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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/31830-8.txt b/31830-8.txt new file mode 100644 index 0000000..91dae51 --- /dev/null +++ b/31830-8.txt @@ -0,0 +1,1062 @@ +Project Gutenberg's Genera of Leptodactylid Frogs in México, 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: Genera of Leptodactylid Frogs in México + +Author: John D. Lynch + +Release Date: March 30, 2010 [EBook #31830] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK LEPTODACTYLID FROGS IN MEXICO *** + + + + +Produced by Chris Curnow, Joseph Cooper, Diane Monico, and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS +MUSEUM OF NATURAL HISTORY + +Volume 17, No. 11, pp. 503-515, 5 figs. +March 20, 1968 + + + +Genera of Leptodactylid Frogs in México + + +BY + +JOHN D. LYNCH + + +UNIVERSITY OF KANSAS +LAWRENCE +1968 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Frank B. Cross + + +Volume 17, No. 11, pp. 503-515, 5 figs. +Published March 20, 1968 + + +UNIVERSITY OF KANSAS +Lawrence, Kansas + + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1968 + +31-9418 + + + + +Genera of Leptodactylid Frogs in México + +BY JOHN D. LYNCH + + +INTRODUCTION + +According to the most recent review of the Mexican amphibian fauna +(Smith and Taylor, 1948), six genera of leptodactylid frogs occur in +México. One other genus, _Pleurodema_, occurs in Lower Central America. +Smith and Taylor recognized one species of _Engystomops_, 28 of +_Eleutherodactylus_, three of _Leptodactylus_, eight of +_Microbatrachylus_, 12 of _Syrrhophus_, and five of _Tomodactylus_. +Subsequent to the publication of their checklist of the Mexican +amphibia (1948), numerous taxonomic changes have been proposed. Many +species of _Eleutherodactylus_ have been added to the fauna, either +through the extension of their recorded ranges into México from +Guatemala or by the recognition of species unknown in 1948, whereas +some nominal species have been synonymized. _Microbatrachylus_ has been +regarded as synonymous with _Eleutherodactylus_ (Lynch, 1965); four +species of _Microbatrachylus_ currently are regarded as valid +(Duellman, 1961, Lynch, 1965). _Syrrhophus_ was revised in part by +Duellman (1958) and Firschein (1954), and a species of _Tomodactylus_ +transferred to _Syrrhophus_ by Dixon (1957), who redefined +_Tomodactylus_ and added more species to the genus. + +Since beginning my studies of the Mexican leptodactylids in 1962, I +have become acutely aware of difficulties involved in defining the +genera. A revision of _Eleutherodactylus_ and a review of _Syrrhophus_ +are nearing completion, but prior to their publication it is desirable +to redefine the genera of the Mexican leptodactylids, and in so doing +recognize an heretofore unnamed genus. The definitions of +_Eleutherodactylus_ and _Leptodactylus_ may need to be altered in the +future, since both are widespread in South America and occur in the +West Indies. Their definitions as given here are as precise as present +knowledge permits. _Syrrhophus_ and _Tomodactylus_ are small +assemblages that occur only in southwestern United States, México, and +Guatemala. + +Taylor (1952) synonymized _Engystomops_ with _Eupemphix_ which, +although related, should be regarded as generically distinct (Gallardo, +1965). Perhaps the most conservative classification is that of Myers +(1962) who, without published evidence, combined _Eleutherodactylus_, +_Syrrhophus_, and the South American _Lithodytes_ in a single genus. + +The major problem for students working with the Mexican leptodactylids +has not been the separation of _Engystomops_ or _Leptodactylus_ from +other genera but the separation and definition of the eleutherodactyline +frogs currently placed in three genera, _Eleutherodactylus_, +_Syrrhophus_, and _Tomodactylus_. As will be shown in this paper, these +are more conveniently placed in four genera. Once a fourth genus is +recognized, certain phylogenetic problems disappear and a reasonable +zoogeographic interpretation is possible for Middle American +leptodactylid distribution. + + + + +ANALYSIS OF CHARACTERS + + +In México and northern Central America approximately 55 species of +eleutherodactyline frogs (_Eleutherodactylus_, _Syrrhophus_, and +_Tomodactylus_) are known. Four genera can be recognized on the basis +of the nature of inguinal glands, morphology of the hands and feet, and +certain osteological features. + +[Illustration: FIG. 1. _Tomodactylus angustidigitorum_ (UMMZ 114305, +× 4.5) illustrating the lumbo-inguinal gland typical of members of the +genus. From a kodachrome by Wm. E. Duellman.] + + +Glands + +Leptodactylids have a variety of glands that have been used as generic +characters. Smith and Taylor (1948) regarded the so-called inguinal +gland as a generic character in Mexican eleutherodaycty-lines. Lynch +(1965) showed that _Eleutherodactylus_ and _Microbatrachylus_ cannot +be separated by the nature of the gland or the condition of the +prevomers (dentate or not). _Syrrhophus_ and _Tomodactylus_, as defined +by Smith and Taylor (1948), are not generically distinct because of +overlap in the condition of the prevomers and in the development of the +gland. Firschein (1954) stated that _Syrrhophus_ differed from +_Tomodactylus_ by having an axillary gland, but it is now known that +one species of _Syrrhophus_ lacks the gland. + +The inguinal glands of _Eleutherodactylus_ and _Syrrhophus_, if +present, are diffuse, irregular in outline, and generally not +prominent; in _Tomodactylus_ the gland is higher on the body (a +lumbo-inguinal gland), compact, oval in outline, and prominent (Fig. +1). Axillary glands occur in most _Syrrhophus_ but are not known in +_Tomodactylus_ or _Eleutherodactylus_. + + +Hands and feet + +The tips of the digits are laterally expanded in most +_Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_. Two species +of _Eleutherodactylus_ (_augusti_ and _tarahumarensis_) and two +_Tomodactylus_ (_angustidigitorum_ and _grandis_) lack any expansion +of the digital tips. All but two of the species of eleutherodactyline +frogs (_E. augusti_ and _E. tarahumarensis_) have a transverse groove +across the tips of the digits (Fig. 2). + +[Illustration: FIG. 2. Palmar views of the hands and lateral views of +the tip of the third digits of _Eleutherodactylus alfredi_ (left, KU +93994, × 5) and _Hylactophryne augusti_ (right, KU 102594, × 3).] + +Supernumerary tubercles rarely are present on the feet of +_Eleutherodactylus_, but are present and numerous in every species of +_Syrrhophus_, _Tomodactylus_, and in the members of the _augusti_ group +of _Eleutherodactylus_ (Fig. 3). The tubercles are small and numerous +in _Syrrhophus_ and larger in _Tomodactylus_ and the _Eleutherodactylus +augusti_ group. Most species of _Eleutherodactylus_ have no plantar +supernumerary tubercles; a few species have such tubercles, which never +extend between the metatarsal tubercles as in _Syrrhophus_ and +_Tomodactylus_. + +[Illustration: FIG. 3. Plantar views of feet of _Eleutherodactylus +alfredi_ (left, KU 93994, × 4.5), _Syrrhophus pipilans nebulosus_ +(middle, KU 58900, × 7.5), and _Hylactophryne augusti_ (right, KU +102594, × 3) showing differences in size and arrangement of +supernumerary tubercles.] + +Tarsal folds and tubercles are lacking in _Syrrhophus_, _Tomodactylus_, +and the _augusti_ group of _Eleutherodactylus_. Several species of +_Eleutherodactylus_ lack tarsal folds and tubercles, but in nearly +every species group, one or more species possess either an inner tarsal +fold, inner tarsal tubercle(s), or outer tarsal tubercles. + +The terminal phalanges of _Syrrhophus_, _Tomodactylus_, and all +_Eleutherodactylus_ (except the frogs of the _augusti_ group) are +distinctly T-shaped. In the latter, the bones are knob-shaped distally +(Fig. 4). T-shaped terminal phalanges also are present in _Lithodytes_ +and _Trachyphrynus_ but not in other leptodactylid genera. At least one +species of _Eupsophus_ (_E. quixensis_) has terminal phalanges that +resemble those of the _Eleutherodactylus augusti_ group. Several +species of _Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_ with +slender fingers have T-shaped terminal phalanges although the terminal +dilations proportionately are only scarcely wider than the finger tips +in the _Eleutherodactylus augusti_ group. The presence of a terminal +groove at the tip of the finger is an external indicator of the +T-shaped terminal phalanges. + +[Illustration: FIG. 4. Terminal phalanges of four leptodactylid frogs +(all × 13.5). (a) _Eleutherodactylus mexicanus_, KU 55593; (b) +_Eupsophus roseus_, KU 84731; (c) _Eupsophus quixensis_, UIMNH 59643; +and (d) _Hylactophryne augusti_, KU 56192.] + + +Skull + +All Mexican eleutherodactyline frogs have quadratojugal-maxillary +articulations, completely roofed skulls in adults, median contact of +the nasals, separated occipital condyles, and large prevomers. The +premaxillae of all species are visible when the skulls are viewed from +directly above. The pterygoid lacks a medioventral flange and does not +meet the palatine. In no species is the anterior arm of the squamosal +in contact with the maxillary. Of the numerous species examined (30 +_Eleutherodactylus_, four _Syrrhophus_, and four _Tomodactylus_), the +species in the _Eleutherodactylus augusti_ group are unique in having a +sphenethmoid with a blunt anterior edge. + + +Pectoral Girdle + +All species have large cartilaginous plates in the pectoral girdles; +none possesses a bony style. No divergent modifications of the clavicle +and coracoid bones are known in the family. + + + + +GENERIC ACCOUNTS + + +Genus ~Eleutherodactylus~ Dumeril and Bibron, 1841 + + _Type-species._--_Hylodes martinicensis_ Tschudi, 1838 + + _Diagnosis and definition._--Small to large frogs (12 to 110 + mm. snout-vent length) having slightly to widely expanded + digital pads, each pad bearing a terminal transverse groove; + lumbo-inguinal, inguinal, and axillary glands absent, or if + present, diffuse, irregular in outline, not compact; plantar + supernumerary tubercles absent, or if present, six or fewer, + restricted to distal area of plantar surface, and not + extending between metatarsal tubercles; tarsus bearing inner + or outer tubercles or folds or not; toes free to one-half + webbed; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + contact; anterior arm of squamosal not in contact with + maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + dentigerous processes present or not, dentate or not; + maxillary and premaxillary bones dentate; occipital condyles + separated; development direct. + + _Composition._--About 420 