94.1 AMPHIBIA: SALIENTIA: BUFONIDAE VALLICEPS

Catalogue of Americau and Reptiles• • DEFINITIONANDDIAGNOSIS.A member of the tropical and subtropical Bujo valliceps group (Blair, 1966) with PORTER, KENNETH R. 1970. Bujo valliceps. strongly developed cranial crests, that produce a deep con• cavity down the center of the head, and a linear series of Bufo valliceps Wiegmann prominent warts on each side of the body bounded above by Gulf Coast a light stripe and below by a dark band; a light vertebral stripe is usually present; breeding males have a yellow-green Bujo valliceps Wiegmann, 1833:658. Type-localities, "Mexico" throat. The parotoid glands are relatively small and sub• (Zool. Mus. Berlin 3525-3527) and "Vera Cruz, Mexico" triangular to ellipsoid. Body size varies geographically, (ZMB 3532); restricted to Veracruz, Veracruz, Mexico medium in the north (females to 130 mm snout-vent), smaller (Smith and Taylor, 1950). Syntypes (4 females, 1 male) in southern range (females to 75 mm snout-vent); males are collected by Ferdinand Deppe (not seen by author). smaller than females. The mating call is a trill with a carrier Bujo trachypus Weigmann, 1833:657, 658, footnote. Apparently frequency that varies from about 1400 cps in the north to the name under which specimens of B. valliceps were sold about 2100 cps in the south; its pulse rate changes about 2 by the brother of Ferdinand Deppe and a nomen nudum. notes per see per degree centigrade change in body tempera• Bujo granulosus Baird and Girard, 1852:173 (not of Spix, ture and is about 38 notes per see at 25°C (30-46); call 1824). Type-locality, "between Indianola and San An• length is variable (1-7 see) but averages about 4 seconds. tonio," Texas; restricted to San Antonio, Bexar County, The dorsum of the tadpole tail musculature has 8-10 black Texas (Smith and Taylor, 1950). Holotype (male), U. S. bars with intervening areas of pale buff and there is an Natl. Mus. 2595, collected by John H. Clark (examined irregular black or brown band on the tail. by author). The combination of well-developed parietal cranial crests, Bujo nebulijer Girard, 1854:87. Replacement name for a linear series of prominent warts on each side of the body granulosus Baird and Girard which is preoccupied. bounded above by a light stripe and below by a dark band, and a light vertebral stripe will distinguish Bujo valliceps Bujo sternosignatus (part) Gunther, 1858:68, 69. Type• from any species with which it is sympatric. The similar localities, "Cordova" (British Mus. 56.3.17.25) and "Mex• Bujo gemmijer and B. mazatlanensis occur only on Pacific ico" (BM 58.9.6.13). Syntypes (2 juveniles), BM drainages of central and northern Mexico, are allopatric with 56.3.17.25, collected by Auguste Salle, and 58.9.6.13, pur• B. valliceps, and have poorly developed parietal crests. Bujo chased from Emile Parzudaki, a dealer at Paris (not seen cavijrons, whose range possibly overlaps that of B. valliceps by author). at higher elevations in Mexico, lacks a distinct light stripe on Chilophryne nebulijera: Cope, 1862:357. Emendation of the side of the body and has more elongated parotoid glands. nebulijer and transfer to genus Chilophryne. Bujo leutkeni, possibly sympatric with B. valliceps in Middle lncilius nebulijer: Cope, 1863:50. Transfer of nebulijer to America, has weak parietal crests and lacks a vertebral stripe. genus . Bujo coccijer is sympatric with B. valliceps in Mexico and Bujo valliceps valliceps: Werner, 1896:350. First use of tri• probably in Middle America but differs in having very short nomial; this combination implied by reference to "Type" parietal crests, more spinous warts, spherical parotoid glands, in description of micro tis. and lacking the linear series of warts on each side of the body. Bujo valliceps micro tis Werner, 1896:350. Type-locality, "Hon• duras." Holotype, Zool. Mus. Berlin 13200, collected by S. Schluter. Bujo valliceps macrocristatus Firschein and Smith, 1957:219. Type-locality, "between La Gloria and Cerro Azul," Oaxaca, Mexico. Holotype, Univ. Illinois Mus. Nat. Hist. 35583, collected by Thomas MacDougall, 24-26 March 1950. Bujo valliceps wilsoni Baylor and Stuart, 1961:195. Type• locality, "Jacaltenango (ca. 50 air-line kilometers north• west of Juehuetenango), Huehuetenango, Guatemala." Holotype, Univ. Michigan Mus. Zool. 119391, collected by L. C. Stuart on 6 June 1955.

