Plateau in Michoacan. All specimens from the state have spotted venters.
In living toads the dorsal ground color was gray or grayish tan with
olive green spots. The vocal sac was brownish gray; the iris was a
bright golden color.
On June 11, 1958, many individuals were calling from shallow water in a
flooded field at Emiliano Zapata. The call is a slow trill, in which the
individual notes are discernible.
~Bufo marinus~ (Linnaeus)
_Rana marina_ Linnaeus, Systema naturae, ed. 10, 1:211,
1758.--America.
_Bufo horribilis_ Wiegmann, Isis von Oken, 26:654,
1833.--Misantla and Veracruz, Veracruz, Mexico. Taylor and
Smith, Proc. U. S. Natl. Mus., 95:551, January 30, 1945.
_Bufo angustipes_ Taylor and Smith, Proc. U. S. Natl. Mus.,
95:553, January 30, 1945.--La Esperanza, Chiapas, Mexico.
Aguililla; Apatzingan (3); Barranca de Bejuco; Capirio;
Charapendo; Chichihuas; Coahuayana (2); Coalcoman (7);
Cofradia (2); 25 km. S of Cuatro Caminos; El Sabino (10);
Huahua, La Playa (13); Ojos de Agua de San Telmo; Ostula;
Playa Azul (2); Pomaro (2).
This large toad is characteristically found in areas supporting tropical
scrub forest to elevations of about 1000 meters. The species is much
more abundant than the numbers listed above suggest. In the dry season
individuals have been observed in patios, along streams, and by
irrigation ditches. In the rainy season the loud, rattling call of the
males is heard at night throughout the Tepalcatepec Valley and the
coastal lowlands.
Taylor and Smith (1945:552) revived Wiegmann's _Bufo horribilis_ for the
large toads of Mexico that are here referred to _B. marinus_. Their
action was based upon the supposition that the "species _marinus_" is
composite. Although probably true, this supposition has yet to be
proved. Until the large, and apparently related, species of _Bufo_
inhabiting tropical America have been studied systematically as a unit,
the recognition of segments of the population as either species or
subspecies is meaningless. Taylor and Smith (op. cit.:553) based the
description of a new species, _Bufo angustipes_, on one rather
emaciated, formalin-hardened female from La Esperanza, Chiapas. The type
(USNM 116513), when compared with numerous specimens of _Bufo marinus_
from throughout the range of the species in Mexico and northern Central
America, displays no combination of characters to set it off from the
others. Therefore, I suggest that _Bufo horribilis_ Wiegmann and _Bufo
angustipes_ Taylor and Smith be placed in the synonymy of _Bufo marinus_
(Linnaeus) until future systematic study of the genus and this species
in particular establishes the existence of recognizable taxa.
~Bufo marmoreus~ Wiegmann
_Bufo marmoreus_ Wiegmann, Isis von Oken, 26:66,
1833.--Veracruz, Veracruz, Mexico.
Barranca de Bejuco; Coahuayana (11); El Diezmo (2); La
Placita (9); La Orilla (12); Motin del Oro; Ostula (9);
Playa Azul (5); Pomaro (15); Salitre de Estopilas; San Pedro
Naranjestila.
In Michoacan this species is confined to elevations of less than 1000
meters on the coast and foothills of the Sierra de Coalcoman. In this
region in the months of June and July, breeding congregations have been
found in temporary pools and along streams.
Smith and Taylor (1948:39), in their key to the Mexican species of
_Bufo_, placed emphasis on the nature of the supraorbital and
postorbital crests (whether they form a curve or a sharp angle) in
distinguishing _Bufo marmoreus_ from _Bufo perplexus_. In the original
description of _perplexus_, Taylor (1943a:347) characterized the species
as follows: supraorbital and postorbital crests forming a sharp angle,
instead of a curve as in _marmoreus_; supratympanic crest smaller than
in _marmoreus_; diagonal lateral stripe lacking in females;
concentration of dorsal tubercles as found in _marmoreus_ lacking in
males. The discovery of specimens in which the crests form a curve and
others in which the crests form an angle in both the Tepalcatepec Valley
and in the coastal lowlands prompted an investigation of these
characters and others throughout the ranges of the species. An
examination of 410 specimens has resulted in the following conclusions.
