The Project Gutenberg EBook of A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope, by John D. Lynch This eBook is for the use of anyone anywhere at no cost and with almost no restrictions whatsoever. You may copy it, give it away or re-use it under the terms of the Project Gutenberg License included with this eBook or online at www.gutenberg.org Title: A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope Author: John D. Lynch Release Date: October 21, 2011 [EBook #37809] Language: English Character set encoding: ISO-8859-1 *** START OF THIS PROJECT GUTENBERG EBOOK A TAXONOMIC REVISION OF THE *** Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and the Online Distributed Proofreading Team at http://www.pgdp.net UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Volume 20, No. 1, pp. 1-45, 22 figs. February 20, 1970 A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope BY JOHN D. LYNCH UNIVERSITY OF KANSAS LAWRENCE 1970 University of Kansas Publications, Museum of Natural History Editors of this number: Frank B. Cross, Philip S. Humphrey, William E. Duellman [Pg 2] Volume 20, No. 1, pp. 1-45, 22 figs. Published February 20, 1970 University of Kansas Lawrence, Kansas PRINTED BY THE UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS 1970 A Taxonomic Revision of the Leptodactylid Frog Genus Syrrhophus Cope BY JOHN D. LYNCH INTRODUCTION Cope (1878) proposed the genus Syrrhophus for a medium-sized leptodactylid frog from central Texas; in the ensuing 75 years the genus was expanded to include a heterogeneous group of frogs ranging from Texas to Peru. Taylor (1952) and Firschein (1954) limited the genus to several species of frogs occurring in Guatemala, México, and Texas. Lynch (1968) provided a definition of the previously loosely-defined genus. With the exception of Taylor (1952), who treated the Costa Rican species, none of these authors dealt with the present status of the nineteen species erroneously assigned to Syrrhophus. These species are listed in Tables 1 and 2 with the name currently applied. Some of them are new combinations and their justifications will be published elsewhere. Gorham (1966) is the most recent author to include South American species in the genus Syrrhophus. Smith and Taylor (1948) recognized two species groups of the genus in México, an eastern and a western group (here termed complexes for purposes of discussion), separated on the basis of the number of palmar (metacarpal) tubercles (three palmar tubercles in the members of the eastern complex and two in those of the western complex). Duellman (1958) reviewed the species of the genus occurring in western México and concluded that there were five species (two polytypic). Dixon and Webb (1966) described an additional species from Jalisco, México. The distributions of some species have been extended, but otherwise the western complex of species remains unchanged since Duellman's review. Smith and Taylor (1948) recognized seven species of the genus in eastern México. Firschein revised the eastern complex (as then understood), and in so doing added one new species and treated Syrrhophus verruculatus as a [Pg 3] nomen dubium. Dixon (1957) redefined the related genus Tomodactylus and transferred T. macrotympanum Taylor to the genus Syrrhophus. Neill (1965) described a new subspecies of S. leprus from British Honduras. Two species (S. gaigeae and S. marnockii) were recognized in Texas until Milstead, Mecham, and McClintock (1950) synonymized S. gaigeae with S. marnockii. Thus, at present, nine species (one polytypic) are recognized on the eastern slopes and lowlands from central Texas to British Honduras. These are currently placed on one species group equivalent to the western complex reviewed by Duellman (1958). Table 1—Species Described as Members of the Genus Syrrhophus but Now Placed in Other Genera. Trivial name and author Current combination areolatus Boulenger, 1898 Eleutherodactylus areolatus calcaratus Andersson, 1945 Eleutherodactylus anderssoni caryophyllaceus Barbour, 1928 Eleutherodactylus caryophyllaceus coeruleus Andersson, 1945 Eleutherodactylus coeruleus ineptus Barbour, 1928 Eleutherodactylus diastema juninensis Shreve, 1938 Eupsophus juninensis lutosus Barbour and Dunn, 1921 Eleutherodactylus lutosus