summaryrefslogtreecommitdiff
path: root/31830-8.txt
diff options
context:
space:
mode:
authorRoger Frank <rfrank@pglaf.org>2025-10-14 19:56:30 -0700
committerRoger Frank <rfrank@pglaf.org>2025-10-14 19:56:30 -0700
commit397660dadbf82c77860660740b16d8e336da7999 (patch)
tree44a5982f4af5a816e81a5bb0ce03f9d9f414c4f1 /31830-8.txt
initial commit of ebook 31830HEADmain
Diffstat (limited to '31830-8.txt')
-rw-r--r--31830-8.txt1062
1 files changed, 1062 insertions, 0 deletions
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. Lynch
+
+*** END OF THIS PROJECT GUTENBERG EBOOK LEPTODACTYLID FROGS IN MEXICO ***
+
+***** This file should be named 31830-8.txt or 31830-8.zip *****
+This and all associated files of various formats will be found in:
+ https://www.gutenberg.org/3/1/8/3/31830/
+
+Produced by Chris Curnow, Joseph Cooper, Diane Monico, and
+the Online Distributed Proofreading Team at
+https://www.pgdp.net
+
+
+Updated editions will replace the previous one--the old editions
+will be renamed.
+
+Creating the works from public domain print editions means that no
+one owns a United States copyright in these works, so the Foundation
+(and you!) can copy and distribute it in the United States without
+permission and without paying copyright royalties. Special rules,
+set forth in the General Terms of Use part of this license, apply to
+copying and distributing Project Gutenberg-tm electronic works to
+protect the PROJECT GUTENBERG-tm concept and trademark. Project
+Gutenberg is a registered trademark, and may not be used if you
+charge for the eBooks, unless you receive specific permission. If you
+do not charge anything for copies of this eBook, complying with the
+rules is very easy. You may use this eBook for nearly any purpose
+such as creation of derivative works, reports, performances and
+research. They may be modified and printed and given away--you may do
+practically ANYTHING with public domain eBooks. Redistribution is
+subject to the trademark license, especially commercial
+redistribution.
+
+
+
+*** START: FULL LICENSE ***
+
+THE FULL PROJECT GUTENBERG LICENSE
+PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK
+
+To protect the Project Gutenberg-tm mission of promoting the free
+distribution of electronic works, by using or distributing this work
+(or any other work associated in any way with the phrase "Project
+Gutenberg"), you agree to comply with all the terms of the Full Project
+Gutenberg-tm License (available with this file or online at
+https://gutenberg.org/license).
+
+
+Section 1. General Terms of Use and Redistributing Project Gutenberg-tm
+electronic works
+
+1.A. By reading or using any part of this Project Gutenberg-tm
+electronic work, you indicate that you have read, understand, agree to
+and accept all the terms of this license and intellectual property
+(trademark/copyright) agreement. If you do not agree to abide by all
+the terms of this agreement, you must cease using and return or destroy
+all copies of Project Gutenberg-tm electronic works in your possession.
+If you paid a fee for obtaining a copy of or access to a Project
+Gutenberg-tm electronic work and you do not agree to be bound by the
+terms of this agreement, you may obtain a refund from the person or
+entity to whom you paid the fee as set forth in paragraph 1.E.8.
+
+1.B. "Project Gutenberg" is a registered trademark. It may only be
+used on or associated in any way with an electronic work by people who
+agree to be bound by the terms of this agreement. There are a few
+things that you can do with most Project Gutenberg-tm electronic works
+even without complying with the full terms of this agreement. See
+paragraph 1.C below. There are a lot of things you can do with Project
+Gutenberg-tm electronic works if you follow the terms of this agreement
+and help preserve free future access to Project Gutenberg-tm electronic
+works. See paragraph 1.E below.
+
+1.C. The Project Gutenberg Literary Archive Foundation ("the Foundation"
+or PGLAF), owns a compilation copyright in the collection of Project
+Gutenberg-tm electronic works. Nearly all the individual works in the
+collection are in the public domain in the United States. If an
+individual work is in the public domain in the United States and you are
+located in the United States, we do not claim a right to prevent you from
+copying, distributing, performing, displaying or creating derivative
+works based on the work as long as all references to Project Gutenberg
+are removed. Of course, we hope that you will support the Project
+Gutenberg-tm mission of promoting free access to electronic works by
+freely sharing Project Gutenberg-tm works in compliance with the terms of
+this agreement for keeping the Project Gutenberg-tm name associated with
+the work. You can easily comply with the terms of this agreement by
+keeping this work in the same format with its attached full Project
+Gutenberg-tm License when you share it without charge with others.