TABLE 1.--VARIATION IN THE SHAPE OF THE SUPRAORBITAL AND POSTORBITAL
CRANIAL CRESTS IN BUFO MARMOREUS AND B. PERPLEXUS.
+------------------------+---+-----------+------------+----------+
|LOCALITY | N | Curved |Intermediate| Angular |
+------------------------+---+-----------+------------+----------+
|Tepalcatepec Valley | 50| 10 (20.0%)| 17 (34.0%) |23 (46.0%)|
+------------------------+---+-----------+------------+----------+
|Morelos | 12| 2 (16.6%)| 5 (41.7%) | 5 (41.7%)|
+------------------------+---+-----------+------------+----------+
|Izucar, Puebla | 4| 2 (50.0%)| 0 (0.0%) | 2 (50.0%)|
+------------------------+---+-----------+------------+----------+
|Southern Sinaloa | 1| 1(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Puerto Vallarta, Jalisco| 2| 2(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Colima | 45| 25 (55.0%)| 18 (40.0%) | 2 (5.0%)|
+------------------------+---+-----------+------------+----------+
|Coast of Michoacan | 55| 35 (63.6%)| 17 (30.9%) | 3 (5.5%)|
+------------------------+---+-----------+------------+----------+
|Acapulco, Guerrero | 7| 7(100.0%)| 0 (0.0%) | 0 (0.0%)|
+------------------------+---+-----------+------------+----------+
|Chilpancingo, Guerrero | 10| 1 (10.0%)| 4 (40.0%) | 5 (50.0%)|
+------------------------+---+-----------+------------+----------+
|Pochutla, Oaxaca | 13| 6 (46.2%)| 6 (46.2%) | 1 (7.6%)|
+------------------------+---+-----------+------------+----------+
|Tehuantepec, Oaxaca |177| 81 (45.8%)| 67 (37.8%) |29 (16.4%)|
+------------------------+---+-----------+------------+----------+
|Tonola, Chiapas | 1| 0 (0.0%)| 0 (0.0%) | 1(100.0%)|
+------------------------+---+-----------+------------+----------+
|Veracruz | 33| 26 (78.8%)| 6 (18.2%) | 1 (3.0%)|
+------------------------+---+-----------+------------+----------+
| Total |410|198 (48.3%)|140 (34.2%) |72 (17.5%)|
+------------------------+---+-----------+------------+----------+
1. Although the highest percentage of individuals having the
supraorbital and postorbital crests forming a sharp angle is
from localities in the Balsas-Tepalcatepec Basin, numerous
individuals from throughout the range of _marmoreus_ have the
crests forming an angle (Table 1).
2. In all samples of ten or more specimens, some toads have
the supraorbital and postorbital crests forming a sharp
angle, some have the crests forming a curve, and some have
an intermediate condition.
3. The relative size of the supratympanic crest is highly
variable in all samples examined.
[Illustration: FIG. 3. Adult male of _Bufo perplexus_ from Apatzingan,
Michoacan. x 1.5.]
[Illustration: FIG. 4. Adult male of _Bufo marmoreus_ from Pomaro,
Michoacan. x 1.5.]
[Illustration: PLATE 1
Hatchling of _Pseudoeurycea belli_ from San Juan de Parangaricutiro,
Michoacan. x 8.]
[Illustration: PLATE 2
FIG. 1. Nest and eggs of _Pseudoeurycea belli_ beneath a
rock at San Juan de Parangaricutiro. Approx. natural size.
FIG. 2. Multiple egg clutches of _Phyllomedusa
dacnicolor_ from Coalcoman, Michoacan. 1/3 x.
[Illustration: PLATE 3
FIG. 1. Adult male of _Tomodactylus angustidigitorum_
from Paracho, Michoacan. x 4.
FIG. 2. Adult male of _Tomodactylus fuscus_ from Los
Cantiles, Michoacan. x4.
[Illustration: PLATE 4
FIG. 1. Adult male of _Tomodactylus nitidus_ nitidus from
Tuxpan, Michoacan. x.
FIG. 2. Adult male of _Tomodactylus nitidus orarius_ from
Tecolapa, Colima. x 4.]
[Illustration: PLATE 5
FIG. 1. Adult male of _Tomodactylus nitidus_ petersi from
Apatzingan, Michoacan. x 4.