molinoi Barbour, 1928 Eleutherodactylus molinoi montium Shreve, 1938 Niceforonia montia mystaceus Barbour, 1922 Eleutherodactylus rhodopis obesus Barbour, 1928 Eleutherodactylus punctariolus omiltemanus Gunther, 1900 Eleutherodactylus omiltemanus[1] pardalis Barbour, 1928 Eleutherodactylus pardalis [1] New combination. Table 2—Species Incorrectly Regarded as Members of the Genus Syrrhophus but Described as Members of Other Genera. Trivial name, original generic assignment, and author Current combination chalceus (Phyllobates) Peters, 1873 Eleutherodactylus chalceus festae (Paludicola) Peracca, 1904 Niceforonia festae hylaeformis (Phyllobates) Cope, 1875 Eleutherodactylus hylaeformis palmatus (Phyllobates) Werner, 1899 Colostethus palmatus ridens (Phyllobates) Cope, 1866 Eleutherodactylus ridens simonsii (Paludicola) Boulenger, 1900 Niceforonia simonsii Table 3—Nominal Species of Syrrhophus (sensu strictu) and the Name Used Herein. Original combination Current combination campi, Syrrhophus cystignathoides campi cholorum, Syrrhophus leprus leprus cystigathoides, Phyllobates cystignathoides cystignathoides dennisi, Syrrhophus dennisi new species gaigeae, Syrrhophus guttilatus guttilatus, Malachylodes guttilatus interorbitalis, Syrrhophus interorbitalis latodactylus, Syrrhophus longipes leprus, Syrrhophus leprus longipes, Batrachyla longipes macrotympanum, Tomodactylus verrucipes [Pg 4] marnockii, Syrrhophus marnockii modestus, Syrrhophus modestus nebulosus, Syrrhophus pipilans nebulosus nivocolimae, Syrrhophus nivocolimae pallidus, Syrrhophus modestus pallidus petrophilus, Syrrhophus guttilatus pipilans, Syrrhophus pipilans pipilans rubrimaculatus, Syrrhophus rubrimaculatus smithi, Syrrhophus guttilatus teretistes, Syrrhophus teretistes verrucipes, Syrrhophus verrucipes verruculatus, Phyllobates Nomen dubium In the course of preparing an account of the species of Eleutherodactylus occurring in México and northern Central America, it became necessary to reëxamine the status of the genus Syrrhophus and its nominal species. It soon became evident that there were more names than species, that some previously regarded species were geographic variants, and that the eastern and western groups (complexes here) were artificial divisions of the genus. I conclude that there are seven species (one polytypic) of Syrrhophus in eastern México, Texas, and El Petén of Guatemala, and seven species (one polytypic) in western México. The current status of each of the 23 names correctly assigned to the genus is presented in Table 3. The fourteen species recognized by me are placed in five species groups. Two of these groups are presently placed in the western complex (modestus and pipilans groups) and three in the eastern complex (leprus, longipes and marnockii groups). The two complexes do not correspond exactly with the eastern and western groups of Smith and Taylor (1948), Firschein (1954), and Duellman (1958) since S. rubrimaculatus is now associated with the eastern leprus group. The definitions and contents of the five species groups are as follows: leprus group: digital pads not or only slightly expanded, rounded in outline; first finger longer or shorter than second; snout acuminate or subacuminate, not rounded; outer metatarsal tubercle conical; digits lacking distinct lateral fringes. content: cystignathoides, leprus and rubrimaculatus. longipes group: digital pads widely expanded, triangular in outline; first finger shorter than second; snout acuminate; outer metatarsal tubercle not conical; digits bearing lateral fringes. content: dennisi and longipes. marnockii group: digital pads expanded, rounded to truncate in outline; first finger equal in length to second or slightly shorter; snout rounded; outer metatarsal tubercle not conical; digits lacking lateral fringes; generally stout-bodied frogs. content: guttilatus, marnockii, and verrucipes. modestus group: digital pads expanded, truncate in outline; first and second fingers subequal in length, first usually slightly shorter than second; snout subacuminate; inner metatarsal tubercle twice as large (or larger) as outer metatarsal tubercle; digits bearing poorly-defined lateral fringes. content: interorbitalis, modestus, nivocolimae, pallidus, and teretistes. pipilans group: digital pads not or only slightly expanded, truncate in outline; first finger equal in length to second; snout subacuminate; metatarsal tubercles subequal in size; digits lacking lateral fringes. content: pipilans. Acknowledgments.