• CONTENT.No subspecies are recognized, see Comment.

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6

2

o .2 .4 .6 .8 1.0 1.2 1.4 1.6 TIME IN SECONDS

FIGURE. Audiospectrogram (narrow band, 45 cycles per K?O 290 3qO 4C?O 590 second on left; wide band, 300 cycles per second on right) b 2bo' 400 . sOo . abo of part of mating call of Bujo valliceps: 7 mi. W Rincon, Veracruz, Mexico, 13 June 1964, body temperature 24°C MAP. The solid symbol marks the type locality, open symbols (K. R. Porter recording). indicate other records. The star marks a fossil locality. 94.2

Bulo ibarrai tends to replace B. valliceps at higher elevations were studied by Blair (953) and Thornton (960). Inter• in Guatemala and differs in lacking lateral body stripes and specific mating and hybridization are discussed in Orton in having very low parietal crests and generally more spinous (1951), Liner (954), Thornton (955), Volpe 0956, 1959, warts. Both B. cocciler and B. ibarrai tend to have very 1960), and Blair 0956, 1959, 1961b, 1963, 1964, 1966). narrow vertebral lines rather than stripes as in B. valliceps. Gratzer and Allison (960) mention the haemoglobin of The enigmatic Bulo cristatus, known only from Jalapa, Vera• B. valliceps. Chu and McCain (969) studied learning in cruz, and Tezuitlan, Puebla, differs from B. valliceps in having B. valliceps. much larger parotoid glands (20% of snout-vent length) and bypertrophied parietal and postorbital crests that fill the • ETYMOLOGY.The name valliceps, from Latin vallis (hol· space between the parotoid gland and eye. The population low) and caput (head), refers to the median dorsal longi• of B. valliceps from the Cuautlapan region of Veracruz also tudinal concavity of the head produced by the high cranial crests. has hypertrophied parietal crests but the postorbital crests are not expanded nor are the parotoid glands swollen. Other COMMENT than the above mentioned species, no other sympatric toad is likely to be confusedwith Bulo valliceps. A thorough study of geographical variation in B. valliceps is lacking and, consequently, the evaluation of subspecies is • DESCRIPTIONS.General accounts are in Dickerson (906), impossible. With the possible exception of wilsoni, a rela• Wright and Wright (949), and Conant (958); details of tively local and slightly.differentiated population, none of the adult morphology are in Kellogg (932) and Porter 0964a, named subspecies is well-defined morphologically or geo· 1964b). Osteological descriptions are in Baldauf (958), graphically. The validity of macrocristatus as a populational Sanders (961), and Tihen (962). A detailed description of entity is doubtful (Duellman 1960, Baylor and Stuart 1961, tadpole developmentis in Limbaugh and Volpe (957). Mat• Porter 1964a). The type·locality of microtis is not well· ing call parameters and variation are in Porter 0964b); the defined and specimens from Honduras will key to valliceps vocal apparatus is described by McAlister (961) . Venom (Lynch and Fugler, 1965). constituents are described by Chen and Chen (933), Barbier et al. (961), Porter 0964c), and Wittliff (964). LITERATURECITED • ILLUSTRATIONS.Photographs of adults are in Dickerson Baird, Spencer F. 1859. Reptiles of the boundary. In Report of the United States and Mexican Boundary (906), Wright and Wright (949), Conant (958), Cochran (1961), and Porter 0964a). The dorsal surface of the head Survey.... , Pt. II: 1-35, pIs. 1-41. is photographed in Baylor and Stuart (961); photographs of -, and Charles Girard. 1852. Characteristics of some new skulls are in Sanders (961) and Tihen (962). A lithograph reptiles in the museum of the Smithsonian Institution. of dorsal and ventral body surfaces is in Baird (859); draw• Proc. Acad. Nat. Sci. Philadelphia 6:68-70, 173. ings of osteologicalfeatures are in Baldauf (958). Excellent Baldauf, Richard J. 1958. Contributions to the cranial drawings of all developmental stages through metamorphosis morphology of Bulo valliceps Wiegmann. Texas J. Sci. are in Limbaugh and Volpe (957). A drawing of tadpole 10(2) :172-186. mouth parts is in Wright (929). Barbier, M., M. Bharucha, K. K. Chen, V. Deulofeu, E. Iseli, Herb. Jager, M. Kotake, R. Rees, T. Reichstein, O. • DISTRIBUTION.Bulo valliceps occurs on both Atlantic and Schindler, and Ek. Weiss. 1961. Papierchromatograph• Pacific drainages of Middle America from Bebedero (Wett· ische Priifung weiterer Krotensekrete. Helvetica Chim. stein, 1934) and the Rio Frio (USNM specimen 19581, Acta 44(44) :362-367. exaIuined by author) in extreme northern Costa Rica through Baylor, Edward R., and L. C. Stuart. 1961. A new race of the Isthmus of Tehuantepec; it is widespread along lowlands Bulo valliceps from Guatemala. Proc. BioI. Soc. Wash• of the Gulf coast from the Isthmus of Tehuantepec to Louisiana ington 74:195-202. and as far north as Calion, Arkansas (Smith and Langebartel, Blair, Albert P. 1941. Variation, isolating mechanisms, and 1949); it is not present on the Pacific coast of Mexico west hydribization in certain . Genetics 26(July) :398-417. of the Isthmus of Tehuantepec nor on the Central Plateau; it is found primarily on lowlands but to nearly 1800 meters in Blair, W. Frank. 1953. Growth, dispersal, and age at sexual moist habitats. maturity of the Mexican toad (Bulo valliceps Wiegmann). Important distributional data are in Noble (918), Kellogg Copeia 1953(4):208-212. (932), Smith (938), Smith and Taylor (948), Brown 1956. Comparative survival of hybrid toads (B. wood· (950), Milstead, Mecham, and McClintock (950), Taylor housei X B. valliceps) in nature. Copeia 1956(4) :259• (953), Firschein and SIuith (957), Milstead (960), Porter 260. 0964a), and Lynch and Fugler (965). Numerous other 1959. Genetic compatibility and species groups in U. S. publications list collection localities. Habitat notes are in toads (Bulo). Texas J. Sci. 11(4) :427-453. Dickerson (906), Ruthven (912), Brown (950), Raun 1960. A breeding population of the Mexican toad (Bulo (959), Duellman 0960, 1963, 1965), Blair 0960, 1961a), valliceps) in relation to its environment. Ecology 410): Baylor and Stuart (961), Porter 0964a), and Reddell 165-174. (967) . 1961a. Calling and spawning seasons in a mixed popula• tion of anurans. Ibid. 420) :99-110. • FOSSILRECORD.Langebartel (953) lists B. valliceps remains from Yucatan cave deposits that are of questionable 1961b. Further evidence bearing on intergroup and intra• group genetic compatibility in toads (genus Bulo). Texas age from very recent to Pleistocene. No other fossils have J. Sci. 13(2) :16:>-175. been assigned to this species. 1963. Evolutionary relationships of North American toads • PERTINENTLITERATURE.Relationships to other species are of the genus Bulo: A progress report. Evolution 170): discussed in Blair - 0959, 1961b, 1963, 1964, 1966), Porter 1-16. 0964a, 1964b, 1964c), and Porter and Porter (967). Em· 1964. Evolution at populational and interpopulational bryonic temperature tolerance and rates of development are levels: Isolating mechanisms and interspecies interactions discussed in Limbaugh and Volpe (957), Volpe (957), and in anuran amphibians. Quart. Rev. BioI. 39(4) :33:>-344. Hubbs et al. (963). Brattstrom 0963, 1968) has provided 1966. Genetic compatibility in the Bulo valliceps and data on thermal requirements of adults. Notes on feeding closely related groups of toads. Texas J. Sci. 18(4): habits are in Strecker (937). Morphologicalvariation is dis· 33:>-351. cussed in Blair (941), Taylor and SIuith (945), Taylor Brattstrom, Bayard H. 1963. A preliIuinary review of the 0949, 1952), Duellman 0960, 1963), Baylor and Stuart thermal requirements of amphibians.. Ecology 44(2) :238• (961), and Porter 0964a, 1964b). Population dynamics 255. 94.3