names have been applied to frogs + of this genus; many of these names are synonyms, and many + other species remain undescribed and unnamed. Perhaps the + genus contains 350 species. Thirty-one species occur in + México and northern Central America. + + _Distribution._--From Tamaulipas and Sinaloa, México, + exclusive of the Mexican Plateau, to at least Peru and + southernmost Brazil and throughout the West Indies. + Introduced into Florida. + + _Etymology._--Greek (_eleuthero_ + _dactylus_) meaning + free-toed. + + +Genus ~Engystomops~ Jiménez de la Espada, 1872 + + _Type species._--_Engystomops petersi_ Jiménez de la Espada, + 1872 + + _Diagnosis and definition._--Small frogs (20 to 40 mm. + snout-vent length) having undilated digital tips lacking + transverse grooves; lumbo-inguinal or inguinal glands + absent; plantar supernumerary tubercles present, extending + between metatarsal tubercles; tarsus bearing spinelike + tubercle on inner edge; toes free; terminal phalanges + pointed; sternum bearing bony style; spenethmoid not + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers moderate in size, lacking teeth; + maxillary and premaxillary bones edentate; occipital + condyles separated; tadpole free living. + + _Composition._--Four nominal species (_E. petersi_, _E. + pustulatus_, _E. pustulosus_ and _E. schereri_). + + _Distribution._--Central Veracruz and eastern Oaxaca, + México, to Trinidad, Bolivia, and Peru, east of the Andes. + + _Etymology._--Greek (_engys_ + _stoma_) meaning narrow-mouthed. + + +Genus ~Hylactophryne~ new genus + + _Type-species._--_Hylodes augusti_ Dugés, 1879 + + _Diagnosis and definition._--Medium to large frogs (37 to 94 + mm. snout-vent length) having undilated digital tips lacking + terminal grooves; lumbo-inguinal or inguinal glands absent; + plantar supernumerary tubercles present, prominent, + extending to but not between metatarsal tubercles; tarsus + lacking tubercles or folds; toes free of webbing; terminal + phalanges knob-shaped, lacking elongate lateral expansions; + sternum cartilaginous, lacking bony style; sphenethmoid + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers large, bearing dentigerous + processes; maxillary and premaxillary bones dentate; + occipital condyles separated; development direct. + + _Composition._--Two species, _H. augusti_ and _H. + tarahumarensis_, the former composed of four subspecies + (Zweifel, 1956). + + _Distribution._--From Arizona, New Mexico, and Texas to + Guerrero and Puebla, México, and a relict population on + Cerro Quingola (just west of the Isthmus of Tehuantepec, + México). + + _Etymology._--Greek (_hylactor_ + _phryne_) meaning barking + toad; in reference to the voice and common name. + + +Genus ~Leptodactylus~ Fitzinger, 1826 + + _Type-species._--_Leptodactylus typhonia_ Fitzinger, 1826 + + _Diagnosis and definition._--Small to large frogs (30 to + about 200 m., snout-vent length) having undilated to + slightly expanded digital tips bearing pads, no transverse + groove at tips of digits; lumbo-inguinal, axillary, and/or + ventral glands present or not, low, diffuse; plantar + supernumerary tubercles generally absent, if present not + extending between metatarsal tubercles; tarsus bearing + tarsal folds or not; toes free of webbing, extensive lateral + fringes present in some species; terminal phalanges pointed, + not T-shaped; sternum bearing bony style; sphenethmoid not + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers large, bearing dentigerous + processes; maxillary and premaxillary bones dentate; + occipital condyles separated; tadpole free living. + + _Composition._--Sixty species according to Smith and Taylor + (1948); 54 according to Gorham (1963); Argentinian authors + have described several more in recent years. + + _Distribution._--Southern Sonora, México, and southern Texas + throughout the Central and South American lowlands to + Argentina. Also known from Hispaniola and Puerto Rico in the + Greater Antilles and a few islands in the Lesser Antilles. + + _Entymology._--Greek (_leptos_ + _dactylus_) meaning slender + toes. + + +Genus ~Syrrhophus~ Cope, 1878 + + _Type-species._--_Syrrhophus marnockii_ Cope, 1878 + + _Diagnosis and definition._--Small to medium sized frogs (18 + to 40 mm. snout-vent) having slight to prominent digital + expansions with transverse groove at tip of each digit; + lumbo-inguinal and inguinal gland flattened, irregular in + outline, not compact and oval; axillary glands present or + not; plantar supernumerary tubercles numerous, more than + eight, usually extending between metatarsal tubercles; + tarsus lacking tubercles or folds; toes free or basally + webbed; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + articular contact; anterior arm of squamosal not in contact + with maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + usually lacking dentigerous processes and teeth; maxillary + and premaxillary bones dentate; occipital condyles + separated; development direct. + + _Composition._--Thirteen species; the species described as, + or later referred to, _Syrrhophus_ from Lower Central + America and South America are _Eleutherodactylus_ or + _Eupsophus_. + + _Distribution._--Low to moderate elevations from Sinaloa, + México, to Guatemala on the Pacific versant; from the + Edwards and Stockton plateaus of Texas to British Honduras + on the Caribbean versant. + + _Etymology._--Greek, emendation of _syrrhaptos_, meaning + sewn together in reference to the united outer metatarsals. + + +Genus ~Tomodactylus~ Günther, 1900 + + _Type-species._--_Tomodactylus amulae_ Günther, 1900. + + _Diagnosis and definition._--Small frogs (20 to 35 mm. + snout-vent length) having digital expansions or not, with + transverse groove across tip of each digit; lumbo-inguinal + gland prominently elevated, compact, oval, often patterned; + axillary glands absent; plantar supernumerary tubercles + numerous, more than eight, usually extending between + metatarsal tubercles; tarsus lacking tubercles or folds; toes + free; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + articular contact; anterior arm of squamosal not in contact + with maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + usually bearing dentigerous processes; maxillary and + premaxillary bones dentate; occipital condyles separated; + development direct. + + _Composition._--Ten species. + + _Distribution._--The southern edge of the Mexican Plateau + from Sinaloa to Veracruuz and onto the Oaxaca highlands and + Sierra Madre del Sur. + + _Etymology._--Greek (_tomis_ + _dactylus_) meaning knife + toe; in reference to either the sharp subarticular tubercles + or the unwebbed toes. + + + + +DISCUSSION + + +The preceding definitions only slightly alter the present generic +limits of Mexican leptodactylids. Two species, previously regarded as +_Eleutherodactylus_, are transferred to the new genus _Hylactophryne_. +The arrangement of the species of _Syrrhophus_ and _Tomodactylus_ +remains the same as concluded by Dixon (1957), Duellman (1958), and +Firschein (1954) in their reviews of the genera. + +Lumbo-inguinal glands are most prominent in the genera _Pleurodema_ and +_Tomodactylus_. Various nondescript glands are present in many genera, +but none is so well developed as those of _Pleurodema_ and +_Tomodactylus_. + +At least nine leptodactylid genera are either known or thought to be +terrestrial breeders lacking a free-living tadpole stage +(_Eleutherodactylus_, _Euparkerella_, _Hylactophryne_, _Niceforonia_, +_Noblella_, _Sminthillus_, _Syrrhophus_, _Tomodactylus_ and +_Trachyphrynus_). _Niceforonia_ and _Trachyphrynus_, and probably +_Hylactophryne_, are not closely related to the other genera. Direct +development probably is an adaptation to adverse environmental +conditions since many of the species occur in semi-arid or cold (Andean +páramos) areas. _Eleutherodactylus_ is generally thought to be the +stock from which _Euparkerella_, _Noblella_, and _Sminthillus_ evolved +(Griffiths, 1959) and from which _Syrrhophus_ and _Tomodactylus_ are +derived (Firschein, 1954). + +The present distribution of _Hylactophryne_ (isolated on the Mexican +Plateau) and its digital form (like that of Papuan and many primitive +South American leptodactylids) suggest that the genus was isolated in +México throughout the Tertiary, whereas the other Central American +genera are either post-Pliocene derivatives of _Eleutherodactylus_ or +invaders of Central America from South America since the mid-Pliocene +land bridge was formed (Lloyd, 1963). + +Piatt (1934) presented arguments against assigning _Eleutherodactylus +latrans_ to the genus _Lithodytes_ and concluded that it was a "true" +_Eleutherodactylus_. Contrary to his arguments, _latrans_ (= _augusti_ +of Zweifel) and _E. tarahumarensis_ Taylor differ from all other +_Eleutherodactylus_ (and _Syrrhophus_ and _Tomodactylus_) in the nature +of the tips of the digits (external and skeletal). The digits of +_Hylactophryne_ are like those of _Eupsophus_. My study of nearly all +genera of leptodactylids indicates that Noble (1925) was correct in +suggesting that _Borborocoetes_ (= _Eupsophus_) is a close relative of +_Eleutherodactylus latrans_, although Noble's arguments were based in +part upon false evidence concerning the breeding habits of _E. +latrans_, then thought to have a free-living tadpole. + +Kellogg (1932) and Piatt (1934) argued that the terminal phalanges of +_E. latrans_ were typically eleutherodactyline. The variation of this +character in _Eupsophus_ (see Fig. 4) ranges from knobbed to bifurcate +or Y-shaped (T-shaped in _Eleutherodactylus_, _Syrrhophus_ and +_Tomodactylus_) and encompasses the nature of the character represented +in _Hylactophryne_. _Eupsophus_ differs from _Hylactophryne_ in +possessing a frontoparietal fontanelle, in generally having a +maxillary-quadratojugal gap, and in having a free swimming tadpole +stage. + +[Illustration: FIG. 5. Outline drawings of _Leptodactylus melanonotus_ +(left, KU 65704, × 2) and _Eleutherodactylus alfredi_ (right, KU 93994, +× 2).] + + + + +KEY TO MEXICAN LEPTODACTYLID GENERA + + +1. Small (20-40 mm.), pustular, toadlike frogs; + maxillary and premaxillary bones not bearing teeth _Engystomops_ + + Large (20-110 mm.), smooth skinned and non-toadlike + frogs; maxillary and premaxillary bones bearing teeth 2 + +2. No conspicuous waist (Fig. 5); sternum bearing + bony style, _Leptodactylus_ + + Constrictions at waist (Fig. 5); sternum + cartilaginous, no bony style 3 + +3. Few (less than six), if any, supernumerary + tubercles on plantar surface _Eleutherodactylus_ + + Many (more than 8) supernumerary tubercles on + plantar surfaces 4 + +4. Terminal, transverse groove across tip of + digits, especially outer two fingers, digits + expanded or not; small frogs (18 to 40 mm.) 5 + + Tips of digits lacking transverse groove; + digits unexpanded; medium-sized to large frogs + (37 to 94 mm.) _Hylactophryne_ + +5. Lumbo-inguinal gland compact, oval _Tomodactylus_ + + Lumbo-inguinal or inguinal gland absent or + diffuse and irregular in outline _Syrrhophus_ + + + + +LITERATURE CITED + + +DIXON, J. R. + + 1957. Geographic variation and distribution of the genus + _Tomodactylus_ in Mexico. Texas Jour. Sci., 9:379-409, + December. + +DUELLMAN, W. E. + + 1958. A review of the frogs of the genus Syrrhophus in + western Mexico. Occas. Papers Mus. Zool. Univ. + Michigan, 594:1-15, June 6. + + 1961. The amphibians and reptiles of Michoacan, Mexico. Univ. + Kansas Publ. Mus. Nat. Hist., 15:1-148, December 20. + +FIRSCHEIN, I. L. + + 1954. Definition of some little understood members of the + leptodactylid genus _Syrrhophus_, with a description + of a new species. Copeia, 1:48-58, February 19. + +GALLARDO, J. M. + + 1965. A proposito de los Leptodactylidae (Amphibia Anura). + Papeis Avulsos, 17:77-87, January 30. + +GORHAM, S. W. + + 1963. The comparative number of species of amphibians in + Canada and other countries. III. Summary of species + of anurans. Canadian Field-Nat., 77:13-48, March. + +GRIFFITHS, I. + + 1959. The phylogeny of Sminthillus limbatus and the status of + the Brachycephalidae (Amphibia Salientia). Proc. Zool. + Soc. London, 132:457-87, May. + +KELLOGG, R. + + 1932. Mexican tailless amphibians in the United States + National Museum. Bull. U. S. Natl. Mus., 160:224 pp., + March 31. + +LLOYD, J. J. + + 1963. Tectonic history of the south Central-American orogen, + _in_ Childs and Beebe eds., Backbone of the Americas. + Amer. Assoc. Petroleum Geol., pp. 88-100. + +LYNCH, J. D. + + 1965. A review of the eleutherodactylid frog genus + _Microbatrachylus_ (Leptodactylidae). Nat. Hist. Misc., + 182:1-12, December 15. + +MYERS, G. S. + + 1962. The American leptodactylid frog genus _Eleutherodactylus_, + _Hylodes_ (= _Elosia_), and _Caudiverbera_ + (= _Calytocephalus_). Copeia, 1:195-202, April 11. + +NOBLE, G. K. + + 1925. An outline of the relation of the ontogeny to + phylogeny within the Amphibia. I. Amer. Mus. Nov., + 165:1-17, April 16. + +PIATT, J. + + 1934. The systematic status of Eleutherodactylus latrans + (Cope). Amer. Midl. Nat., 15:89-91, February 15. + +SMITH, H. M. and TAYLOR, E. H. + + 1948. An annotated checklist and key to the Amphibia of + Mexico. Bull. U. S. Natl. Mus., 194:1-118. June 7. + +TAYLOR, E. H. + + 1952. A review of the frogs and toads of Costa Rica. Univ. + Kansas Sci. Bull., 35:577-942, July 1. + +ZWEIFEL, R. G. + + 1956. A survey of the frogs of the _augusti_ group, genus + _Eleutherodactylus_. Amer. Mus. Novitates, 1813:1-35, + December 23. + + +_Transmitted July 11, 1967._ + + +31-9418 + + + + + * * * * * + +Transcriber's Notes + + +Bold text is shown within ~tildes~. + +Italicized text is shown within _underscores_. + +Illustrations have been moved to avoid breaking up paragraphs of text. + +Page 507: Changed know to known (are not know in Tomodactylus). + +Page 507: Added closing parenthesis in Fig. 2 caption after × 3. + +Page 509: Changed compeltely to completely (compeltely roofed skulls). + +Page 512: Veracruuz may be a typo for Veracruz (Sinaloa to Veracruuz). + +Page 514: Changed two occurrences of Hylatophryne to Hylactophryne. + + + + + + + + +End of the Project Gutenberg EBook of Genera of Leptodactylid Frogs in México, by +John D. 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Lynch. + </title> + <style type="text/css"> + +body { + margin-left: 10%; + margin-right: 10%; +} + + h1,h2,h3 { + text-align: center; /* all headings centered */ + clear: both; +} + +p { + margin-top: .75em; + text-align: justify; + margin-bottom: .75em; +} + + p.title { text-align: center; text-indent: 0; + font-weight: bold; + line-height: 1.4; margin-bottom: 3em; } + + +hr { + width: 33%; + margin-top: 2em; + margin-bottom: 2em; + margin-left: auto; + margin-right: auto; + clear: both; +} + +table { + margin-left: auto; + margin-right: auto; +} + +.pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: smaller; + text-align: right; +} /* page numbers */ + +.blockquot { + margin-left: 5%; + margin-right: 10%; +} + +.i4 {display: block; margin-left: 2.5em; + padding-left: 2.5em; text-indent: -2.5em;} + +.center {text-align: center;} + +.smcap {font-variant: small-caps;} + +.u {text-decoration: underline;} + +.caption {font-weight: bold;} + +.figcenter { + margin: auto; + text-align: center; +} + + + </style> + </head> +<body> + + +<pre> + +Project Gutenberg's Genera of Leptodactylid Frogs in México, 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: Genera of Leptodactylid Frogs in México + +Author: John D. Lynch + +Release Date: March 30, 2010 [EBook #31830] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK LEPTODACTYLID FROGS IN MEXICO *** + + + + +Produced by Chris Curnow, Joseph Cooper, Diane Monico, and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + +</pre> + + + + + + + +<p class="title"><span class="smcap">University of Kansas Publications</span><br /> +<span class="smcap">Museum of Natural History</span><br /><br /> + +Volume 17, No. 11, pp. 503-515, 5 figs.<br /> +March 20, 1968</p> +<hr style="width: 25%;" /> + + + +<h1>Genera of Leptodactylid Frogs in México</h1> + + +<p class="title">BY<br /><br /> + +<big>JOHN D. LYNCH</big><br /><br /><br /> + + +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1968<br /> +</p> +<hr style="width: 65%;" /> + + + +<p class="title"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Frank B. Cross<br /> +<br /> +<br /> +Volume 17, No. 11, pp. 503-515, 5 figs.<br /> +Published March 20, 1968<br /> +<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +<br /> +<small>PRINTED BY<br /> +ROBERT R. (BOB) SANDERS, STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1968<br /> +<br /> +31-9418<br /></small> +</p> + + + +<hr style="width: 65%;" /><p><span class="pagenum"><a name="Page_505" id="Page_505">[Pg 505]</a></span></p> +<h2>Genera of Leptodactylid Frogs in México</h2> + +<p class="center"><b>BY JOHN D. LYNCH</b></p> + + +<h2>INTRODUCTION</h2> + +<p>According to the most recent review of the Mexican amphibian +fauna (Smith and Taylor, 1948), six genera of leptodactylid frogs +occur in México. One other genus, <i>Pleurodema</i>, occurs in Lower +Central America. Smith and Taylor recognized one species of <i>Engystomops</i>, +28 of <i>Eleutherodactylus</i>, three of <i>Leptodactylus</i>, eight of +<i>Microbatrachylus</i>, 12 of <i>Syrrhophus</i>, and five of <i>Tomodactylus</i>. Subsequent +to the publication of their checklist of the Mexican amphibia +(1948), numerous taxonomic changes have been proposed. Many +species of <i>Eleutherodactylus</i> have been added to the fauna, either +through the extension of their recorded ranges into México from +Guatemala or by the recognition of species unknown in 1948, +whereas some nominal species have been synonymized. <i>Microbatrachylus</i> +has been regarded as synonymous with <i>Eleutherodactylus</i> +(Lynch, 1965); four species of <i>Microbatrachylus</i> currently are +regarded as valid (Duellman, 1961, Lynch, 1965). <i>Syrrhophus</i> was +revised in part by Duellman (1958) and Firschein (1954), and a +species of <i>Tomodactylus</i> transferred to <i>Syrrhophus</i> by Dixon (1957), +who redefined <i>Tomodactylus</i> and added more species to the genus.</p> + +<p>Since beginning my studies of the Mexican leptodactylids in 1962, +I have become acutely aware of difficulties involved in defining the +genera. A revision of <i>Eleutherodactylus</i> and a review of <i>Syrrhophus</i> +are nearing completion, but prior to their publication it is desirable +to redefine the genera of the Mexican leptodactylids, and in so doing +recognize an heretofore unnamed genus. The definitions of <i>Eleutherodactylus</i> +and <i>Leptodactylus</i> may need to be altered in the future, +since both are widespread in South America and occur in the +West Indies. Their definitions as given here are as precise as +present knowledge permits. <i>Syrrhophus</i> and <i>Tomodactylus</i> are +small assemblages that occur only in southwestern United States, +México, and Guatemala.</p> + +<p>Taylor (1952) synonymized <i>Engystomops</i> with <i>Eupemphix</i> which, +although related, should be regarded as generically distinct (Gallardo, +1965). Perhaps the most conservative classification is that of +Myers (1962) who, without published evidence, combined <i>Eleutherodactylus</i>,<span class="pagenum"><a name="Page_506" id="Page_506">[Pg 506]</a></span> +<i>Syrrhophus</i>, and the South American <i>Lithodytes</i> in a +single genus.</p> + +<p>The major problem for students working with the Mexican leptodactylids +has not been the separation of <i>Engystomops</i> or <i>Leptodactylus</i> +from other genera but the separation and definition of the +eleutherodactyline frogs currently placed in three genera, <i>Eleutherodactylus</i>, +<i>Syrrhophus</i>, and <i>Tomodactylus</i>. As will be shown in this +paper, these are more conveniently placed in four genera. Once a +fourth genus is recognized, certain phylogenetic problems disappear +and a reasonable zoogeographic interpretation is possible for Middle +American leptodactylid distribution.</p> + + + +<hr style="width: 65%;" /> +<h2><a name="ANALYSIS_OF_CHARACTERS" id="ANALYSIS_OF_CHARACTERS"></a>ANALYSIS OF CHARACTERS</h2> + + +<p>In México and northern Central America approximately 55 species +of eleutherodactyline frogs (<i>Eleutherodactylus</i>, <i>Syrrhophus</i>, and +<i>Tomodactylus</i>) are known. Four genera can be recognized on the +basis of the nature of inguinal glands, morphology of the hands and +feet, and certain osteological features.</p> + + +<p class="figcenter" style="width: 600px;"> +<a name="fig1" id="fig1"></a> +<img src="images/image001.jpg" width="600" height="370" alt="Fig. 1. Tomodactylus angustidigitorum (UMMZ 114305, × 4.5) illustrating +the lumbo-inguinal gland typical of members of the genus. From a kodachrome +by Wm. E. Duellman." title="Fig. 1. Tomodactylus angustidigitorum (UMMZ 114305, × 4.5) illustrating +the lumbo-inguinal gland typical of members of the genus. From a kodachrome +by Wm. E. Duellman." /> +<span class="caption">Fig. 1. Tomodactylus angustidigitorum (UMMZ 114305, × 4.5) illustrating +the lumbo-inguinal gland typical of members of the genus. From a kodachrome +by Wm. E. Duellman.