+
+1.D. The copyright laws of the place where you are located also govern
+what you can do with this work. Copyright laws in most countries are in
+a constant state of change. If you are outside the United States, check
+the laws of your country in addition to the terms of this agreement
+before downloading, copying, displaying, performing, distributing or
+creating derivative works based on this work or any other Project
+Gutenberg-tm work. The Foundation makes no representations concerning
+the copyright status of any work in any country outside the United
+States.
+
+1.E. Unless you have removed all references to Project Gutenberg:
+
+1.E.1. The following sentence, with active links to, or other immediate
+access to, the full Project Gutenberg-tm License must appear prominently
+whenever any copy of a Project Gutenberg-tm work (any work on which the
+phrase "Project Gutenberg" appears, or with which the phrase "Project
+Gutenberg" is associated) is accessed, displayed, performed, viewed,
+copied or distributed:
+
+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
+
+1.E.2. If an individual Project Gutenberg-tm electronic work is derived
+from the public domain (does not contain a notice indicating that it is
+posted with permission of the copyright holder), the work can be copied
+and distributed to anyone in the United States without paying any fees
+or charges. If you are redistributing or providing access to a work
+with the phrase "Project Gutenberg" associated with or appearing on the
+work, you must comply either with the requirements of paragraphs 1.E.1
+through 1.E.7 or obtain permission for the use of the work and the
+Project Gutenberg-tm trademark as set forth in paragraphs 1.E.8 or
+1.E.9.
+
+1.E.3. If an individual Project Gutenberg-tm electronic work is posted
+with the permission of the copyright holder, your use and distribution
+must comply with both paragraphs 1.E.1 through 1.E.7 and any additional
+terms imposed by the copyright holder. Additional terms will be linked
+to the Project Gutenberg-tm License for all works posted with the
+permission of the copyright holder found at the beginning of this work.
+
+1.E.4. Do not unlink or detach or remove the full Project Gutenberg-tm
+License terms from this work, or any files containing a part of this
+work or any other work associated with Project Gutenberg-tm.
+
+1.E.5. Do not copy, display, perform, distribute or redistribute this
+electronic work, or any part of this electronic work, without
+prominently displaying the sentence set forth in paragraph 1.E.1 with
+active links or immediate access to the full terms of the Project
+Gutenberg-tm License.
+
+1.E.6. You may convert to and distribute this work in any binary,
+compressed, marked up, nonproprietary or proprietary form, including any
+word processing or hypertext form. However, if you provide access to or
+distribute copies of a Project Gutenberg-tm work in a format other than
+"Plain Vanilla ASCII" or other format used in the official version
+posted on the official Project Gutenberg-tm web site (www.gutenberg.org),
+you must, at no additional cost, fee or expense to the user, provide a
+copy, a means of exporting a copy, or a means of obtaining a copy upon
+request, of the work in its original "Plain Vanilla ASCII" or other
+form. Any alternate format must include the full Project Gutenberg-tm
+License as specified in paragraph 1.E.1.
+
+1.E.7. Do not charge a fee for access to, viewing, displaying,
+performing, copying or distributing any Project Gutenberg-tm works
+unless you comply with paragraph 1.E.8 or 1.E.9.
+
+1.E.8. You may charge a reasonable fee for copies of or providing
+access to or distributing Project Gutenberg-tm electronic works provided
+that
+
+- You pay a royalty fee of 20% of the gross profits you derive from
+ the use of Project Gutenberg-tm works calculated using the method
+ you already use to calculate your applicable taxes. The fee is
+ owed to the owner of the Project Gutenberg-tm trademark, but he
+ has agreed to donate royalties under this paragraph to the
+ Project Gutenberg Literary Archive Foundation. Royalty payments
+ must be paid within 60 days following each date on which you
+ prepare (or are legally required to prepare) your periodic tax
+ returns. Royalty payments should be clearly marked as such and
+ sent to the Project Gutenberg Literary Archive Foundation at the
+ address specified in Section 4, "Information about donations to
+ the Project Gutenberg Literary Archive Foundation."