1968. Thermal acclimation in anuran amphibians as a Noble, G. Kingsley. 1918. The amphibians collected by the function of latitude and altitude. Compo Biochem. Physiol. American Museum expedition to Nicaragua in 1916. Bull. 24(1) :93-111. Amer. Mus. Nat. Hist. 38:311-347. Brown, Bryce C. 1950. An annotated check list of the Orton, Grace L. 1951. An example of interspecific mating reptiles and amphibians of Texas" Baylor Univ. Press, in toads. Copeia 1951(1) :78. Waco, Texas, xii + 257 pp. Porter, Kenneth R. 1964a. Distribution and taxonomic status Chen, K. K., and A. L. Chen. 1933. A study of the poisonous of seven species of Mexican Bulo. Herpetologica 19(4): secretions of five North American species of toads. J. 229-247. Pharmacol. ExptI. Therap. 49(4) :526-542. 1964b. Morphological and mating call comparisons in the Chu, Philip K., and Garvin McCain. 1969. Discrimination Bulo val/ieeps complex. Amer Midland Nat. 71 (1) :232• learning and extinction in toads. Psychon. Sci. 14(1): 245. 14-15. 1964c. Chromatographic comparisons of the parotoid Cochran, Doris M. 1961. Living amphibians of the world. gland secretions of six species in the Bulo val/ieeps group. Doubleday and Company, Garden City, New York, 199 p. P. 451-456. In Charles A. Leone, Ed., Taxonomic bio• Conant, Roger. 1958. A field guide to reptiles and amphib• chemistry and serology. Ronald Press Co., New York. ians of the United States and Canada east of the 100th -, and Wendy F. Porter. 1967. Venom comparisons and Meridian. Houghton-Mifflin Company, Boston, xviii + relationships of twenty species of New World toads (genus 366 p. Bulo). Copeia 1967(2) :298-307. Cope, Edward Drinker. 1862. Catalogues of the reptiles Raun, Gerald G. 1959. Terrestrial and aquatic vertebrates obtained during the exploration of the Parana, Paraguay, of a moist relict area in central Texas. Texas J. Sci. Vermejo, and Uruguay, by Capt. Thos. J. Page, U.S.N.; 11(2) :158-171. and of those procured by Lieut. N. Michler, U.s. Top. Reddell, James R. 1967. A checklist of the cave fauna of Engr., Commander of the expedition conducting the survey Texas. III. Vertebrata. Texas J, Sci. 19(2) :184-226. of the Atrato River. Proc. Acad. Nat. Sci. Philadelphia Ruthven, Alexander G. 1912. The amphibians and reptiles 14:346-359. collected by the University of Michigan-Walker expedition 1863. On Traehyeephalus, Seaphiopus and other Ameri• in southern Vera Cruz, Mexico. ZooI. Jahr. 32(4) :295-332. can Batrachia. Ibid. 15:43-54. Sanders, OUys. 1961. Indications for the hybrid origin of Dickerson, Mary C. 1906. The book. Doubleday, Page, Bulo terrestris Bonnaterre. Herpetologica 17(3) :145-156. and Company, Garden City, New York, xvii + 253 p. Smith, Hobart M. 1938. Notes on reptiles and amphibians Duellman, William· E. 1960. A distributional study of the from Yucatan and Campeche, Mexico. Occ. Pap. Mus. amphibians of the Isthmus of Tehuantepec, Mexico. Univ. ZooI. Univ. Michigan 388:1-22. Kansas Mus. Nat. Hist. Publ. 13(2) :19-72. -, and Dave A. LangebarteI. 1949. The toad Bulo val/ieeps 1963. Amphibians and reptiles of the rainforests of in Arkansas. Copeia 1949(3) :230. southern EI Peten, Guatemala. Ibid. 15(5) :205-249. -, and Edward H. Taylor. 1948. An annotated checklist 1965. Amphibians and reptiles from the Yucatan Penin• and key to the Amphibia of Mexico. U. S. Nat!. Mus. Bull. sula, Mexico. Ibid. 15 (12) :577-614. (194) : ix + 118 pp. Firschein, I. Lester, and Hobart M. Smith. 1957. A high -, and -. 