</span> +</p> + +<h3>Glands</h3> + +<p>Leptodactylids have a variety of glands that have been used as +generic characters. Smith and Taylor (1948) regarded the so-called +inguinal gland as a generic character in Mexican eleutherodaycty-lines. +Lynch (1965) showed that <i>Eleutherodactylus</i> and <i>Microbatrachylus</i><span class="pagenum"><a name="Page_507" id="Page_507">[Pg 507]</a></span> +cannot be separated by the nature of the gland or the +condition of the prevomers (dentate or not). <i>Syrrhophus</i> and <i>Tomodactylus</i>, +as defined by Smith and Taylor (1948), are not generically +distinct because of overlap in the condition of the prevomers and in +the development of the gland. Firschein (1954) stated that <i>Syrrhophus</i> +differed from <i>Tomodactylus</i> by having an axillary gland, +but it is now known that one species of <i>Syrrhophus</i> lacks the gland.</p> + +<p>The inguinal glands of <i>Eleutherodactylus</i> and <i>Syrrhophus</i>, if present, +are diffuse, irregular in outline, and generally not prominent; +in <i>Tomodactylus</i> the gland is higher on the body (a lumbo-inguinal +gland), compact, oval in outline, and prominent (<a href="#fig1">Fig. 1</a>). Axillary +glands occur in most <i>Syrrhophus</i> but are not known in <i>Tomodactylus</i> +or <i>Eleutherodactylus</i>.</p> + + +<h3>Hands and feet</h3> + +<p>The tips of the digits are laterally expanded in most <i>Eleutherodactylus</i>, +<i>Syrrhophus</i>, and <i>Tomodactylus</i>. Two species of <i>Eleutherodactylus</i> +(<i>augusti</i> and <i>tarahumarensis</i>) and two <i>Tomodactylus</i> (<i>angustidigitorum</i> +and <i>grandis</i>) lack any expansion of the digital tips. +All but two of the species of eleutherodactyline frogs (<i>E. augusti</i> +and <i>E. tarahumarensis</i>) have a transverse groove across the tips of +the digits (<a href="#fig2">Fig. 2</a>).</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig2" id="fig2"></a> +<img src="images/image002.jpg" width="600" height="410" alt="Fig. 2. Palmar views of the hands and lateral views of the tip of the third +digits of Eleutherodactylus alfredi (left, KU 93994, × 5) and Hylactophryne +augusti (right, KU 102594, × 3)." title="Fig. 2. Palmar views of the hands and lateral views of the tip of the third +digits of Eleutherodactylus alfredi (left, KU 93994, × 5) and Hylactophryne +augusti (right, KU 102594, × 3)." /> +<span class="caption">Fig. 2. Palmar views of the hands and lateral views of the tip of the third +digits of Eleutherodactylus alfredi (left, KU 93994, × 5) and Hylactophryne +augusti (right, KU 102594, × 3).</span> +</p> +<p><span class="pagenum"><a name="Page_508" id="Page_508">[Pg 508]</a></span></p> + +<p>Supernumerary tubercles rarely are present on the feet of <i>Eleutherodactylus</i>, +but are present and numerous in every species of +<i>Syrrhophus</i>, <i>Tomodactylus</i>, and in the members of the <i>augusti</i> group +of <i>Eleutherodactylus</i> (<a href="#fig3">Fig. 3</a>). The tubercles are small and numerous +in <i>Syrrhophus</i> and larger in <i>Tomodactylus</i> and the <i>Eleutherodactylus +augusti</i> group. Most species of <i>Eleutherodactylus</i> have no +plantar supernumerary tubercles; a few species have such tubercles, +which never extend between the metatarsal tubercles as in <i>Syrrhophus</i> +and <i>Tomodactylus</i>.</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig3" id="fig3"></a> +<img src="images/image003.jpg" width="600" height="490" alt="Fig. 3. Plantar views of feet of Eleutherodactylus alfredi (left, KU 93994, +× 4.5), Syrrhophus pipilans nebulosus (middle, KU 58900, × 7.5), and Hylactophryne +augusti (right, KU 102594, × 3) showing differences in size and +arrangement of supernumerary tubercles." title="Fig. 3. Plantar views of feet of Eleutherodactylus alfredi (left, KU 93994, +× 4.5), Syrrhophus pipilans nebulosus (middle, KU 58900, × 7.5), and Hylactophryne +augusti (right, KU 102594, × 3) showing differences in size and +arrangement of supernumerary tubercles." /> +<span class="caption">Fig. 3. Plantar views of feet of Eleutherodactylus alfredi (left, KU 93994, +× 4.5), Syrrhophus pipilans nebulosus (middle, KU 58900, × 7.5), and Hylactophryne +augusti (right, KU 102594, × 3) showing differences in size and +arrangement of supernumerary tubercles.</span> +</p> + +<p>Tarsal folds and tubercles are lacking in <i>Syrrhophus</i>, <i>Tomodactylus</i>, +and the <i>augusti</i> group of <i>Eleutherodactylus</i>. Several species of +<i>Eleutherodactylus</i> lack tarsal folds and tubercles, but in nearly +every species group, one or more species possess either an inner +tarsal fold, inner tarsal tubercle(s), or outer tarsal tubercles.</p> + +<p>The terminal phalanges of <i>Syrrhophus</i>, <i>Tomodactylus</i>, and all +<i>Eleutherodactylus</i> (except the frogs of the <i>augusti</i> group) are distinctly<span class="pagenum"><a name="Page_509" id="Page_509">[Pg 509]</a></span> +T-shaped. In the latter, the bones are knob-shaped distally +(<a href="#fig4">Fig. 4</a>). T-shaped terminal phalanges also are present in <i>Lithodytes</i> +and <i>Trachyphrynus</i> but not in other leptodactylid genera. At +least one species of <i>Eupsophus</i> (<i>E. quixensis</i>) has terminal phalanges +that resemble those of the <i>Eleutherodactylus augusti</i> group. +Several species of <i>Eleutherodactylus</i>, <i>Syrrhophus</i>, and <i>Tomodactylus</i> +with slender fingers have T-shaped terminal phalanges although the +terminal dilations proportionately are only scarcely wider than the +finger tips in the <i>Eleutherodactylus augusti</i> group. The presence of +a terminal groove at the tip of the finger is an external indicator of +the T-shaped terminal phalanges.</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig4" id="fig4"></a> +<img src="images/image004.jpg" width="600" height="356" alt="Fig. 4. Terminal phalanges of four leptodactylid frogs (all × 13.5). (a) +Eleutherodactylus mexicanus, KU 55593; (b) Eupsophus roseus, KU 84731; +(c) Eupsophus quixensis, UIMNH 59643; and (d) Hylactophryne augusti, +KU 56192." title="Fig. 4. Terminal phalanges of four leptodactylid frogs (all × 13.5). (a) +Eleutherodactylus mexicanus, KU 55593; (b) Eupsophus roseus, KU 84731; +(c) Eupsophus quixensis, UIMNH 59643; and (d) Hylactophryne augusti, +KU 56192." /> +<span class="caption">Fig. 4. Terminal phalanges of four leptodactylid frogs (all × 13.5). (a) +Eleutherodactylus mexicanus, KU 55593; (b) Eupsophus roseus, KU 84731; +(c) Eupsophus quixensis, UIMNH 59643; and (d) Hylactophryne augusti, +KU 56192.</span> +</p> + + +<h3>Skull</h3> + +<p>All Mexican eleutherodactyline frogs have quadratojugal-maxillary +articulations, completely roofed skulls in adults, median contact of +the nasals, separated occipital condyles, and large prevomers. The +premaxillae of all species are visible when the skulls are viewed from +directly above. The pterygoid lacks a medioventral flange and does +not meet the palatine. In no species is the anterior arm of the +squamosal in contact with the maxillary. Of the numerous species +examined (30 <i>Eleutherodactylus</i>, four <i>Syrrhophus</i>, and four <i>Tomodactylus</i>), +the species in the <i>Eleutherodactylus augusti</i> group are +unique in having a sphenethmoid with a blunt anterior edge.</p> + +<p><span class="pagenum"><a name="Page_510" id="Page_510">[Pg 510]</a></span></p> +<h3>Pectoral Girdle</h3> + +<p>All species have large cartilaginous plates in the pectoral girdles; +none possesses a bony style. No divergent modifications of the +clavicle and coracoid bones are known in the family.</p> + + + +<hr style="width: 65%;" /> +<h2><a name="GENERIC_ACCOUNTS" id="GENERIC_ACCOUNTS"></a>GENERIC ACCOUNTS</h2> + + +<h3><span style="font-weight: normal">Genus</span> <b>Eleutherodactylus</b> <span style="font-weight: normal">Dumeril and Bibron, 1841</span></h3> + +<div class="blockquot"><p><i>Type-species.</i>—<i>Hylodes martinicensis</i> Tschudi, 1838</p> + +<p><i>Diagnosis and definition.</i>—Small to large frogs (12 to 110 mm. snout-vent +length) having slightly to widely expanded digital pads, each pad bearing a +terminal transverse groove; lumbo-inguinal, inguinal, and axillary glands absent, +or if present, diffuse, irregular in outline, not compact; plantar supernumerary +tubercles absent, or if present, six or fewer, restricted to distal area of plantar +surface, and not extending between metatarsal tubercles; tarsus bearing inner +or outer tubercles or folds or not; toes free to one-half webbed; terminal phalanges +T-shaped; sternum cartilaginous, lacking bony style; sphenethmoid not +truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in +contact; anterior arm of squamosal not in contact with maxillary; dermal cranial +elements not involved in integumentary-cranial co-ossification; prevomers large, +dentigerous processes present or not, dentate or not; maxillary and premaxillary +bones dentate; occipital condyles separated; development direct.</p> + +<p><i>Composition.</i>—About 420 names have been applied to frogs of this genus; +many of these names are synonyms, and many other species remain undescribed +and unnamed. Perhaps the genus contains 350 species. Thirty-one +species occur in México and northern Central America.</p> + +<p><i>Distribution.</i>—From Tamaulipas and Sinaloa, México, exclusive of the Mexican +Plateau, to at least Peru and southernmost Brazil and throughout the West +Indies. Introduced into Florida.</p> + +<p><i>Etymology.</i>—Greek (<i>eleuthero</i> + <i>dactylus</i>) meaning free-toed.</p></div> + + +<h3><span style="font-weight: normal">Genus</span> <b>Engystomops</b> <span style="font-weight: normal">Jiménez de la Espada, 1872</span></h3> + +<div class="blockquot"><p><i>Type species.</i>—<i>Engystomops petersi</i> Jiménez de la Espada, 1872</p> + +<p><i>Diagnosis and definition.</i>—Small frogs (20 to 40 mm. snout-vent length) +having undilated digital tips lacking transverse grooves; lumbo-inguinal or +inguinal glands absent; plantar supernumerary tubercles present, extending between +metatarsal tubercles; tarsus bearing spinelike tubercle on inner edge; toes +free; terminal phalanges pointed; sternum bearing bony style; spenethmoid +not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal +in articular contact; anterior arm of squamosal not in contact with maxillary; +dermal cranial elements not involved in integumentary-cranial co-ossification; +prevomers moderate in size, lacking teeth; maxillary and premaxillary bones +edentate; occipital condyles separated; tadpole free living.</p> + +<p><i>Composition.</i>—Four nominal species (<i>E. petersi</i>, <i>E. pustulatus</i>, <i>E. pustulosus</i> +and <i>E. schereri</i>).</p> + +<p><i>Distribution.</i>—Central Veracruz and eastern Oaxaca, México, to Trinidad, +Bolivia, and Peru, east of the Andes.