+
+- You provide a full refund of any money paid by a user who notifies
+ you in writing (or by e-mail) within 30 days of receipt that s/he
+ does not agree to the terms of the full Project Gutenberg-tm
+ License. You must require such a user to return or
+ destroy all copies of the works possessed in a physical medium
+ and discontinue all use of and all access to other copies of
+ Project Gutenberg-tm works.
+
+- You provide, in accordance with paragraph 1.F.3, a full refund of any
+ money paid for a work or a replacement copy, if a defect in the
+ electronic work is discovered and reported to you within 90 days
+ of receipt of the work.
+
+- You comply with all other terms of this agreement for free
+ distribution of Project Gutenberg-tm works.
+
+1.E.9. If you wish to charge a fee or distribute a Project Gutenberg-tm
+electronic work or group of works on different terms than are set
+forth in this agreement, you must obtain permission in writing from
+both the Project Gutenberg Literary Archive Foundation and Michael
+Hart, the owner of the Project Gutenberg-tm trademark. Contact the
+Foundation as set forth in Section 3 below.
+
+1.F.
+
+1.F.1. Project Gutenberg volunteers and employees expend considerable
+effort to identify, do copyright research on, transcribe and proofread
+public domain works in creating the Project Gutenberg-tm
+collection. Despite these efforts, Project Gutenberg-tm electronic
+works, and the medium on which they may be stored, may contain
+"Defects," such as, but not limited to, incomplete, inaccurate or
+corrupt data, transcription errors, a copyright or other intellectual
+property infringement, a defective or damaged disk or other medium, a
+computer virus, or computer codes that damage or cannot be read by
+your equipment.
+
+1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the "Right
+of Replacement or Refund" described in paragraph 1.F.3, the Project
+Gutenberg Literary Archive Foundation, the owner of the Project
+Gutenberg-tm trademark, and any other party distributing a Project
+Gutenberg-tm electronic work under this agreement, disclaim all
+liability to you for damages, costs and expenses, including legal
+fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT
+LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE
+PROVIDED IN PARAGRAPH F3. YOU AGREE THAT THE FOUNDATION, THE
+TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE
+LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR
+INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH
+DAMAGE.
+
+1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a
+defect in this electronic work within 90 days of receiving it, you can
+receive a refund of the money (if any) you paid for it by sending a
+written explanation to the person you received the work from. If you
+received the work on a physical medium, you must return the medium with
+your written explanation. The person or entity that provided you with
+the defective work may elect to provide a replacement copy in lieu of a
+refund. If you received the work electronically, the person or entity
+providing it to you may choose to give you a second opportunity to
+receive the work electronically in lieu of a refund. If the second copy
+is also defective, you may demand a refund in writing without further
+opportunities to fix the problem.
+
+1.F.4. Except for the limited right of replacement or refund set forth
+in paragraph 1.F.3, this work is provided to you 'AS-IS' WITH NO OTHER
+WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO
+WARRANTIES OF MERCHANTIBILITY OR FITNESS FOR ANY PURPOSE.
+
+1.F.5. Some states do not allow disclaimers of certain implied
+warranties or the exclusion or limitation of certain types of damages.
+If any disclaimer or limitation set forth in this agreement violates the
+law of the state applicable to this agreement, the agreement shall be
+interpreted to make the maximum disclaimer or limitation permitted by
+the applicable state law. The invalidity or unenforceability of any
+provision of this agreement shall not void the remaining provisions.
+
+1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation, the
+trademark owner, any agent or employee of the Foundation, anyone
+providing copies of Project Gutenberg-tm electronic works in accordance
+with this agreement, and any volunteers associated with the production,
+promotion and distribution of Project Gutenberg-tm electronic works,
+harmless from all liability, costs and expenses, including legal fees,
+that arise directly or indirectly from any of the following which you do
+or cause to occur: (a) distribution of this or any Project Gutenberg-tm
+work, (b) alteration, modification, or additions or deletions to any
+Project Gutenberg-tm work, and (c) any Defect you cause.