1950. Type localities of Mexican reptiles and crested race of toad (Bulo vallieeps) and other noteworthy amphibians. Univ. Kansas Sci. Bull. 33, Pt. 11(8) :313--380. reptiles and amphibians from southern Mexico. Herpe• Spix, Johann Baptist von. 1824. Animalia nova, species tologica 13(3) :219-222. novae testudinorum et ranarum quas in itinere per Girard, Charles. 1854. A list of the North American bu• Brasiliam annis 1817-1820 jussu et asupiciis Maximiliana fonids with diagnosis of new species. Proc. Acad. Nat. Josephi Z., 39 pIs. Leipzig. Sci. Philadelphia 7:86-88. Strecker, John K. 1927. Chapters from the life-histories of Gratzer, W. B., and A. C. Allison. 1960. Multiple haemo• Texas reptiles and amphibians. Part two. Contr. Baylor globins. BioI. Rev. 35:459-506. Univ. Mus. 10:3-14. Giinther, A. C. L. 1858. Catalogue of the Batrachia Salientia Taylor, Edward H. 1949. A preliminary account of the in the collection of the British Museum, xvi + 160 p. herpetology of the state of San Luis Potosi, Mexico. Univ. Hubbs, Clark, Thomas Wright, and Orlando Cuellar. 1963. Kansas Sci. Bull. 33, Pt. I, (2) :169-215. Developmental temperature tolerance of central Texas 1952. Third contribution to the herpetology of the Mexi• populations of two anuran amphibians Bulo val/ieeps and can state of San Luis Potosi. Ibid. 34(13) :793-815. Pseudaens streeken. Southwest. Nat. 8(3) :142-149. 1953. Fourth contribution to the herpetology of San Luis Kellogg, Remington. 1932. Mexican tailless amphibians in Potosi. Ibid. 35(2) :1587-1614. the United States National Museum. U. S. Nat!. Mus. Bull. -, and Hobart M. Smith. 1945. Summary of the collections (160) :1-224. of amphibians made in Mexico under the Walter Rathbone Langebartel, Dave A. 1953. The reptiles and amphibians. Bacon travelling scholarship. Proc. U. S. Nat!. Mus. 95 P. 91-108. In Faunal and Archeological Researches in (3185) :521-647. Yucatan Caves by R. T. Hatt, H. I. Fisher, D. A. Lange• Thornton, Wilmot A. 1955. Interspecific hybridization in bartel, and G. W. Brainerd. Cranbrook Inst. Sci. Bull. Bulo woodhousei and Bulo val/ieeps. Evolution 9(4): (33) . 455-468. Limbaugh, Beverly A., and E. Peter Volpe. 1957. Early 1960. Population dynamics in Bulo woodhousei and Bulo development of the Gulf coast toad, Bulo val/ieeps Wieg• val/ieeps. Texas J. Sci. 12(3/4) :176-200. mann. Amer. Mus. Novitates (1842) :1-32. Tihen, J. A. 1962. Osteological observations on New World Liner, E. A. 1954. The herpetofauna of Lafayette, Terre• Bulo. Amer. Midland Nat. 67(1) :157:-183. bonne and Vermilion Parishes, Louisiana. Proc. Louisiana Volpe, E. Peter. 1956. Experimental F. hybrids between Acad. Sci. 17:65-85. Bulo vallieeps and Bulo lowlen. Tulane Stud. Zool. 4(2) : Lynch, John D., and Charles M. Fugler. 1965. A survey of 61-75. the of Honduras. J. Ohio Herp. Soc. 5(1) :5-18. 1957. Embryonic temperature tolerance and rate of de• McAlister, Wayne H. 1961. The mechanics of sound produc• velopment in Bulo val/ieeps. Physiol. Zool. 30(2) :164-176. tion in North American Bulo. Copeia 1961(1) :86-95. 1959. Hybridization of Bulo vallieeps· with Bulo amen• Milstead, William W. 1960. Relict species of the Chihua• eanus and Bulo terrestns. Texas J. Sci. 11(3) :335-342. huan desert. Southwest. Nat. 5(2) :75-88. 1960. Evolutionary consequences of hybrid sterility and -, John S. Mecham, and Haskell McClintock. 1950. The vigor in toads. Evolution 14(2) :181-193. amphibians and reptiles of the Stockton Plateau in north• Werner, Franz. 1896. Beitrage zur Kenntniss der Reptilien ern Terrell County, Texas. Texas J. Sci. 2(4) :543-562. und Batrachier von Centralamerika und Chile, sowie einiger 94.4