</p> + +<p><i>Etymology.</i>—Greek (<i>engys</i> + <i>stoma</i>) meaning narrow-mouthed.</p></div> + +<p><span class="pagenum"><a name="Page_511" id="Page_511">[Pg 511]</a></span></p> +<h3><span style="font-weight: normal">Genus</span> <b>Hylactophryne</b> <span style="font-weight: normal">new genus</span></h3> + +<div class="blockquot"><p><i>Type-species.</i>—<i>Hylodes augusti</i> Dugés, 1879</p> + +<p><i>Diagnosis and definition.</i>—Medium to large frogs (37 to 94 mm. snout-vent +length) having undilated digital tips lacking terminal grooves; lumbo-inguinal +or inguinal glands absent; plantar supernumerary tubercles present, prominent, +extending to but not between metatarsal tubercles; tarsus lacking tubercles or +folds; toes free of webbing; terminal phalanges knob-shaped, lacking elongate +lateral expansions; sternum cartilaginous, lacking bony style; sphenethmoid +truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in +articular contact; anterior arm of squamosal not in contact with maxillary; dermal +cranial elements not involved in integumentary-cranial co-ossification; prevomers +large, bearing dentigerous processes; maxillary and premaxillary bones +dentate; occipital condyles separated; development direct.</p> + +<p><i>Composition.</i>—Two species, <i>H. augusti</i> and <i>H. tarahumarensis</i>, the former +composed of four subspecies (Zweifel, 1956).</p> + +<p><i>Distribution.</i>—From Arizona, New Mexico, and Texas to Guerrero and +Puebla, México, and a relict population on Cerro Quingola (just west of the +Isthmus of Tehuantepec, México).</p> + +<p><i>Etymology.</i>—Greek (<i>hylactor</i> + <i>phryne</i>) meaning barking toad; in reference +to the voice and common name.</p></div> + + +<h3><span style="font-weight: normal">Genus</span> <b>Leptodactylus</b> <span style="font-weight: normal">Fitzinger, 1826</span></h3> + +<div class="blockquot"><p><i>Type-species.</i>—<i>Leptodactylus typhonia</i> Fitzinger, 1826</p> + +<p><i>Diagnosis and definition.</i>—Small to large frogs (30 to about 200 m., snout-vent +length) having undilated to slightly expanded digital tips bearing pads, +no transverse groove at tips of digits; lumbo-inguinal, axillary, and/or ventral +glands present or not, low, diffuse; plantar supernumerary tubercles generally +absent, if present not extending between metatarsal tubercles; tarsus bearing +tarsal folds or not; toes free of webbing, extensive lateral fringes present in +some species; terminal phalanges pointed, not T-shaped; sternum bearing bony +style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary +and quadratojugal in articular contact; anterior arm of squamosal not in +contact with maxillary; dermal cranial elements not involved in integumentary-cranial +co-ossification; prevomers large, bearing dentigerous processes; maxillary +and premaxillary bones dentate; occipital condyles separated; tadpole free +living.</p> + +<p><i>Composition.</i>—Sixty species according to Smith and Taylor (1948); 54 +according to Gorham (1963); Argentinian authors have described several +more in recent years.</p> + +<p><i>Distribution.</i>—Southern Sonora, México, and southern Texas throughout the +Central and South American lowlands to Argentina. Also known from Hispaniola +and Puerto Rico in the Greater Antilles and a few islands in the Lesser +Antilles.</p> + +<p><i>Entymology.</i>—Greek (<i>leptos</i> + <i>dactylus</i>) meaning slender toes.</p></div> + +<p><span class="pagenum"><a name="Page_512" id="Page_512">[Pg 512]</a></span></p> +<h3><span style="font-weight: normal">Genus</span> <b>Syrrhophus</b> <span style="font-weight: normal">Cope, 1878</span></h3> + +<div class="blockquot"><p><i>Type-species.</i>—<i>Syrrhophus marnockii</i> Cope, 1878</p> + +<p><i>Diagnosis and definition.</i>—Small to medium sized frogs (18 to 40 mm. +snout-vent) having slight to prominent digital expansions with transverse groove +at tip of each digit; lumbo-inguinal and inguinal gland flattened, irregular in +outline, not compact and oval; axillary glands present or not; plantar supernumerary +tubercles numerous, more than eight, usually extending between +metatarsal tubercles; tarsus lacking tubercles or folds; toes free or basally +webbed; terminal phalanges T-shaped; sternum cartilaginous, lacking bony +style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary +and quadratojugal in articular contact; anterior arm of squamosal not in +contact with maxillary; dermal cranial elements not involved in integumentary-cranial +co-ossification; prevomers large, usually lacking dentigerous processes +and teeth; maxillary and premaxillary bones dentate; occipital condyles separated; +development direct.</p> + +<p><i>Composition.</i>—Thirteen species; the species described as, or later referred +to, <i>Syrrhophus</i> from Lower Central America and South America are <i>Eleutherodactylus</i> +or <i>Eupsophus</i>.</p> + +<p><i>Distribution.</i>—Low to moderate elevations from Sinaloa, México, to Guatemala +on the Pacific versant; from the Edwards and Stockton plateaus of Texas +to British Honduras on the Caribbean versant.</p> + +<p><i>Etymology.</i>—Greek, emendation of <i>syrrhaptos</i>, meaning sewn together in +reference to the united outer metatarsals.</p></div> + + +<h3><span style="font-weight: normal">Genus</span> <b>Tomodactylus</b> <span style="font-weight: normal">Günther, 1900</span></h3> + +<div class="blockquot"><p><i>Type-species.</i>—<i>Tomodactylus amulae</i> Günther, 1900.</p> + +<p><i>Diagnosis and definition.</i>—Small frogs (20 to 35 mm. snout-vent length) +having digital expansions or not, with transverse groove across tip of each +digit; lumbo-inguinal gland prominently elevated, compact, oval, often patterned; +axillary glands absent; plantar supernumerary tubercles numerous, more +than eight, usually extending between metatarsal tubercles; tarsus lacking tubercles +or folds; toes free; terminal phalanges T-shaped; sternum cartilaginous, +lacking bony style; sphenethmoid not truncate anteriorly; nasals in contact +medially; maxillary and quadratojugal in articular contact; anterior arm of +squamosal not in contact with maxillary; dermal cranial elements not involved +in integumentary-cranial co-ossification; prevomers large, usually bearing dentigerous +processes; maxillary and premaxillary bones dentate; occipital condyles +separated; development direct.</p> + +<p><i>Composition.</i>—Ten species.</p> + +<p><i>Distribution.</i>—The southern edge of the Mexican Plateau from Sinaloa to +Veracruuz and onto the Oaxaca highlands and Sierra Madre del Sur.</p> + +<p><i>Etymology.</i>—Greek (<i>tomis</i> + <i>dactylus</i>) meaning knife toe; in reference to +either the sharp subarticular tubercles or the unwebbed toes.</p></div> + + + +<hr style="width: 65%;" /> +<h2><a name="DISCUSSION" id="DISCUSSION"></a>DISCUSSION</h2> + + +<p>The preceding definitions only slightly alter the present generic +limits of Mexican leptodactylids. Two species, previously regarded +as <i>Eleutherodactylus</i>, are transferred to the new genus <i>Hylactophryne</i>.<span class="pagenum"><a name="Page_513" id="Page_513">[Pg 513]</a></span> +The arrangement of the species of <i>Syrrhophus</i> and <i>Tomodactylus</i> +remains the same as concluded by Dixon (1957), Duellman +(1958), and Firschein (1954) in their reviews of the genera.</p> + +<p>Lumbo-inguinal glands are most prominent in the genera <i>Pleurodema</i> +and <i>Tomodactylus</i>. Various nondescript glands are present +in many genera, but none is so well developed as those of <i>Pleurodema</i> +and <i>Tomodactylus</i>.</p> + +<p>At least nine leptodactylid genera are either known or thought to +be terrestrial breeders lacking a free-living tadpole stage (<i>Eleutherodactylus</i>, +<i>Euparkerella</i>, <i>Hylactophryne</i>, <i>Niceforonia</i>, <i>Noblella</i>, +<i>Sminthillus</i>, <i>Syrrhophus</i>, <i>Tomodactylus</i> and <i>Trachyphrynus</i>). <i>Niceforonia</i> +and <i>Trachyphrynus</i>, and probably <i>Hylactophryne</i>, are not +closely related to the other genera. Direct development probably +is an adaptation to adverse environmental conditions since many of +the species occur in semi-arid or cold (Andean páramos) areas. +<i>Eleutherodactylus</i> is generally thought to be the stock from which +<i>Euparkerella</i>, <i>Noblella</i>, and <i>Sminthillus</i> evolved (Griffiths, 1959) +and from which <i>Syrrhophus</i> and <i>Tomodactylus</i> are derived (Firschein, +1954).</p> + +<p>The present distribution of <i>Hylactophryne</i> (isolated on the Mexican +Plateau) and its digital form (like that of Papuan and many +primitive South American leptodactylids) suggest that the genus +was isolated in México throughout the Tertiary, whereas the other +Central American genera are either post-Pliocene derivatives of +<i>Eleutherodactylus</i> or invaders of Central America from South America +since the mid-Pliocene land bridge was formed (Lloyd, 1963).</p> + +<p>Piatt (1934) presented arguments against assigning <i>Eleutherodactylus +latrans</i> to the genus <i>Lithodytes</i> and concluded that it was a +"true" <i>Eleutherodactylus</i>. Contrary to his arguments, <i>latrans</i> +(= <i>augusti</i> of Zweifel) and <i>E. tarahumarensis</i> Taylor differ from +all other <i>Eleutherodactylus</i> (and <i>Syrrhophus</i> and <i>Tomodactylus</i>) +in the nature of the tips of the digits (external and skeletal). The +digits of <i>Hylactophryne</i> are like those of <i>Eupsophus</i>. My study of +nearly all genera of leptodactylids indicates that Noble (1925) was +correct in suggesting that <i>Borborocoetes</i> (= <i>Eupsophus</i>) is a close +relative of <i>Eleutherodactylus latrans</i>, although Noble's arguments +were based in part upon false evidence concerning the breeding +habits of <i>E. latrans</i>, then thought to have a free-living tadpole.</p> + +<p>Kellogg (1932) and Piatt (1934) argued that the terminal phalanges +of <i>E. latrans</i> were typically eleutherodactyline. The variation +of this character in <i>Eupsophus</i> (see <a href="#fig4">Fig. 4</a>) ranges from +knobbed to bifurcate or Y-shaped (T-shaped in <i>Eleutherodactylus</i>,<span class="pagenum"><a name="Page_514" id="Page_514">[Pg 514]</a></span> +<i>Syrrhophus</i> and <i>Tomodactylus</i>) and encompasses the nature of the +character represented in <i>Hylactophryne</i>. <i>Eupsophus</i> differs from +<i>Hylactophryne</i> in possessing a frontoparietal fontanelle, in generally +having a maxillary-quadratojugal gap, and in having a free swimming +tadpole stage.