Fig. 2. Adult male of _Tomodactylus rufescens_ from Dos
Aguas, Michoacan. x 4.]
[Illustration: PLATE 6
FIG. 1. Adult male of _Hypopachus caprimimus_ from
Tuxpan, Michoacan. x 2-1/2.
FIG. 2. Adult male of _Hypopachus oxyrrhinus ovis_ from
Tangamandapio, Michoacan. x 3.]
4. A distinct, pale-colored, diagonal lateral stripe is
found in females only from localities outside of the
Balsas-Tepalcatepec Basin; females from the basin have a
spotted dorsum.
5. Males from the Balsas-Tepalcatepec Basin usually have a
broad middorsal line that is yellow or pale tan; those from
outside the basin have either a narrow middorsal line or
none.
6. Males from the Balsas-Tepalcatepec Basin have low,
scattered dorsal tubercles (Fig. 3); males from outside the
basin have a concentration of tubercles in a broad band on
the back (Fig. 4).
Therefore the nature of the cranial crests is of little value in
separating two populations, but the color pattern of the females and the
nature of the dorsal tubercles of the males do show distinct
differences. Furthermore, certain differences in size and proportion are
evident; _Bufo marmoreus_ is a slightly larger toad and has a relatively
longer tibia and longer head than _perplexus_ (Table 2).
TABLE 2.--COMPARISON OF CERTAIN MEASUREMENTS AND PROPORTIONS IN BUFO
MARMOREUS AND B. PERPLEXUS. (MEANS ARE GIVEN IN PARENTHESES BELOW THE
RANGES.)
+-------------------+------------+--+----------+------------+-----------+
| | | | |Tibia length|Head length|
| Species | Sex | N|Snout-vent+------------+-----------+
| | | | length | Snout-vent |Snout-vent |
| | | | | length | length |
+-------------------+------------+--+----------+------------+-----------+
| _B. marmoreus_ | Male |15|61.5-72.5 | 35.9-41.6 | 28.3-33.3 |
| | | | (65.2) | (39.0) | (31.6) |
+-------------------+------------+--+----------+------------+-----------+
| _B. perplexus_ | Male |20|50.0-59.0 | 33.7-38.1 | 26.4-31.1 |
| | | | (54.9) | (36.4) | (29.5) |
+-------------------+------------+--+----------+------------+-----------+
| _B. marmoreus_ | Female | 7|68.0-76.0 | 33.0-36.8 | 26.8-32.6 |
| | | | (70.7) | (34.7) | (29.6) |
+-------------------+------------+--+----------+------------+-----------+
| _B. perplexus_ |Female | 6|64.1-69.8 | 32.4-36.9 | 25.1-29.0 |
| | | | (66.8) | (35.5) | (27.5) |
+-------------------+------------+--+----------+------------+-----------+
Taylor (1943a:347) described _Bufo perplexus_ from Mexcala on the Rio
Balsas in Guerrero. Among the many paratypes are specimens from Tonola,
Chiapas, and Tehuantepec, Oaxaca. These apparently were referred to
_perplexus_ solely on the nature of the cranial crests. All of the
specimens examined during the course of the present study from the
lowlands of Veracruz and from the Pacific lowlands from Sinaloa
southward to Chiapas are referable to _Bufo marmoreus_; those from the
Balsas-Tepalcatepec Basin are referable to _Bufo perplexus_, as defined
above. Ten specimens from Chilpancingo, Guerrero (UMMZ 115352), do not
readily fit either species. Perhaps there is gene exchange between the
inland and coastal populations through the relatively low pass at
Chilpancingo, at the mouth of the Rio Balsas, and near the convergent
headwaters of the Rio Coahuayana and Rio Tepalcatepec in southern
Jalisco. If this can be demonstrated, then _Bufo perplexus_ would have
to be considered as a subspecies of _Bufo marmoreus_, instead of an
allopatric species.
~Bufo perplexus~ Taylor
_Bufo perplexus_ Taylor, Univ. Kansas Sci. Bull., 29:347,
October 15, 1943.--Balsas River near Mexcala, Guerrero,
Mexico.