seltenerer Schlangenarten. Verhandl. Zool.·Bot. Ges. Wien Wright, Albert Hazen, and Anna Allen Wright. 1949. Hand• 46:344-365. book of frogs and toads of the United States and Canada. Wettstein, Otto. 1934. Ergebnisse der osterreichischen bio• Comstock Publ. Company, Ithaca, New York, vii + 640 pp. logischen Costa Rica-Expedition 1930. Die Amphibien und Wright, Anna Allen. 1929. Synopsis and description of Reptilien. Sitzungsb. Akad, Wiss. Wien, Mathem.-Naturw. North American tadpoles. Proc. U. S. Natl. Mus. 74(11): 1-70. Kl. Abt. I, 143(1-2) :1-39. Wiegmann, A. F. A. 1833. Herpetologische Beitrage. Isis K. R. PORTER, UNIVERSITY OF DENVER, DENVER, COLORADO 26:651-662. 80210. Wittliff, James L. 1964. Venom constituents of Hulo lowleri, Published 11 September 1970 by the American Society of Hulo valliceps, and their natural hybrids analyzed by Ichthyologists and Herpetologists. Publication is supported by electrophoresis and chromatography. P. 457-464. In National Science Foundation grant G24231. Charles A. Leone, Ed., Taxonomic biochemistry and serology. Ronald Press Co., New York. Primary editor for this account, Richard G. Zweifel.