSyrrhophusis represented in the lowlands of western México (modestusgroup).SyrrhophusandTomodactylusexhibit essentially parapatric distributions. The two genera as now composed can be characterized as low to moderate elevation frogs (Syrrhophus) and moderate to intermediate elevation frogs (Tomodactylus).

Literature Cited

Baird, S. F.

1859. Reptiles of the Boundary. United States and Mexican Boundary Survey, pp. 1-35, pls. 1-41.

Barbour, T.

1923. The reappearance of Batrachyla longipes. Proc. New England Zool. Club, 8:81-83.

Barbour, T., andA. Loveridge

1946. Typical reptiles and amphibians; supplement. Bull. Mus. Comp. Zool., 96:59-214.

Boulenger, G. A.

1882. Catalogue of the Batrachia Salientia ... British Museum., 2nd ed.

1888. Note on the classification of the Ranidae. Proc. Zool. Soc. London, 1888, pt. 2:204-06.

Campbell, H. W., andR. S. Simmons

1962. Notes on some reptiles and amphibians from western Mexico. Bull. So. California Acad. Sci., 61:193-203.

Conant, R.

1958. A field guide to reptiles and amphibians. Houghton-Mifflin Co. Boston. 366 pp.

Cope, E. D.

1866. On the structures and distribution of the genera of the arciferous Anura. J. Acad. Nat. Sci. Philadelphia, n. ser., 6:67-112.

1877. Tenth contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc., 17:85-98.

1878. New genus of Cystignathidae from Texas. Amer. Nat., 12:252-53.

1879. Eleventh contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc., 18:261-77.

1885. A contribution to the herpetology of Mexico.Ibid., 22:379-404.

Davis, W. B., andJ. R. Dixon

1957. Notes on Mexican amphibians, with description of a newMicrobatrachylus. Herpetologica, 13:145-47.

1965. Amphibians of the Chilpancingo Region, Mexico.Ibid., 20:225-33.

Díaz de León, J.

1904. Indice de los Batracios que se enquentran en la Republica Méxicana. Imprenta de Ricardo Rodriquez Romo. Aguascalientes. 40 pp.

Dixon, J. R.

1957. Geographic variation and distribution of the genus Tomodactylus in Mexico. Texas J. Sci., 9:379-409.

Dixon, J. R., andR. G. Webb

1966. A newSyrrhophusfrom Mexico (Amphibia: Leptodactylidae). Cont. Sc., Los Angeles Co. Mus., 102:1-5.

Duellman, W. E.

1958. A review of the frogs of the genusSyrrhophusin western Mexico. Occ. Pap. Mus. Zool. Univ. Michigan, 594:1-15.

1960. A distributional study of the amphibians of the Isthmus of Tehuantepec, Mexico. Univ. Kansas Publs. Mus. Nat. Hist., 13:19-72.

Firschein, I. L.

1954. Definition of some little-understood members of the leptodactylid genusSyrrhophus, with a description of a new species. Copeia, (1):48-58.

Fouquette, M. J.

1960. Call structure in frogs of the family Leptodactylidae. Texas J. Sci., 12:201-15.

Gadow, H.

1905. The distribution of Mexican amphibians and reptiles. Proc. Zool. Soc. London, 1905, pt. 2:191-244.

Gorham, S. W.

1966. Liste der rezenten Amphibien und Reptilien.... Das Tierreich. Lief, 85:1-222.

Günther, A. C. L. G.

1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. 326 pp., 76 pls. Syrrhophus section dated 1900.

Kellogg, R.

1932. Mexican tailless amphibians in the United States National Museum. Bull. U.S. Natl. Mus., 160.

Langebartel, D. A., andF. A. Shannon

1956. A new frog (Syrrhophus) from the Sinoloan lowlands of Mexico. Herpetologica, 12:161-65.

Lynch, J. D.

1963. The status ofEleutherodactylus longipes(Baird) of Mexico (Amphibia: Leptodactylidae). Copeia, (3):580-81.

1964. Additional hylid and leptodactylid remains from the Pleistocene of Texas and Florida Herpetologica. 20:141-42.

1967.EpirhexisCope, 1866 (Amphibia: Salientia): request for suppression under the plenary powers. I. N. (S). Bull. Zool. Nomencl., 24:313-15.

