Part 2

_Voice._--The voices of all _Syrrhophus_ can be described as a single short chirp or peep; without audiospectrographic analyses the significance of the differences between a chirp, peep, or short whistle cannot be appreciated. Martin (1958) and Wright and Wright (1949) reported multi-noted calls, and one collector of _S. verrucipes_ noted the frog "trilled."

Fouquette (1960) presented analyses of two species (_marnockii_ and _pipilans nebulosus_). The voices were very similar; both frogs were reported to "trill" and "chirp."

SYSTEMATIC ACCOUNT

The genus _Syrrhophus_ has been defined (Lynch, 1968) and limited to the group of species occurring in Guatemala, México and the United States. The closest relatives of _Syrrhophus_ are the frogs of the genus _Tomodactylus_ (Dixon, 1957; Firschein, 1954). Lynch (1968) implied there were no osteological bases for the separation of _Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_. At that time, I believed such to be the case and derived _Syrrhophus_ and _Tomodactylus_ from the _rhodopis_ complex of _Eleutherodactylus_, with which they share terrestrial habits and relatively short limbs. In the _rhodopis_ complex there is a tendency for the loss of the outer palmar tubercle, a not uncommon condition in _Syrrhophus_ and _Tomodactylus_.

However, the skulls of _Syrrhophus_ and _Tomodactylus_ show departures from the pattern observed in the Middle American _Eleutherodactylus_, as well as many of those species in western South America. Baldauf and Tanzer (1965) reported that the frontoparietals and prootics were fused in _Syrrhophus marnockii_ and that the prootics and exoccipitals appeared to be one bone (otoccipital). The otoccipital is not uncommon in eleutherodactyline frogs, but the fusion of the frontoparietals with the prootics (regardless of the fusion of the latter with the exoccipital) is uncommon in the family. I have found the frontoparietal-prootic fusion only in _Syrrhophus_ (all species), _Tomodactylus_ (all species), and _Eleutherodactylus_ (West Indies species). None of the Middle American _Eleutherodactylus_ has the two bones fused. Examination of the character is difficult in dried skeletal preparations. Cleared and stained or macerated preparations are satisfactory for checking this character.

Thus, in addition to the presence of numerous plantar supernumerary tubercles in the frogs of the genera _Syrrhophus_ and _Tomodactylus_, these two genera can be separated from other Middle American eleutherodactylines by the fusion of the frontoparietals and prootics. This character not only further strengthens the argument that the two genera are closely related but poses a problem of zoogeographic analysis of the distribution of the character, which will be discussed fully elsewhere.

Key to the Species of the Frog Genus _Syrrhophus_

1. Three large, well-developed palmar tubercles 2

Two large palmar tubercles; outer (third) palmar tubercle reduced in size or absent 9

2. Digital pads more than twice (usually three or more) times width of digit 3

Digital pads less than twice width of digit 4

3. Males having vocal slits; dorsum vermiculate; diameter of tympanum in males about one-half diameter of eye _S. dennisi_

Males lacking vocal slits; dorsum flecked, spotted, or blotched; diameter of tympanum in male about three-fourths that of eye _S. longipes_

4. First finger longer than second 5

First finger shorter than or equal to second 7

5. Venter smooth; dorsum spotted or vermiculate _S. leprus_

Venter areolate, or if smooth, dorsum flecked and interorbital bar lacking 6

6. Venter areolate; interorbital bar present; ground color yellowish _S. cystignathoides cystignathoides_

Venter smooth; interorbital bar absent; ground color brown _S. cystignathoides campi_

7. First finger shorter than second; digital tips only slightly dilated; green in life with darker green spots _S. verrucipes_

First finger equal to second; digital tips slightly to moderately expanded 8

8. Dorsum vermiculate; interorbital bar present; ground color cream to brown in life _S. guttilatus_

Dorsum punctate or flecked; interorbital bar absent; ground color green in life _S. marnockii_

9. Dorsum dark with pale (red in life) spots; digital pads not expanded _S. rubrimaculatus_

Dorsum pale with dark markings and digital pads slightly to widely expanded 10

10. Digital tips not widely expanded; tympanum well-defined; outer metatarsal tubercle more than one-half size of inner 11

Digital tips widely expanded, truncate in outline; tympanum poorly defined; outer metatarsal tubercle less than one-half size of inner 12

11. Dorsum dark brown with large light spots or blotches; tympanum/eye ratio usually greater than 43 percent _S. pipilans pipilans_

Dorsum dark brown with small light spots; tympanum/eye ratio less than 48 percent _S. pipilans nebulosus_

12. Light interorbital bar present 13

Light interorbital bar absent 14

13. Adults small, less than 22 mm. snout-vent length with a broad mid-dorsal stripe; dark bands on shank narrower than light interspaces _S. nivocolimae_

Adults larger, more than 22 mm. snout-vent length; dorsum vermiculate; dark bands on shank broader than light interspaces _S. interorbitalis_

14. Dorsum spotted with discrete black spots; pattern definite _S. modestus_

Dorsum reticulate or vermiculate, pattern poorly defined 15

15. Adults small, less than 21 mm. snout-vent length; upper arm not banded _S. pallidus_

Adults larger, usually greater than 21 mm. snout-vent length; upper arm banded _S. teretistes_

SPECIES ACCOUNTS

The following accounts do not include complete descriptions of each taxon, because a more than adequate number of descriptions is available in the recent (1940-1966) literature. An abbreviated synonymy, in which are listed all combinations and emendations of names and significant contributions to our knowledge of the taxon, is given for each. For each species and subspecies the following are given: descriptive diagnosis, statement of range, remarks on taxonomy, list of specimens examined, illustration of color pattern, and distribution map.