Aguililla (2); Apatzingan (42); Buena Vista (5); Capirio
(3); La Playa (25); Lombardia (6); Nueva Italia (9); Rio
Cancita, 14 km. E of Apatzingan; Rio Tepalcatepec, 27 km. S
of Apatzingan; San Salvador (4); Tzitzio; Volcan Jorullo.
~Bufo occidentalis~ Camerano
_Bufo occidentalis_ Camerano, Atti R. Accad. Sci. Torino,
14:887, December 31, 1878.--Mexico. Type locality restricted
to Guanajuato, Guanajuato, Mexico, by Smith and Taylor
(1950a:330). Firschein, Copeia, no. 3:220, September 15,
1950.
_Bufo simus_, Smith and Taylor, Bull. U. S. Natl. Mus.,
194:42, 1948.
Barranca Seca (32); Cerro de Barolosa (4); Cerro Tancitaro,
3 km. E of Apo (2); Cerro Tancitaro, 19 km. E. of Apo (10);
Charapendo; Coalcoman (7); Dos Aguas (4); Jacona, Jaramillo
(2); Las Tecatas; Los Reyes (181); Tancitaro (10); Uruapan
(3).
This toad is an inhabitant of pine and oak forests between 900 and 2400
meters. Near Charapendo on the slopes of the Sierra de los Tarascos and
at Coalcoman it apparently reaches its lowest altitudinal limits. At
both of these localities the pine-oak forest is replaced by arid
tropical scrub forest on the lower slopes.
Twenty-four tadpoles were collected on May 3 in a quiet section of a
fast stream near Barranca Seca. The tadpoles have a robust body,
broadest about two-thirds the distance from the snout to the posterior
edge of the body, half again as broad as deep. Eyes dorsolateral;
nostrils dorsal, somewhat directed forward, and about three-fifths the
distance from the tip of the snout to the eye; spiracle sinistral and
lateral, located at about midbody; anus median; tail long and slender;
tail-musculature extends nearly to tip of tail; depth of
tail-musculature at mid-length about one-third total depth of tail;
dorsal tail-fin not extending onto body (Fig. 5); average body length of
ten tadpoles having small hind limb buds, 14.4 mm.; average tail length,
22.0 mm.
Mouth ventral, nearly terminal, about one-third as wide as widest part
of body; anterior lip has no papillae; lower lip bordered by two rows of
papillae and lateral lips by one row of papillae; beaks moderately well
developed, the upper forming a broad arch and finely denticulate; tooth
rows 2/3, the upper rows extending to the edge of the lips, subequal in
length, and slightly longer than lower rows, which also are subequal in
length; inner upper tooth row broken medially; inner lower tooth row
sometimes broken (Fig. 6).
The body is black dorsally and laterally, and bluish gray ventrally; the
tail musculature is brown and stippled with darker brown. The fins are
transparent and stippled with brown, the stippling being most pronounced
on the posterior two-thirds of the upper tail-fin.
[Illustration: FIG. 5. Tadpole of _Bufo occidentalis_ (UMMZ 94269) from
Barranca Seca, Michoacan. x 3.]
[Illustration: FIG. 6. Mouthparts of larval _Bufo occidentalis_ (UMMZ
94269) from Barranca Seca, Michoacan. x 20.]
Forty recently metamorphosed individuals average 18.9 mm. in snout-vent
length.
The relationships of this toad seem to be with _Bufo bocourti_ Brocchi,
an inhabitant of pine and oak forests in the uplands of Chiapas and
Guatemala. In _Bufo occidentalis_ the tympanum usually is indistinct and
sometimes completely covered, and it is absent in _bocourti_. _Bufo
occidentalis_ has a broader interorbital area and relatively shorter and
more rounded parotid glands than _bocourti_. The tadpoles of the two
species are nearly identical (see Stuart, 1943:12).
~Leptodactylus labialis~ (Cope)
_Cystignathus labialis_ Cope, Proc. Amer. Philos. Soc.,
17:90, 1877.--No type locality designated; type locality
restricted to Potrero Viejo, Veracruz, Mexico, by Smith and
Taylor (1950a:350).
_Leptodactylus labialis_, Brocchi, Mission Scientifique au
Mexique et dans l'Amerique Centrale, pt. 3, sec. 2, livr.
1:20, 1881.