—For loan of specimens, I am indebted to Richard J. Baldauf, Texas A & M University (TCWC); W. Frank Blair, University of Texas (TNHC); Charles M. Bogert and Richard G. Zweifel, American Museum of Natural History (AMNH); James E. Böhlke and Edmond V. Malnate, Academy of Natural Sciences of Philadelphia (ANSP); Robert F. Inger and Hymen Marx, Field Museum of Natural History (FMNH); Ernest A. Liner (EAL); Michael Ovchynnyk, Michigan State University collection (MSU); James A. Peters, United States National Museum (USNM); Douglas A. Rossman, Louisiana State University Museum of Zoology (LSUMZ); Hobart M. Smith, University of Illinois Museum of Natural History (UIMNH); Charles F. Walker, University of Michigan Museum of Zoology (UMMZ); and John W. Wright, Los Angeles County Museum (LACM). Specimens in the collection at the University of Kansas Museum of Natural History are identified as KU. The abbreviations EHT-HMS refer to the Edward H. Taylor-Hobart M. Smith collection and FAS to the Frederick A. Shannon collection. The type-specimens from these collections are now in the Field Museum of Natural History and the University of Illinois Museum of Natural History. I have profited from discussions concerning this problem with several persons, most notably William E. Duellman, Hobart M. Smith, Edward H. Taylor and Charles F. Walker. Nevertheless, the ideas and conclusions presented here should not be construed as necessarily reflecting their opinions. [Pg 5] [Pg 6] David M. Dennis executed all of the figures, and my wife, Marsha, typed the manuscript. Materials and Methods.—In the course of this study, 1003 specimens of the genus were examined. The holotypes of 21 of the 23 nominal species are extant; I have examined 19 of these. Nine measurements were taken, and five ratios computed for each of 338 specimens. Females are available for all species but one; thus, measurements were taken on individuals of both sexes. ANALYSIS OF CHARACTERS Size and proportions.—Frogs of this genus range in size from 16 to 40 mm. in snout-vent length. Five species are relatively small: S. cystignathoides, modestus, nivocolimae, pallidus and rubrimaculatus; one, S. longipes, is relatively large, and the remaining eight species are intermediate in size (22-30 mm.). Males are generally smaller than females and have proportionately longer heads and usually larger tympani. No significant differences were found among proportions, except that S. longipes has a larger tympanum/eye ratio than any other species. Frogs in the Syrrhophus marnockii group tend to have shorter shanks and feet, thereby giving those species a more stocky appearance. However, the differences are not significant. A summary of the data on size and proportions for the frogs of the genus Syrrhophus is given in Tables 4, 5, and 6. Hands and Feet.—Taylor and Smith (1945), Smith and Taylor (1948), Firschein (1954) and Duellman (1958) discussed the value of the palmar tubercles in identifying frogs of this genus. The eastern complex in general has a well-developed outer palmar tubercle (Fig. 1) in distinction to the western complex in which the outer palmar tubercle is reduced or absent (Fig. 2). Dixon and Webb (1966) imply that the outer palmar tubercle is rarely absent but is usually smaller than the first supernumerary tubercle of the fourth finger. My study of the western species demonstrates that the outer palmar tubercle is indeed usually present and smaller than the first supernumerary tubercle. Differences in interpretation of the terms "unexpanded" and "narrow," as well as differences in techniques of preservation, have led to confusion of the reported digital shapes in various species. Constant specific differences are evident in the hands (Fig. 1). Except in the cases of excessive uptake of fluids, all species have a terminal transverse groove at the tip of each digit. Taylor (1940b) stated that S. smithi lacked grooves, but examination of the holotype reveals faint grooves at the tops of the digits. Syrrhophus guttilatus, leprus, pipilans, and verrucipes