+
+
+Section 2. Information about the Mission of Project Gutenberg-tm
+
+Project Gutenberg-tm is synonymous with the free distribution of
+electronic works in formats readable by the widest variety of computers
+including obsolete, old, middle-aged and new computers. It exists
+because of the efforts of hundreds of volunteers and donations from
+people in all walks of life.
+
+Volunteers and financial support to provide volunteers with the
+assistance they need are critical to reaching Project Gutenberg-tm's
+goals and ensuring that the Project Gutenberg-tm collection will
+remain freely available for generations to come. In 2001, the Project
+Gutenberg Literary Archive Foundation was created to provide a secure
+and permanent future for Project Gutenberg-tm and future generations.
+To learn more about the Project Gutenberg Literary Archive Foundation
+and how your efforts and donations can help, see Sections 3 and 4
+and the Foundation web page at https://www.pglaf.org.
+
+
+Section 3. Information about the Project Gutenberg Literary Archive
+Foundation
+
+The Project Gutenberg Literary Archive Foundation is a non profit
+501(c)(3) educational corporation organized under the laws of the
+state of Mississippi and granted tax exempt status by the Internal
+Revenue Service. The Foundation's EIN or federal tax identification
+number is 64-6221541. Its 501(c)(3) letter is posted at
+https://pglaf.org/fundraising. Contributions to the Project Gutenberg
+Literary Archive Foundation are tax deductible to the full extent
+permitted by U.S. federal laws and your state's laws.
+
+The Foundation's principal office is located at 4557 Melan Dr. S.
+Fairbanks, AK, 99712., but its volunteers and employees are scattered
+throughout numerous locations. Its business office is located at
+809 North 1500 West, Salt Lake City, UT 84116, (801) 596-1887, email
+business@pglaf.org. Email contact links and up to date contact
+information can be found at the Foundation's web site and official
+page at https://pglaf.org
+
+For additional contact information:
+ Dr. Gregory B. Newby
+ Chief Executive and Director
+ gbnewby@pglaf.org
+
+
+Section 4. Information about Donations to the Project Gutenberg
+Literary Archive Foundation
+
+Project Gutenberg-tm depends upon and cannot survive without wide
+spread public support and donations to carry out its mission of
+increasing the number of public domain and licensed works that can be
+freely distributed in machine readable form accessible by the widest
+array of equipment including outdated equipment. Many small donations
+($1 to $5,000) are particularly important to maintaining tax exempt
+status with the IRS.
+
+The Foundation is committed to complying with the laws regulating
+charities and charitable donations in all 50 states of the United
+States. Compliance requirements are not uniform and it takes a
+considerable effort, much paperwork and many fees to meet and keep up
+with these requirements. We do not solicit donations in locations
+where we have not received written confirmation of compliance. To
+SEND DONATIONS or determine the status of compliance for any
+particular state visit https://pglaf.org
+
+While we cannot and do not solicit contributions from states where we
+have not met the solicitation requirements, we know of no prohibition
+against accepting unsolicited donations from donors in such states who
+approach us with offers to donate.
+
+International donations are gratefully accepted, but we cannot make
+any statements concerning tax treatment of donations received from
+outside the United States. U.S. laws alone swamp our small staff.
+
+Please check the Project Gutenberg Web pages for current donation
+methods and addresses. Donations are accepted in a number of other
+ways including including checks, online payments and credit card
+donations. To donate, please visit: https://pglaf.org/donate
+
+
+Section 5. General Information About Project Gutenberg-tm electronic
+works.
+
+Professor Michael S. Hart was the originator of the Project Gutenberg-tm
+concept of a library of electronic works that could be freely shared
+with anyone. For thirty years, he produced and distributed Project
+Gutenberg-tm eBooks with only a loose network of volunteer support.
+
+
+Project Gutenberg-tm eBooks are often created from several printed
+editions, all of which are confirmed as Public Domain in the U.S.
+unless a copyright notice is included. Thus, we do not necessarily
+keep eBooks in compliance with any particular paper edition.
+
+
+Most people start at our Web site which has the main PG search facility:
+
+ https://www.gutenberg.org
+
+This Web site includes information about Project Gutenberg-tm,
+including how to make donations to the Project Gutenberg Literary
+Archive Foundation, how to help produce our new eBooks, and how to
+subscribe to our email newsletter to hear about new eBooks.