Apatzingan (26); Capirio (5); Cofradia (9); El Sabino (4);
Lombardia; Rio Tepalcatepec, 27 km. S of Apatzingan (2).
In the Tepalcatepec Valley this frog reaches the northernmost known
limit of its range in western Mexico. Although the species is abundant
in the valley, it apparently is absent from the coastal lowlands. In the
Tepalcatepec Valley _Leptodactylus melanonotus_ seems to be more
abundant than _labialis_. In the rainy season both species have been
heard calling from the same ponds and flooded fields.
There are only slight differences in size between the sexes;
measurements of 20 males and eight females are, respectively: snout-vent
length, 32.3-39.5 (35.1), 34.1-39.2 (37.2); tibia length, 14.3-17.0
(15.4), 14.9-16.8 (15.8); head width, 11.0-13.6 (12.0), 12.2-13.2
(12.6); head length, 12.8-15.1 (13.3), 12.8-14.6 (13.7).
~Leptodactylus melanonotus~ (Hallowell)
_Cystignathus melanonotus_ Hallowell, Proc. Acad. Nat. Sci.
Philadelphia, 12:485, 1861.--Nicaragua. Type locality
restricted to Recero, Nicaragua, by Smith and Taylor
(1950a:320).
_Leptodactylus melanonotus_, Brocchi, Mission Scientifique
au Mexique et dans l'Amerique Centrale, pt. 3, sec. 2, livr.
1:20, 1881.
Apatzingan (103); Capirio; Charapendo (7); Coahuayana;
Cofradia (10); El Sabino (21); La Playa (3); Lombardia (5);
Maruata; Nueva Italia (7); Ostula (9); Playa Azul (11); Rio
Marquez, 10 km. S of Lombardia; Rio Marquez, 13 km. SE of
Nueva Italia (6); Rio Tepalcatepec, 27 km. S of Apatzingan.
This species is widespread in the lowlands of the state; it has been
collected up to elevations of 1050 meters in the Tepalcatepec Valley. In
the dry season individuals were discovered beneath rocks along streams
and in damp arroyos; in the rainy season they were found wherever there
was water. Males were heard calling from flooded fields, ditches, rocky
streams, and small puddles. The call is a series of individual notes:
"woink, woink, woink."
Adult males are noticeably smaller than females; measurements for 20
males and ten females from Apatzingan are, respectively: snout-vent
length, 29.6-34.6 (32.3), 36.3-44.1 (40.8); tibia length, 12.6-15.1
(14.0), 16.5-19.0 (17.8); head width, 10.8-11.9 (11.3), 12.6-14.8
(13.7); head length, 11.2-13.2 (11.9), 13.1-14.8 (14.0). Brownish yellow
ventral glands are present in some juveniles and in some adults
collected in the dry season as well as in the rainy season.
~Leptodactylus occidentalis~ Taylor
_Leptodactylus occidentalis_ Taylor, Trans. Kansas Acad.
Sci., 39:349, 1937.--Tepic, Nayarit, Mexico.
Five km. W of Tangamandapio.
On the night of June 11, 1958, this species was calling from a
hyacinth-choked ditch. Although numerous individuals were heard, only
one specimen was obtained. The frogs were calling from the tangled mat
of hyacinths along with _Hyla eximia_, _Hypopachus oxyrrhinus ovis_, and
_Rana pipiens_.
Taylor (1936a:352) characterized this species as follows: "The narrow
head, small maximum size (38 mm. for females, 33 mm. for males), the
character of the postaxillary and postfemoral glands, the narrower
groups of vomerine teeth, clearly distinguish this western Mexican form
from the more robust, larger _melanonotus_ to the south. The call is
likewise fainter and different in quality." Concerning the glands,
Taylor (_loc. cit._) remarked: "There is a possibility that the horny
excrescence covering the glands may appear only during the breeding
season. This character is quite as strongly marked in females as in
males." Bogert and Oliver (1945:324) concluded that the population of
_Leptodactylus_ in northwestern Mexico could not be distinguished from
_melanonotus_ in other parts of the country and thus synonymized
_Leptodactylus occidentalis_ with _melanonotus_. Bogert and Oliver (op.
cit.: 324) stated that the extent as well as the presence or absence of
ventral glands was highly variable in all samples examined by them.