lack lateral fringes on the fingers. Lateral fringes are well developed in the longipes and modestus groups but poorly defined or absent in the other members of the genus. The digital pads of the frogs of the longipes group are much broader than those of the other species and are narrowest in the frogs of the leprus group. Supernumerary tubercles are present on the palmar surfaces of all species of the genus. Table 4—Size and Proportions in the Frogs of the Syrrhophus leprus Group. Species Sex N Snout-vent length (SVL) Tibia length/ SVL Head width/ SVL Tympanum/ Eye Eyelid/ Interorbital cystignathoides campi ♂ 33 16.3-23.5 41.3-49.6 34.0-40.1 43.7-66.5 43.2-89.6 (45.8) (37.0) (56.2) (61.5) ♀ 12 16.0-25.8 41.5-51.0 33.0-38.0 42.8-60.0 48.2-69.2 (45.8) (35.0) (51.2) (60.1) c. cystignathoides ♂ 15 16.8-22.1 45.1-50.4 33.2-40.7 44.3-68.7 44.6-65.4 (47.3) (37.8) (54.8) (60.0) ♀ 6 19.6-24.2 46.4-50.0 34.1-38.1 43.3-56.5 53.2-65.4 (47.6) (36.2) (46.9) (59.2) leprus ♂ 14 20.6-26.4 42.3-52.3 35.0-40.3 47.5-62.5 58.2-72.5 (46.8) (37.4) (56.5) (67.3) ♀ 15 22.1-29.2 43.4-53.3 32.6-38.9 38.6-57.9 50.2-86.9 (47.1) (35.8) (47.1) (68.1) rubrimaculatus ♂ 12 18.2-23.5 40.4-46.2 31.8-35.5 35.5-46.5 65.1-78.5 (43.4) (33.8) (41.7) (71.7) Table 5—Size and Proportions in the Frogs of the Syrrhophus longipes and S. marnockii Groups. [Pg 7] [Pg 8] Species Sex N Snout-vent length (SVL) Tibia length/ SVL Head width/ SVL Tympanum/ Eye Eyelid/ Interorbital dennisi ♂ 16 22.8-28.4 43.9-49.7 35.3-41.2 53.9-64.2 55.3-74.0 (47.4) (38.8) (58.9) (65.1) ♀ 10 25.9-32.0 46.3-50.8 35.6-40.3 50.6-58.7 58.1-70.9 (48.2) (37.7) (54.9) (63.6) longipes ♂ 22 22.1-33.2 45.8-51.7 38.7-44.4 61.1-87.2 61.5-83.0 (48.4) (41.8) (72.0) (72.0) ♀ 19 26.8-39.6 44.3-51.0 36.3-40.8 49.5-72.1 55.3-85.9 (47.2) (39.1) (59.5) (67.9) guttilatus ♂ 19 20.6-29.0 41.2-48.1 36.9-44.9 55.1-75.7 53.3-79.5 (44.5) (40.6) (64.1) (66.0) ♀ 5 25.7-31.0 41.4-46.8 35.9-42.3 47.6-61.7 62.3-79.8 (43.6) (38.5) (54.0) (72.9) marnockii ♂ 14 18.4-28.9 42.3-47.2 36.1-43.0 47.2-68.3 51.6-74.4 (44.1) (39.6) (61.2) (66.3) ♀ 29 20.4-35.4 38.7-46.4 35.9-41.3 45.8-73.3 52.1-70.5 (42.7) (38.2) (60.3) (60.7) verrucipes ♂ 29 17.5-29.2 42.7-49.5 36.2-42.4 56.1-82.2 56.8-82.8 (46.3) (39.1) (67.8) (70.4) ♀ 6 26.5-31.7 42.4-47.7 36.0-38.1 45.8-57.8 61.0-77.9 (44.6) (37.0) (53.9) (69.0) Table 6—Size and Proportions in the Frogs of the Syrrhophus pipilans and S. modestus Groups. Species Sex N Snout-vent length (SVL) Tibia length/ SVL Head width/ SVL Tympanum/ Eye Eyelid/ Interorbital pipilans nebulosus ♂ 17 22.9-28.5 38.1-42.0 34.4-37.2 36.6-47.8 56.1-82.4 ♀ ♀ ♀ (40.0) (35.4) (43.6) (68.2) ♀ 3 21.1-22.7 42.1-44.5 33.2-35.8 36.6-47.6 64.3-65.4 pipilans pipilans ♂ 18 22.6-27.8 37.9-44.0 32.2-36.5 38.0-54.0 56.1-79.5 (41.4) (33.0) (46.2) (67.3) ♀ 1 29.4 38.4 32.5 44.6 55.0 modestus ♂ 8 15.8-20.1 38.5-42.6 32.1-38.1 26.8-39.3 57.0-86.9 (40.6) (34.2) (31.5) (69.1) ♀ 1 18.5 44.2 36.0 24.0 52.1 pallidus ♂ 6 17.9-19.3 41.0-44.9 32.6-36.2 27.0-35.6 59.4-67.7 (43.4) (35.2) (30.9) (65.2) teretistes ♂ 18 19.2-23.2 41.5-45.3 32.5-36.4 28.6-43.8 51.2-75.0 (43.7) (34.0) (33.7) (62.2) ♀ 1 24.8 41.8 30.8 37.9 60.5 nivocolimae ♂ 15 18.9-21.1 42.2-48.6 30.9-37.1 30.0-39.3 42.6-69.1 (45.0) (33.7) (34.7) (55.0) ♀ 1 24.1 40.9 33.5 27.6 56.5 interorbitalis ♂ 1 25.6 43.0 —— 39.4 57.6 ♀ 9 20.2-26.7 39.9-47.1 32.6-39.3 29.1-41.2 58.2-76.9 (43.2) (35.8) (36.4) (69.2) [Pg 9] Fig. 1: Palmar views of hands of six species of the eastern complex of Syrrhophus. (A) verrucipes (UIMNH 15995), (B) rubrimaculatus (KU 58911), (C) dennisi sp. nov. (holotype, UMMZ 101121), (D) guttilatus (UIMNH 55520), (E) marnockii (TCWC 4782), and (F) longipes (TCWC 12179). All ×6.5. Fig. 2: Palmar views of hands of two species of the western complex of Syrrhophus. pipilans (left, KU 58908, ×6) and teretistes (center, KU 75269, and right, KU 75263, respectively, ×9). In S. cystignathoides and leprus, the first finger is longer than the second, and the first two fingers are equal in length in guttilatus and marnockii. In the other species the first finger is shorter than the second. Supernumerary tubercles are well developed on the plantar surfaces in all species, except S. guttilatus, in which they are poorly defined (Fig. 3). The relative sizes of the metatarsal