</p> + +<p class="figcenter" style="width: 600px;"> +<a name="fig5" id="fig5"></a> +<img src="images/image005.png" width="600" height="518" alt="Fig. 5. Outline drawings of Leptodactylus melanonotus (left, KU 65704, × 2) +and Eleutherodactylus alfredi (right, KU 93994, × 2)." title="Fig. 5. Outline drawings of Leptodactylus melanonotus (left, KU 65704, × 2) +and Eleutherodactylus alfredi (right, KU 93994, × 2)." /> +<span class="caption">Fig. 5. Outline drawings of Leptodactylus melanonotus (left, KU 65704, × 2) +and Eleutherodactylus alfredi (right, KU 93994, × 2).</span> +</p> + + + +<hr style="width: 65%;" /> +<h2><a name="KEY_TO_MEXICAN_LEPTODACTYLID_GENERA" id="KEY_TO_MEXICAN_LEPTODACTYLID_GENERA"></a>KEY TO MEXICAN LEPTODACTYLID GENERA</h2> + + + +<div class="center"> +<table border="0" cellpadding="4" cellspacing="0" summary="key"> +<tr><td align="left">1.</td><td align="left"> Small (20-40 mm.), pustular, toadlike frogs; maxillary and premaxillary bones not bearing teeth</td><td align="right"><i>Engystomops</i></td></tr> +<tr><td align="left"> </td><td align="left">Large (20-110 mm.), smooth skinned and non-toadlike frogs; maxillary and premaxillary bones bearing teeth</td><td align="right">2</td></tr> +<tr><td align="left">2.</td><td align="left"> No conspicuous waist (<a href="#fig5">Fig. 5</a>); sternum bearing bony style,</td><td align="right"><i>Leptodactylus</i></td></tr> +<tr><td align="left"> </td><td align="left">Constrictions at waist (<a href="#fig5">Fig. 5</a>); sternum cartilaginous, no bony style</td><td align="right">3</td></tr> +<tr><td align="left">3.</td><td align="left"> Few (less than six), if any, supernumerary tubercles on plantar surface</td><td align="right"><i>Eleutherodactylus</i></td></tr> +<tr><td align="left"> </td><td align="left">Many (more than 8) supernumerary tubercles on plantar surfaces</td><td align="right">4</td></tr> +<tr><td align="left">4.</td><td align="left"> Terminal, transverse groove across tip of digits, especially outer two fingers, digits expanded or not; small frogs (18 to 40 mm.)</td><td align="right">5</td></tr> +<tr><td align="left"> </td><td align="left">Tips of digits lacking transverse groove; digits unexpanded; medium-sized to large frogs (37 to 94 mm.)</td><td align="right"><i>Hylactophryne</i></td></tr> +<tr><td align="left">5.</td><td align="left"> Lumbo-inguinal gland compact, oval</td><td align="right"><i>Tomodactylus</i></td></tr> +<tr><td align="left"> </td><td align="left">Lumbo-inguinal or inguinal gland absent or diffuse and irregular in outline</td><td align="right"><i>Syrrhophus</i></td></tr> +</table></div> + + +<p><span class="pagenum"><a name="Page_515" id="Page_515">[Pg 515]</a></span></p><hr style="width: 65%;" /> +<h2><a name="LITERATURE_CITED" id="LITERATURE_CITED"></a>LITERATURE CITED</h2> + + +<p><span class="smcap">Dixon, J. R.</span></p> + +<p class="i4">1957. Geographic variation and distribution of the genus <i>Tomodactylus</i> +in Mexico. Texas Jour. Sci., 9:379-409, December.</p> + +<p><span class="smcap">Duellman, W. E.</span></p> + +<p class="i4">1958. A review of the frogs of the genus Syrrhophus in western Mexico. +Occas. Papers Mus. Zool. Univ. Michigan, 594:1-15, June 6.</p> + +<p class="i4">1961. The amphibians and reptiles of Michoacan, Mexico. Univ. Kansas +Publ. Mus. Nat. Hist., 15:1-148, December 20.</p> + +<p><span class="smcap">Firschein, I. L.</span></p> + +<p class="i4">1954. Definition of some little understood members of the leptodactylid +genus <i>Syrrhophus</i>, with a description of a new species. Copeia, +1:48-58, February 19.</p> + +<p><span class="smcap">Gallardo, J. M.</span></p> + +<p class="i4">1965. A proposito de los Leptodactylidae (Amphibia Anura). Papeis +Avulsos, 17:77-87, January 30.</p> + +<p><span class="smcap">Gorham, S. W.</span></p> + +<p class="i4">1963. The comparative number of species of amphibians in Canada and +other countries. III. Summary of species of anurans. Canadian +Field-Nat., 77:13-48, March.</p> + +<p><span class="smcap">Griffiths, I.</span></p> + +<p class="i4">1959. The phylogeny of Sminthillus limbatus and the status of the +Brachycephalidae (Amphibia Salientia). Proc. Zool. Soc. London, +132:457-87, May.</p> + +<p><span class="smcap">Kellogg, R.</span></p> + +<p class="i4">1932. Mexican tailless amphibians in the United States National Museum. +Bull. U. S. Natl. Mus., 160:224 pp., March 31.</p> + +<p><span class="smcap">Lloyd, J. J.</span></p> + +<p class="i4">1963. Tectonic history of the south Central-American orogen, <i>in</i> Childs +and Beebe eds., Backbone of the Americas. Amer. Assoc. Petroleum +Geol., pp. 88-100.</p> + +<p><span class="smcap">Lynch, J. D.</span></p> + +<p class="i4">1965. A review of the eleutherodactylid frog genus <i>Microbatrachylus</i> +(Leptodactylidae). Nat. Hist. Misc., 182:1-12, December 15.</p> + +<p><span class="smcap">Myers, G. S.</span></p> + +<p class="i4">1962. The American leptodactylid frog genus <i>Eleutherodactylus</i>, <i>Hylodes</i> +(= <i>Elosia</i>), and <i>Caudiverbera</i> (= <i>Calytocephalus</i>). Copeia, +1:195-202, April 11.</p> + +<p><span class="smcap">Noble, G. K.</span></p> + +<p class="i4">1925. An outline of the relation of the ontogeny to phylogeny within the +Amphibia. I. Amer. Mus. Nov., 165:1-17, April 16.</p> + +<p><span class="smcap">Piatt, J.</span></p> + +<p class="i4">1934. The systematic status of Eleutherodactylus latrans (Cope). Amer. +Midl. Nat., 15:89-91, February 15.</p> + +<p><span class="smcap">Smith, H. M.</span> and <span class="smcap">Taylor, E. H.</span></p> + +<p class="i4">1948. An annotated checklist and key to the Amphibia of Mexico. Bull. +U. S. Natl. Mus., 194:1-118. June 7.</p> + +<p><span class="pagenum"><a name="Page_516" id="Page_516">[Pg 516]</a></span></p> +<p><span class="smcap">Taylor, E. H.</span></p> + +<p class="i4">1952. A review of the frogs and toads of Costa Rica. Univ. Kansas Sci. +Bull., 35:577-942, July 1.</p> + +<p><span class="smcap">Zweifel, R. G.</span></p> + +<p class="i4">1956. A survey of the frogs of the <i>augusti</i> group, genus <i>Eleutherodactylus</i>. +Amer. Mus. Novitates, 1813:1-35, December 23.</p> + + +<p><i>Transmitted July 11, 1967.</i></p> + + +<p class="center"><small>31-9418</small></p> + + + + +<hr style="width: 65%;" /> + +<h3>Transcriber's Notes</h3> + + +<p>Illustrations have been moved to avoid breaking up paragraphs of text.</p> + +<p>Page <a href="#Page_507">507</a>: Changed know to known (are not know in Tomodactylus).</p> + +<p>Page <a href="#Page_507">507</a>: Added closing parenthesis in <a href="#fig2">Fig. 2</a> caption after × 3.</p> + +<p>Page <a href="#Page_509">509</a>: Changed compeltely to completely (compeltely roofed skulls).</p> + +<p>Page <a href="#Page_512">512</a>: Veracruuz may be a typo for Veracruz (Sinaloa to Veracruuz).</p> + +<p>Page <a href="#Page_514">514</a>: Changed two occurrences of Hylatophryne to Hylactophryne.</p> + + + + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Genera of Leptodactylid Frogs in México, by +John D. 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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: Genera of Leptodactylid Frogs in Mexico + +Author: John D. Lynch + +Release Date: March 30, 2010 [EBook #31830] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK LEPTODACTYLID FROGS IN MEXICO *** + + + + +Produced by Chris Curnow, Joseph Cooper, Diane Monico, and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS +MUSEUM OF NATURAL HISTORY + +Volume 17, No. 11, pp. 503-515, 5 figs. +March 20, 1968 + + + +Genera of Leptodactylid Frogs in Mexico + + +BY + +JOHN D. LYNCH + + +UNIVERSITY OF KANSAS +LAWRENCE +1968 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Frank B. Cross + + +Volume 17, No. 11, pp. 503-515, 5 figs. +Published March 20, 1968 + + +UNIVERSITY OF KANSAS +Lawrence, Kansas + + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1968 + +31-9418 + + + + +Genera of Leptodactylid Frogs in Mexico + +BY JOHN D. LYNCH + + +INTRODUCTION + +According to the most recent review of the Mexican amphibian fauna +(Smith and Taylor, 1948), six genera of leptodactylid frogs occur in +Mexico. One other genus, _Pleurodema_, occurs in Lower Central America. +Smith and Taylor recognized one species of _Engystomops_, 28 of +_Eleutherodactylus_, three of _Leptodactylus_, eight of +_Microbatrachylus_, 12 of _Syrrhophus_, and five of _Tomodactylus_. +Subsequent to the publication of their checklist of the Mexican +amphibia (1948), numerous taxonomic changes have been proposed. Many +species of _Eleutherodactylus_ have been added to the fauna, either +through the extension of their recorded ranges into Mexico from +Guatemala or by the recognition of species unknown in 1948, whereas +some nominal species have been synonymized. _Microbatrachylus_ has been +regarded as synonymous with _Eleutherodactylus_ (Lynch, 1965); four +species of _Microbatrachylus_ currently are regarded as valid +(Duellman, 1961, Lynch, 1965). _Syrrhophus_ was revised in part by +Duellman (1958) and Firschein (1954), and a species of _Tomodactylus_ +transferred to _Syrrhophus_ by Dixon (1957), who redefined +_Tomodactylus_ and added more species to the genus. + +Since beginning my studies of the Mexican leptodactylids in 1962, I +have become acutely aware of difficulties involved in defining the +genera. A revision of _Eleutherodactylus_ and a review of _Syrrhophus_ +are nearing completion, but prior to their publication it is desirable +to redefine the genera of the Mexican leptodactylids, and in so doing +recognize an heretofore unnamed genus. The definitions of +_Eleutherodactylus_ and _Leptodactylus_ may need to be altered in the +future, since both are widespread in South America and occur in the +West Indies. Their definitions as given here are as precise as present +knowledge permits. _Syrrhophus_ and _Tomodactylus_ are small +assemblages that occur only in southwestern United States, Mexico, and +Guatemala. + +Taylor (1952) synonymized _Engystomops_ with _Eupemphix_ which, +although related, should be regarded as generically distinct (Gallardo, +1965). Perhaps the most conservative classification is that of Myers +(1962) who, without published evidence, combined _Eleutherodactylus_, +_Syrrhophus_, and the South American _Lithodytes_ in a single genus. + +The major problem for students working with the Mexican leptodactylids +has not been the separation of _Engystomops_ or _Leptodactylus_ from +other genera but the separation and definition of the eleutherodactyline +frogs currently placed in three genera, _Eleutherodactylus_, +_Syrrhophus_, and _Tomodactylus_. As will be shown in this paper, these +are more conveniently placed in four genera. Once a fourth genus is +recognized, certain phylogenetic problems disappear and a reasonable +zoogeographic interpretation is possible for Middle American +leptodactylid distribution. + + + + +ANALYSIS OF CHARACTERS + + +In Mexico and northern Central America approximately 55 species of +eleutherodactyline frogs (_Eleutherodactylus_, _Syrrhophus_, and +_Tomodactylus_) are known. Four genera can be recognized on the basis +of the nature of inguinal glands, morphology of the hands and feet, and +certain osteological features. + +[Illustration: FIG. 1. _Tomodactylus angustidigitorum_ (UMMZ 114305, +x 4.5) illustrating the lumbo-inguinal gland typical of members of the +genus. From a kodachrome by Wm. E. Duellman.] + + +Glands + +Leptodactylids have a variety of glands that have been used as generic +characters. Smith and Taylor (1948) regarded the so-called inguinal +gland as a generic character in Mexican eleutherodaycty-lines. Lynch +(1965) showed that _Eleutherodactylus_ and _Microbatrachylus_ cannot +be separated by the nature of the gland or the condition of the +prevomers (dentate or not). _Syrrhophus_ and _Tomodactylus_, as defined +by Smith and Taylor (1948), are not generically distinct because of +overlap in the condition of the prevomers and in the development of the +gland. Firschein (1954) stated that _Syrrhophus_ differed from +_Tomodactylus_ by having an axillary gland, but it is now known that +one species of _Syrrhophus_ lacks the gland. + +The inguinal glands of _Eleutherodactylus_ and _Syrrhophus_, if +present, are diffuse, irregular in outline, and generally not +prominent; in _Tomodactylus_ the gland is higher on the body (a +lumbo-inguinal gland), compact, oval in outline, and prominent (Fig. +1). Axillary glands occur in most _Syrrhophus_ but are not known in +_Tomodactylus_ or _Eleutherodactylus_. + + +Hands and feet + +The tips of the digits are laterally expanded in most +_Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_. Two species +of _Eleutherodactylus_ (_augusti_ and _tarahumarensis_) and two +_Tomodactylus_ (_angustidigitorum_ and _grandis_) lack any expansion +of the digital tips. All but two of the species of eleutherodactyline +frogs (_E. augusti_ and _E. tarahumarensis_) have a transverse groove +across the tips of the digits (Fig. 2). + +[Illustration: FIG. 2. Palmar views of the hands and lateral views of +the tip of the third digits of _Eleutherodactylus alfredi_ (left, KU +93994, x 5) and _Hylactophryne augusti_ (right, KU 102594, x 3).] + +Supernumerary tubercles rarely are present on the feet of +_Eleutherodactylus_, but are present and numerous in every species of +_Syrrhophus_, _Tomodactylus_, and in the members of the _augusti_ group +of _Eleutherodactylus_ (Fig. 3). The tubercles are small and numerous +in _Syrrhophus_ and larger in _Tomodactylus_ and the _Eleutherodactylus +augusti_ group. Most species of _Eleutherodactylus_ have no plantar +supernumerary tubercles; a few species have such tubercles, which never +extend between the metatarsal tubercles as in _Syrrhophus_ and +_Tomodactylus_. + +[Illustration: FIG. 3. Plantar views of feet of _Eleutherodactylus +alfredi_ (left, KU 93994, x 4.5), _Syrrhophus pipilans nebulosus_ +(middle, KU 58900, x 7.5), and _Hylactophryne augusti_ (right, KU +102594, x 3) showing differences in size and arrangement of +supernumerary tubercles.] + +Tarsal folds and tubercles are lacking in _Syrrhophus_, _Tomodactylus_, +and the _augusti_ group of _Eleutherodactylus_. Several species of +_Eleutherodactylus_ lack tarsal folds and tubercles, but in nearly +every species group, one or more species possess either an inner tarsal +fold, inner tarsal tubercle(s), or outer tarsal tubercles. + +The terminal phalanges of _Syrrhophus_, _Tomodactylus_, and all +_Eleutherodactylus_ (except the frogs of the _augusti_ group) are +distinctly T-shaped. In the latter, the bones are knob-shaped distally +(Fig. 4). T-shaped terminal phalanges also are present in _Lithodytes_ +and _Trachyphrynus_ but not in other leptodactylid genera. At least one +species of _Eupsophus_ (_E. quixensis_) has terminal phalanges that +resemble those of the _Eleutherodactylus augusti_ group. Several +species of _Eleutherodactylus_, _Syrrhophus_, and _Tomodactylus_ with +slender fingers have T-shaped terminal phalanges although the terminal +dilations proportionately are only scarcely wider than the finger tips +in the _Eleutherodactylus augusti_ group. The presence of a terminal +groove at the tip of the finger is an external indicator of the +T-shaped terminal phalanges. + +[Illustration: FIG. 4. Terminal phalanges of four leptodactylid frogs +(all x 13.5). (a) _Eleutherodactylus mexicanus_, KU 55593; (b) +_Eupsophus roseus_, KU 84731; (c) _Eupsophus quixensis_, UIMNH 59643; +and (d) _Hylactophryne augusti_, KU 56192.] + + +Skull + +All Mexican eleutherodactyline frogs have quadratojugal-maxillary +articulations, completely roofed skulls in adults, median contact of +the nasals, separated occipital condyles, and large prevomers. The +premaxillae of all species are visible when the skulls are viewed from +directly above. The pterygoid lacks a medioventral flange and does not +meet the palatine. In no species is the anterior arm of the squamosal +in contact with the maxillary. Of the numerous species examined (30 +_Eleutherodactylus_, four _Syrrhophus_, and four _Tomodactylus_), the +species in the _Eleutherodactylus augusti_ group are unique in having a +sphenethmoid with a blunt anterior edge. + + +Pectoral Girdle + +All species have large cartilaginous plates in the pectoral girdles; +none possesses a bony style. No divergent modifications of the clavicle +and coracoid bones are known in the family. + + + + +GENERIC ACCOUNTS + + +Genus ~Eleutherodactylus~ Dumeril and Bibron, 1841 + + _Type-species._--_Hylodes martinicensis_ Tschudi, 1838 + + _Diagnosis and definition._--Small to large frogs (12 to 110 + mm. snout-vent length) having slightly to widely expanded + digital pads, each pad bearing a terminal transverse groove; + lumbo-inguinal, inguinal, and axillary glands absent, or if + present, diffuse, irregular in outline, not compact; plantar + supernumerary tubercles absent, or if present, six or fewer, + restricted to distal area of plantar surface, and not + extending between metatarsal tubercles; tarsus bearing inner + or outer tubercles or folds or not; toes free to one-half + webbed; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + contact; anterior arm of squamosal not in contact with + maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + dentigerous processes present or not, dentate or not; + maxillary and premaxillary bones dentate; occipital condyles + separated; development direct. + + _Composition._--About 420 names have been applied to frogs + of this genus; many of these names are synonyms, and many + other species remain undescribed and unnamed. Perhaps the + genus contains 350 species. Thirty-one species occur in + Mexico and northern Central America. + + _Distribution._--From Tamaulipas and Sinaloa, Mexico, + exclusive of the Mexican Plateau, to at least Peru and + southernmost Brazil and throughout the West Indies. + Introduced into Florida. + + _Etymology._--Greek (_eleuthero_ + _dactylus_) meaning + free-toed. + + +Genus ~Engystomops~ Jimenez de la Espada, 1872 + + _Type species._--_Engystomops petersi_ Jimenez de la Espada, + 1872 + + _Diagnosis and definition._--Small frogs (20 to 40 mm. + snout-vent length) having undilated digital tips lacking + transverse grooves; lumbo-inguinal or inguinal glands + absent; plantar supernumerary tubercles present, extending + between metatarsal tubercles; tarsus bearing spinelike + tubercle on inner edge; toes free; terminal phalanges + pointed; sternum bearing bony style; spenethmoid not + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers moderate in size, lacking teeth; + maxillary and premaxillary bones edentate; occipital + condyles separated; tadpole free living. + + _Composition._--Four nominal species (_E. petersi_, _E. + pustulatus_, _E. pustulosus_ and _E. schereri_). + + _Distribution._--Central Veracruz and eastern Oaxaca, + Mexico, to Trinidad, Bolivia, and Peru, east of the Andes. + + _Etymology._--Greek (_engys_ + _stoma_) meaning narrow-mouthed. + + +Genus ~Hylactophryne~ new genus + + _Type-species._--_Hylodes augusti_ Duges, 1879 + + _Diagnosis and definition._--Medium to large frogs (37 to 94 + mm. snout-vent length) having undilated digital tips lacking + terminal grooves; lumbo-inguinal or inguinal glands absent; + plantar supernumerary tubercles present, prominent, + extending to but not between metatarsal tubercles; tarsus + lacking tubercles or folds; toes free of webbing; terminal + phalanges knob-shaped, lacking elongate lateral expansions; + sternum cartilaginous, lacking bony style; sphenethmoid + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers large, bearing dentigerous + processes; maxillary and premaxillary bones dentate; + occipital condyles separated; development direct. + + _Composition._--Two species, _H. augusti_ and _H. + tarahumarensis_, the former composed of four subspecies + (Zweifel, 1956). + + _Distribution._--From Arizona, New Mexico, and Texas to + Guerrero and Puebla, Mexico, and a relict population on + Cerro Quingola (just west of the Isthmus of Tehuantepec, + Mexico). + + _Etymology._--Greek (_hylactor_ + _phryne_) meaning barking + toad; in reference to the voice and common name. + + +Genus ~Leptodactylus~ Fitzinger, 1826 + + _Type-species._--_Leptodactylus typhonia_ Fitzinger, 1826 + + _Diagnosis and definition._--Small to large frogs (30 to + about 200 m., snout-vent length) having undilated to + slightly expanded digital tips bearing pads, no transverse + groove at tips of digits; lumbo-inguinal, axillary, and/or + ventral glands present or not, low, diffuse; plantar + supernumerary tubercles generally absent, if present not + extending between metatarsal tubercles; tarsus bearing + tarsal folds or not; toes free of webbing, extensive lateral + fringes present in some species; terminal phalanges pointed, + not T-shaped; sternum bearing bony style; sphenethmoid not + truncate anteriorly; nasals in contact medially; maxillary + and quadratojugal in articular contact; anterior arm of + squamosal not in contact with maxillary; dermal cranial + elements not involved in integumentary-cranial + co-ossification; prevomers large, bearing dentigerous + processes; maxillary and premaxillary bones dentate; + occipital condyles separated; tadpole free living. + + _Composition._--Sixty species according to Smith and Taylor + (1948); 54 according to Gorham (1963); Argentinian authors + have described several more in recent years. + + _Distribution._--Southern