Upon seeing numerous living individuals of _Leptodactylus melanonotus_
from many parts of its range in Mexico and individuals of the population
of _Leptodactylus_ in northwestern Mexico (Nayarit and Sinaloa), I was
immediately impressed not so much by the differences in the development
of the ventral glands, but by the color of the glands. The differences
in color are apparent in freshly preserved specimens. With the exception
of _Leptodactylus_ from northwestern Mexico, specimens of _melanonotus_
from throughout Mexico and northern Central America have yellow or
yellowish brown glands. Specimens from northwestern Mexico have black or
brownish black glands that are conspicuously darker than those found in
_melanonotus_. Examination of 653 preserved specimens of _Leptodactylus
melanonotus_ from Mexico and Guatemala has failed to reveal specimens
with black ventral glands, like those found in specimens from
northwestern Mexico, to which the name _Leptodactylus occidentalis_ has
been applied. Furthermore, in _melanonotus_ the glands are less distinct
and more extensive than in _occidentalis_; in the latter species glands
are absent from the throat and midventral area, where they often are
present in _melanonotus_ (Fig. 7).
In some individuals of both species collected in the dry season and in
some collected in the rainy (breeding) season the glands are absent; the
development of these glands, therefore, does not seem to be correlated
with breeding. Likewise, the glands are present or absent in either sex,
and often as not they are present in juveniles. Presence of the glands,
therefore, cannot be correlated either with sexual or ontogenetic
development. Since the glands are found in individuals from all parts of
the range, it is unlikely that there is a correlation between the
development of the glands and the environment.
[Illustration: FIG. 7. Diagrammatic view of ventral surfaces of
_Leptodactylus melanonotus_ (A) and _Leptodactylus occidentalis_ (B),
showing usual position and size of glandular areas. Approx. natural
size.]
Aside from the differences in the ventral glands, the call is different
in the two populations. The call of _Leptodactylus occidentalis_ is a
rather harsh "wack, wack, wack" as contrasted with the more nasal
"woink, woink, woink" of _melanonotus_. Sound spectrographs are needed
to analyze the differences in calls. None of the specimens of
_occidentalis_ examined approaches in size the largest individuals of
_melanonotus_; possibly the size of the frogs is another valid character
for separating the species. On the basis of the above data it is evident
that the frogs in northwestern Mexico show certain characters that
distinguish them from _Leptodactylus melanonotus_, as it is known
throughout the rest of Mexico. It is not known for certain that
_melanonotus_ and _occidentalis_ are sympatric. Several series of old,
poorly preserved specimens from Nayarit and Sinaloa cannot be placed in
either species, for none has visible ventral glands. _Leptodactylus
melanonotus_ is known from Acaponeta, Nayarit (AMNH 43913-25), and the
following localities in Jalisco: Barro de Navidad (UMMZ 118098), La
Concepcion (UMMZ 113081), La Resolana (UMMZ 102104), and Tenachitlan
(UMMZ 113045-6). Records for _Leptodactylus occidentalis_ are: Alamos,
Sonora (AMNH 51356-65); Culiacan (AMNH 49511-9), Chele (UMMZ 110914),
and Rosario (UMMZ 113062) in Sinaloa; Ixtlan del Rio (UMMZ 102108), San
Blas (UMMZ 112814, 112994, 110892, 115543), and Tepic (UMMZ 115544) in
Nayarit; Ameca (UMMZ 102106-7) and La Cofradia on the south shore of
Lago de Chapala (UMMZ 102105) in Jalisco; and Tangamandapio, Michoacan
(UMMZ 119145). From these scattered records it appears that
_Leptodactylus occidentalis_ in the southern part of its range stays in
the uplands, whereas _melanonotus_ is confined to the lowlands.
~Microbatrachylus hobartsmithi~ (Taylor)
_Eleutherodactylus hobartsmithi_ Taylor, Trans. Kansas Acad.
Sci., 39:355, 1937.--Uruapan, Michoacan, Mexico.
_Microbatrachylus hobartsmithi_ Taylor, Univ. Kansas Sci.
Bull., 26:501, November 27, 1940.
Cascada Tzararacua (6); 21 km. W of Ciudad Hidalgo; 29 km. E
of Morelia; Puerto Hondo; San Jose de la Cumbre (13);
Uruapan (2); Zitacuaro.