tubercles has been used in the classification of the species and species groups of Syrrhophus. The metatarsal tubercles are similar in all species of the eastern complex (including rubrimaculatus); the outer tubercle is always about one-half the size of the ovoid inner metatarsal tubercle. In the leprus group the outer tubercle is conical and compressed. The metatarsal tubercles of pipilans are about the same size, or the outer is slightly smaller than the inner. In the modestus group the outer metatarsal tubercle is about one-third the size of the inner. All species, except guttilatus, have well-defined to poorly defined lateral fringes on the toes. All species have [Pg 10] expanded toe pads. The fifth toe is usually shorter than the third, but the second is equal in length to the fifth in some specimens of S. cystignathoides and S. marnockii. Syrrhophus nivocolimae is the only species with tubercles along the outer edge of the tarsus; this is merely a reflection of the highly tuberculate nature of the skin in this species. Skin texture.—The skin of the dorsum is smooth or very weakly pustular in all species of the genus except nivocolimae and verrucipes. The dorsal surfaces of nivocolimae are warty; in verrucipes the skin is pustular. The skin of the venter is areolate in cystignathoides cystignathoides, dennisi and verrucipes but is smooth in all other species of the genus. Fig. 3: Plantar views of feet of four species of the eastern complex of Syrrhophus. (A) guttilatus (UIMNH 55519, ×6), (B) leprus (UIMNH 42726, ×6), (C) verrucipes (UIMNH 15995, ×6), and (D) longipes (TCWC 12179, ×4.6). Color pattern.—As is evident in the diagnoses, the color patterns of given populations have been regarded as useful in separating the species and subspecies. Duellman (1958) suggested that the coloration, with the exception of modestus, was a dark ground color with pale markings. It is a moot point whether the frogs have light spots on a dark background or have a light background with an extensive reticulate dark pattern. The venters are gray or white, and the vocal sac is nearly black in some species. Interorbital dark bars or triangles are absent in only two species of the eastern complex, cystignathoides campi and marnockii; the latter lacks a supratympanic stripe, which is present in the other members of the eastern complex. Syrrhophus interorbitalis and nivocolimae have light interorbital bars; these bars occur in only one other population of the genus (S. c. cystignathoides). Bars on the thighs are ill defined or absent in the members of the marnockii and part of the modestus groups. The color in life is noted in the species accounts. 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." [Pg 11] [Pg 12] 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 [Pg 13] [Pg 14] 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. [Pg 15] 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) and S. 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. 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[** 1946:170]. [Pg 16] [Pg 17] Fig. 5: Distribution of Syrrhophus cystignathoides campi (solid symbols) and the nominate subspecies (open symbols). Diagnosis.—Venter areolate; interorbital light and dark bars present; ground color yellow to brownish-yellow in life (Fig. 4b). Remarks.—Firschein (1954) briefly considered the status of Peters' (1871) Phyllobates verruculatus and noted that if it was a Syrrhophus it would probably be referrable to S. cystignathoides. Peters' (1871) original description corresponds well with S. cystignathoides, and the type-locality ("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., [Pg 17] [Pg 18] 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. [Pg 19] Fig. 6: Dorsal views of Syrrhophus leprus showing variation in dorsal pattern (left, UMMZ 121244, ×2; right, KU 26106, ×1.7). Side of head (UIMNH 42726, ×7). Fig. 7: Distribution of three species of eastern complex Syrrhophus: leprus (circles), rubrimaculatus (triangles), and verrucipes (squares). Remarks.—My distribution map (Fig. 7) differs somewhat from that of Duellman (1958), who was unaware of specimens reported by Taylor and Smith (1945) from central Veracruz, 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. [Pg 20] [Pg 21]