1968. Genera of leptodactylid frogs in México. Univ. Kansas Publs., Mus. Nat. Hist., 17:503-15.

Martin, P. S.

1958. A biogeography of reptiles and amphibians in the Gomez Farias region, Tamaulipas, Mexico. Misc. Publs. Mus. Zool. Univ. Michigan, 101:1-102.

Milstead, W. M.,J. S. Mecham, andH. McClintock

1950. The amphibians and reptiles of the Stockton Plateau in northern Terrell County, Texas. Texas J. Sci., 2:543-62.

Neill, W. T.

1965. New and noteworthy amphibians and reptiles from British Honduras. Bull. Florida State Mus., 9:77-130.

Nieden, F.

1923. Anura I ... Das Tierreich. Lief., 46:1-584.

Peters, W.

1871. Über neue Amphibien ... des Konigl. Zoologischen Museums. Monatsb. k. k. Preuss. Akad. Wiss. Berlin, 1870:641-52.

Schmidt, K. P., andT. F. Smith

1944. Amphibians and reptiles of the Big Bend Region of Texas. Field Mus. Nat. Hist., zool. ser., 29:75-96.

Smith, H. M.

1947. Notes on Mexican amphibians and reptiles. J. Washington Acad. Sci., 37:408-12.

Smith, H. M., andE. H. Taylor

1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U.S. Natl. Mus., 194:1-118.

Stejneger, L.

1915. A new species of tailless batrachian from North America. Proc. Biol. Soc. Washington, 28:131-32.

Taylor, E. H.

1940a. A new eleutherodactylid frog from Mexico. Proc. New England Zool. Club, 18:13-16.

1940b. Two new anuran amphibians from Mexico. Proc. U.S. Natl. Mus., 89:43-47, 1 pl.

1940c. A new Syrrhophus from Guerrero, Mexico. Proc. Biol. Soc. Washington, 53:95-98, 1 pl.

1940d. New species of Mexican Anura. Univ. Kansas Sci. Bull., 26:385-405.

1940e. Herpetological miscellany no. I.Ibid., 26:489-571.

1942. New Caudata and Salientia from México.Ibid., 28:295-323.

1943. Herpetological novelties from Mexico.Ibid., 29:343-61.

1952. A review of the frogs and toads of Costa Rica.Ibid., 35:577-942.

Taylor, E. H., andH. M. Smith

1945. Summary of the collections of amphibians made in Mexico under the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. Natl. Mus., 95:521-613.

Tihen, J. A.

1960. Notes on Late Cenozoic hylid and leptodactylid frogs from Kansas, Oklahoma and Texas. Southwest. Nat., 5:66-70.

Wright, A. H., andA. A. Wright

1949. Handbook of frogs and toads. 3rd ed. Comstock. 640 pp.

Yarrow, H. C.

1882. Checklist of North American Reptilia and Batrachia, with catalogue of specimens in U.S. National Museum. Bull. U.S. Natl. Mus., 24:1-249.

Zweifel, R. G.

1960. Results of the Puritan-American Museum of Natural History Expedition to Western Mexico. 9. Herpetology of the Tres Marias Islands. Bull. Amer. Mus. Nat. Hist., 119:81-128.

Transcriber's NotesAlthoughSyrrhophus marnockiandSyrrhophus marnockiiboth appear in this text, a literature search shows that both spellings have been used and the two instances where there is only one "i" at the end are in reference to priviously published names. Therefore, they were left as is. With the exception of the list below and a number of silent corrections, the text presented is that of the original printed version. The original cover was modified to include graphics from the article.Typographical CorrectionsPageCorrection3otherwse => otherwise5poltypic => polytypic12interorbtal => interorbital14neublosus => nebulosus16Cuidad => Ciudad161946-170 => 1946:17022rubrimacultaus => rubrimaculatus27resemblence => resemblance

Transcriber's Notes

AlthoughSyrrhophus marnockiandSyrrhophus marnockiiboth appear in this text, a literature search shows that both spellings have been used and the two instances where there is only one "i" at the end are in reference to priviously published names. Therefore, they were left as is. With the exception of the list below and a number of silent corrections, the text presented is that of the original printed version. The original cover was modified to include graphics from the article.

Typographical Corrections

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