=Syrrhophus cystignathoides= (Cope)

_Phyllobates cystignathoides_ Cope, 1877:89-90 [Syntypes.--Originally USNM 32402-32409, (32405 now in MCZ) from Potrero, near Córdoba, Veracruz, México, Francis Sumichrast collector.]

_Diagnosis._--Adults small, males 16.0 to 23.5 mm. in snout-vent length, females 16.0-25.8 mm. in snout-vent length; vocal slits present in males; finger tips slightly expanded; first finger longer than second; outer metatarsal tubercle one-half size of inner, conical, compressed; skin of dorsum weakly pustular, that of venter smooth to areolate; tympanum 44 to 69 per cent diameter of eye (mean 55.5 per cent); ground color yellow to brown in life with brown to black fleckings on dorsum and flanks; limbs banded; interorbital bar present or not.

_Remarks._--Two geographic races (subspecies) are herein recognized; previously these were held by various authors to be species (_campi_ and _cystignathoides_). Intergradation occurs in southern Tamaulipas and eastern San Luis Potosí, México. The two subspecies can be distinguished on the basis of color pattern and the condition of the skin of the venter.

_Distribution._--Low to moderate elevations from the Río Grande embayment to central Veracruz, México (Fig. 5).

=Syrrhophus cystignathoides campi= Stejneger, New combination

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

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

_Remarks._--Martin (1958) was the first author to point out that _S. campi_ was probably a subspecies of the more southern _S. cystignathoides_. Various references in the literature might lead one to believe that the two were sympatric over much of northeastern 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).

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

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

_Remarks._--Firschein (1954) briefly considered the status of Peters' (1871) _Phyllobates verruculatus_ and noted that if it was a _Syrrhophus_ it would probably be referrable to _S. cystignathoides_. Peters' (1871) original description corresponds well with _S. cystignathoides_, and the type-locality ("Huanusco" = Huatusco) is within the range of that species. Firschein (1954) expressed doubt that _verruculatus_ was a _Syrrhophus_, because Peters placed it in another genus. However, Peters described _verruculatus_ a decade before Cope diagnosed the genus Syrrhophus. Most frogs now called _Syrrhophus_, plus a number of lower Central American frogs now placed in a variety of genera were placed in _Phyllobates_ by Boulenger, Cope, and Peters.

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

Smith and Taylor (1948) reported _S. verruculatus_ from Tianguistengo, Hidalgo, México. These specimens are examples of _verrucipes_. Smith (1947) reported a specimen of _verruculatus_ from San Lorenzo, Veracruz. Firschein (1954) referred it to _cystignathoides_, and Duellman (1960) concluded that both authors were in error and that the specimen (USNM 123530) was a _leprus_.

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

_Distribution._--Low and moderate elevations in the foothills along the Sierra Madre Oriental from eastern San Luis 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 leprus= Cope

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

_Diagnosis._--Medium-sized frogs, males 20.5-26.5 mm. in snout-vent, females 22.0-29.3 mm. in snout-vent length; vocal slits present in males; tips of fingers dilated slightly; first finger longer than second; inner metatarsal tubercle twice size of small, conical outer metatarsal tubercle; skin of dorsum pustular, that of venter smooth; snout 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.

_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) regarded _S. leprus_ as having a gray venter. Neill (1965) characterized his new subspecies on the basis of white venter and spots on the dorsum. Some specimens from throughout the range have only small round spots, instead of vermiculations (Fig. 6). The gray ventral coloration is largely restricted to the population in Los Tuxtlas, Veracruz, but only about 80 per cent of the specimens from the Los Tuxtlas have gray venters, whereas specimens from Guatemala, Oaxaca, Tabasco, and central Veracruz, 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, 1.6 km. S Catemaco, UMMZ 118222 (2); Volcán San Martín, south slope, UMMZ 118223; Volcán San Martín, Rancho El Tular, UIMNH 35399-400, 40340-41.

=Syrrhophus rubrimaculatus= Taylor and Smith

_Syrrhophus rubrimaculatus_ Taylor and Smith, 1945:583-85 [Holotype.--USNM 114070, from La Esperanza, near Escuintla, Chiapas, 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.

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

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

_Syrrhophus rubrimaculatus_ is probably best treated as a Pacific derivative of the _leprus_ group, even though the palmar tubercles do not agree. The removal of _rubrimaculatus_ from the western complex results in a more 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 guttilatus_ Cope, 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 smithi_ Taylor, 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 gaigeae_ Schmidt and Smith, 1944:80 [Holotype.--FMNH 27361, from the Basin, Chisos Mountains, Brewster Co., Texas; collected on July 24, 1937, by Walter L. Necker].

_Syrrhophus petrophilus_ Firschein, 1954:50-52 [Holotype.--UIMNH 7807, from 5 km. SW San Luis 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).