Sonora, Mexico, and southern Texas + throughout the Central and South American lowlands to + Argentina. Also known from Hispaniola and Puerto Rico in the + Greater Antilles and a few islands in the Lesser Antilles. + + _Entymology._--Greek (_leptos_ + _dactylus_) meaning slender + toes. + + +Genus ~Syrrhophus~ Cope, 1878 + + _Type-species._--_Syrrhophus marnockii_ Cope, 1878 + + _Diagnosis and definition._--Small to medium sized frogs (18 + to 40 mm. snout-vent) having slight to prominent digital + expansions with transverse groove at tip of each digit; + lumbo-inguinal and inguinal gland flattened, irregular in + outline, not compact and oval; axillary glands present or + not; plantar supernumerary tubercles numerous, more than + eight, usually extending between metatarsal tubercles; + tarsus lacking tubercles or folds; toes free or basally + webbed; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + articular contact; anterior arm of squamosal not in contact + with maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + usually lacking dentigerous processes and teeth; maxillary + and premaxillary bones dentate; occipital condyles + separated; development direct. + + _Composition._--Thirteen species; the species described as, + or later referred to, _Syrrhophus_ from Lower Central + America and South America are _Eleutherodactylus_ or + _Eupsophus_. + + _Distribution._--Low to moderate elevations from Sinaloa, + Mexico, to Guatemala on the Pacific versant; from the + Edwards and Stockton plateaus of Texas to British Honduras + on the Caribbean versant. + + _Etymology._--Greek, emendation of _syrrhaptos_, meaning + sewn together in reference to the united outer metatarsals. + + +Genus ~Tomodactylus~ Guenther, 1900 + + _Type-species._--_Tomodactylus amulae_ Guenther, 1900. + + _Diagnosis and definition._--Small frogs (20 to 35 mm. + snout-vent length) having digital expansions or not, with + transverse groove across tip of each digit; lumbo-inguinal + gland prominently elevated, compact, oval, often patterned; + axillary glands absent; plantar supernumerary tubercles + numerous, more than eight, usually extending between + metatarsal tubercles; tarsus lacking tubercles or folds; toes + free; terminal phalanges T-shaped; sternum cartilaginous, + lacking bony style; sphenethmoid not truncate anteriorly; + nasals in contact medially; maxillary and quadratojugal in + articular contact; anterior arm of squamosal not in contact + with maxillary; dermal cranial elements not involved in + integumentary-cranial co-ossification; prevomers large, + usually bearing dentigerous processes; maxillary and + premaxillary bones dentate; occipital condyles separated; + development direct. + + _Composition._--Ten species. + + _Distribution._--The southern edge of the Mexican Plateau + from Sinaloa to Veracruuz and onto the Oaxaca highlands and + Sierra Madre del Sur. + + _Etymology._--Greek (_tomis_ + _dactylus_) meaning knife + toe; in reference to either the sharp subarticular tubercles + or the unwebbed toes. + + + + +DISCUSSION + + +The preceding definitions only slightly alter the present generic +limits of Mexican leptodactylids. Two species, previously regarded as +_Eleutherodactylus_, are transferred to the new genus _Hylactophryne_. +The arrangement of the species of _Syrrhophus_ and _Tomodactylus_ +remains the same as concluded by Dixon (1957), Duellman (1958), and +Firschein (1954) in their reviews of the genera. + +Lumbo-inguinal glands are most prominent in the genera _Pleurodema_ and +_Tomodactylus_. Various nondescript glands are present in many genera, +but none is so well developed as those of _Pleurodema_ and +_Tomodactylus_. + +At least nine leptodactylid genera are either known or thought to be +terrestrial breeders lacking a free-living tadpole stage +(_Eleutherodactylus_, _Euparkerella_, _Hylactophryne_, _Niceforonia_, +_Noblella_, _Sminthillus_, _Syrrhophus_, _Tomodactylus_ and +_Trachyphrynus_). _Niceforonia_ and _Trachyphrynus_, and probably +_Hylactophryne_, are not closely related to the other genera. Direct +development probably is an adaptation to adverse environmental +conditions since many of the species occur in semi-arid or cold (Andean +paramos) areas. _Eleutherodactylus_ is generally thought to be the +stock from which _Euparkerella_, _Noblella_, and _Sminthillus_ evolved +(Griffiths, 1959) and from which _Syrrhophus_ and _Tomodactylus_ are +derived (Firschein, 1954). + +The present distribution of _Hylactophryne_ (isolated on the Mexican +Plateau) and its digital form (like that of Papuan and many primitive +South American leptodactylids) suggest that the genus was isolated in +Mexico throughout the Tertiary, whereas the other Central American +genera are either post-Pliocene derivatives of _Eleutherodactylus_ or +invaders of Central America from South America since the mid-Pliocene +land bridge was formed (Lloyd, 1963). + +Piatt (1934) presented arguments against assigning _Eleutherodactylus +latrans_ to the genus _Lithodytes_ and concluded that it was a "true" +_Eleutherodactylus_. Contrary to his arguments, _latrans_ (= _augusti_ +of Zweifel) and _E. tarahumarensis_ Taylor differ from all other +_Eleutherodactylus_ (and _Syrrhophus_ and _Tomodactylus_) in the nature +of the tips of the digits (external and skeletal). The digits of +_Hylactophryne_ are like those of _Eupsophus_. My study of nearly all +genera of leptodactylids indicates that Noble (1925) was correct in +suggesting that _Borborocoetes_ (= _Eupsophus_) is a close relative of +_Eleutherodactylus latrans_, although Noble's arguments were based in +part upon false evidence concerning the breeding habits of _E. +latrans_, then thought to have a free-living tadpole. + +Kellogg (1932) and Piatt (1934) argued that the terminal phalanges of +_E. latrans_ were typically eleutherodactyline. The variation of this +character in _Eupsophus_ (see Fig. 4) ranges from knobbed to bifurcate +or Y-shaped (T-shaped in _Eleutherodactylus_, _Syrrhophus_ and +_Tomodactylus_) and encompasses the nature of the character represented +in _Hylactophryne_. _Eupsophus_ differs from _Hylactophryne_ in +possessing a frontoparietal fontanelle, in generally having a +maxillary-quadratojugal gap, and in having a free swimming tadpole +stage. + +[Illustration: FIG. 5. Outline drawings of _Leptodactylus melanonotus_ +(left, KU 65704, x 2) and _Eleutherodactylus alfredi_ (right, KU 93994, +x 2).] + + + + +KEY TO MEXICAN LEPTODACTYLID GENERA + + +1. Small (20-40 mm.), pustular, toadlike frogs; + maxillary and premaxillary bones not bearing teeth _Engystomops_ + + Large (20-110 mm.), smooth skinned and non-toadlike + frogs; maxillary and premaxillary bones bearing teeth 2 + +2. No conspicuous waist (Fig. 5); sternum bearing + bony style, _Leptodactylus_ + + Constrictions at waist (Fig. 5); sternum + cartilaginous, no bony style 3 + +3. Few (less than six), if any, supernumerary + tubercles on plantar surface _Eleutherodactylus_ + + Many (more than 8) supernumerary tubercles on + plantar surfaces 4 + +4. Terminal, transverse groove across tip of + digits, especially outer two fingers, digits + expanded or not; small frogs (18 to 40 mm.) 5 + + Tips of digits lacking transverse groove; + digits unexpanded; medium-sized to large frogs + (37 to 94 mm.) _Hylactophryne_ + +5. Lumbo-inguinal gland compact, oval _Tomodactylus_ + + Lumbo-inguinal or inguinal gland absent or + diffuse and irregular in outline _Syrrhophus_ + + + + +LITERATURE CITED + + +DIXON, J. R. + + 1957. Geographic variation and distribution of the genus + _Tomodactylus_ in Mexico. Texas Jour. Sci., 9:379-409, + December. + +DUELLMAN, W. E. + + 1958. A review of the frogs of the genus Syrrhophus in + western Mexico. Occas. Papers Mus. Zool. Univ. + Michigan, 594:1-15, June 6. + + 1961. The amphibians and reptiles of Michoacan, Mexico. Univ. + Kansas Publ. Mus. Nat. Hist., 15:1-148, December 20. + +FIRSCHEIN, I. L. + + 1954. Definition of some little understood members of the + leptodactylid genus _Syrrhophus_, with a description + of a new species. Copeia, 1:48-58, February 19. + +GALLARDO, J. M. + + 1965. A proposito de los Leptodactylidae (Amphibia Anura). + Papeis Avulsos, 17:77-87, January 30. + +GORHAM, S. W. + + 1963. The comparative number of species of amphibians in + Canada and other countries. III. Summary of species + of anurans. Canadian Field-Nat., 77:13-48, March. + +GRIFFITHS, I. + + 1959. The phylogeny of Sminthillus limbatus and the status of + the Brachycephalidae (Amphibia Salientia). Proc. Zool. + Soc. London, 132:457-87, May. + +KELLOGG, R. + + 1932. Mexican tailless amphibians in the United States + National Museum. Bull. U. S. Natl. Mus., 160:224 pp., + March 31. + +LLOYD, J. J. + + 1963. Tectonic history of the south Central-American orogen, + _in_ Childs and Beebe eds., Backbone of the Americas. + Amer. Assoc. Petroleum Geol., pp. 88-100. + +LYNCH, J. D. + + 1965. A review of the eleutherodactylid frog genus + _Microbatrachylus_ (Leptodactylidae). Nat. Hist. Misc., + 182:1-12, December 15. + +MYERS, G. S. + + 1962. The American leptodactylid frog genus _Eleutherodactylus_, + _Hylodes_ (= _Elosia_), and _Caudiverbera_ + (= _Calytocephalus_). Copeia, 1:195-202, April 11. + +NOBLE, G. K. + + 1925. An outline of the relation of the ontogeny to + phylogeny within the Amphibia. I. Amer. Mus. Nov., + 165:1-17, April 16. + +PIATT, J. + + 1934. The systematic status of Eleutherodactylus latrans + (Cope). Amer. Midl. Nat., 15:89-91, February 15. + +SMITH, H. M. and TAYLOR, E. H. + + 1948. An annotated checklist and key to the Amphibia of + Mexico. Bull. U. S. Natl. Mus., 194:1-118. June 7. + +TAYLOR, E. H. + + 1952. A review of the frogs and toads of Costa Rica. Univ. + Kansas Sci. Bull., 35:577-942, July 1. + +ZWEIFEL, R. G. + + 1956. A survey of the frogs of the _augusti_ group, genus + _Eleutherodactylus_. Amer. Mus. Novitates, 1813:1-35, + December 23. + + +_Transmitted July 11, 1967._ + + +31-9418 + + + + + * * * * * + +Transcriber's Notes + + +Bold text is shown within ~tildes~. + +Italicized text is shown within _underscores_. + +Illustrations have been moved to avoid breaking up paragraphs of text. + +Page 507: Changed know to known (are not know in Tomodactylus). + +Page 507: Added closing parenthesis in Fig. 2 caption after x 3. + +Page 509: Changed compeltely to completely (compeltely roofed skulls). + +Page 512: Veracruuz may be a typo for Veracruz (Sinaloa to Veracruuz). + +Page 514: Changed two occurrences of Hylatophryne to Hylactophryne. + + + + + + + + +End of the Project Gutenberg EBook of Genera of Leptodactylid Frogs in Mexico, by +John D. 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