AMPHIBIA
Caudata
Ambystoma amblycephalum Taylor
Ambystoma amblycephala Taylor, Univ. Kansas Sci. Bull., 26: 420, November 27, 1940.—Fifteen kilometers west of Morelia, Michoacán, México.
Fifteen km. W of Morelia (19); 11 km. SSE of Opopeo (12); 8 km. S of Pátzcuaro; 24 km. S of Pátzcuaro (2); Quiroga (20); Tacícuaro (167).
Taylor and Smith (1945:530) presented data on 137 specimens collected at Tacícuaro on October 1, 1939; these are all larvae and metamorphosing individuals. Aside from these, the largest larva examined (UMMZ 104962 from 15 km. W of Morelia) has a snout-vent length of 70.0 mm. and a tail length of 53.5 mm. The larvae are pale pinkish tan above and somewhat paler below; there is a lateral row of cream colored spots. The tail-fin, which is deepest at mid-length, extends to the back of the head and is flecked with brown. In small larvae the outer edge of the tail-fin is dark brown. The eyes are large. Two small metamorphosed specimens (UMMZ 98967) from 24 kilometers south of Pátzcuaro are tentatively referred to this species. These specimens have body lengths of 49.0 and 45.0 mm. and tail lengths of 36.0 and 31.5 mm., respectively. They have 17-17 and 16-15 vomerine teeth arranged in a broad arch behind the choanae, 10 costal grooves, and 7 intercostal spaces between adpressed toes. The dorsal color is uniform brown; that of the venter is a dusty cream.
Larvae were collected from shallow ponds near Quiroga and 15 kilometers west of Morelia; metamorphosed individuals were taken from beneath logs in pine and fir forests at elevations from 2300 to 2800 meters.
Ambystoma dumerili dumerili (Dugès)
Siredon Dumerili Dugès, La Naturaleza, 1:241, 1870—Lago de Pátzcuaro, Michoacán, México.
Bathysiredon dumerilii, Dunn, Notulae Naturae, 36:1, November 9, 1939.
Bathysiredon dumerilii dumerilii, Maldonado-Koerdell, Mem. y Rev. Acad. Nac. Cien., 56:199, 1948.
Ambystoma (Bathysiredon) dumerili, Tihen, Bull. Florida State Mus., 3:3, June 20, 1958.
Lago de Pátzcuaro (22);? Morelia.
For many years this unusual salamander was known from only a few specimens mostly collected in the last century; Smith and Taylor (1948:7) stated: "It is presumed that this species is extinct owing to the introduction of exotic game and food fishes." In 1951 and in 1955 I had been told that axolotls were sold in the market at Pátzcuaro; nevertheless, none was found on my visits there. In 1956 Charles M. Bogert obtained several large specimens at the market in Pátzcuaro. These establish the continued existence of the salamander in Lago de Pátzcuaro. On January 27, 1955, R. W. Dickerman procured a specimen (KU 41573) in the market at Morelia. Since fish are brought to Morelia from Lago de Pátzcuaro, the specimen probably was from that lake. Nevertheless, the species may occur in other permanent bodies of water in Michoacán. Maldonado-Koerdell (1948) described Bathysiredon dumerili queretarensis from San Juan del Río, Queretaro. This locality is about 200 airline kilometers northeast of Lago de Pátzcuaro and is in the Río Moctezuma drainage.
Ambystoma ordinarium Taylor
Ambystoma ordinaria Taylor, Univ. Kansas Sci. Bull., 26:422, November 27, 1940.—Four miles west of El Mirador, near Puerto Hondo, Michoacán, México.
Axolotl (56); Cerro San Andrés; 22 km. W of Mil Cumbres; 46 km. E of Morelia (34); 8 km. SE of Opopeo (5); Puerto de Garnica (8); Puerto Hondo (41); San Gregorio (16); San José de la Cumbre (20).
Of 16 specimens (KU 51520-35) collected on June 18, 1955, near San Gregorio, 15 are adult females with swollen cloacae and minute ovarian eggs. Possibly these specimens had just recently deposited their mature eggs. In preservative the specimens are black above and dull creamy gray below. Measurements for the 15 females are: snout-vent length, 80.0-102.0 (92.5); tail length, 69.0-93.0 (84.2); head width, 15.8-20.5 (17.7); head length, 22.8-26.6 (24.4). A larval specimen with small gills has a snout-vent length of 72 mm. and a tail length of 62 mm. Three specimens have 12 costal grooves; the other have 11.
Of 20 specimens from San José de la Cumbre (UMMZ 112857 and 115143), 14 are neotenic adults; the others are larvae. In life the salamanders were blackish to olive-brown above with scattered cream-colored dots on the dorsum and flanks but in preservative are dull grayish black with indistinct pale spots and dark reticulations. The belly is pale gray with indistinct dark spots. Eleven females and three males have the following measurements, respectively: snout-vent length, 76.0-90.0 (80.7), 64.0-84.0 (74.3); tail length, 70.0-81.0 (75.0), 58.0-71.0 (66.7); head width, 19.5-23.5 (20.7), 17.5-20.5 (19.3); head length, 22.0-25.0 (23.0), 20.0-22.5 (21.5). The smallest larva has a snout-vent length of 43.0 mm. and a tail length of 38.0 mm. Two individuals have 12 costal grooves; the others have 11. All of the females contained eggs, the largest of which were 1.5 mm. in diameter. The stomachs of most of the specimens were distended with oligochaets, aquatic insect larvae, and small aquatic beetles.
A series of 34 larvae (JRD 5904-37) from 46 kilometers east of Morelia are tentatively referred to this species. These specimens are olive-brown above with cream-colored spots on the flanks; the dorsal tail-fin does not extend onto the body.
This species has been found only at elevations in excess of 2400 meters in pine and fir forests. At Rancho Axolotl James A. Peters collected larvae and neotenic individuals in a rocky stream and adults from beneath rocks and logs in the forest near the stream. Neotenic individuals and larvae were found in a clear stream in pine-fir forest at an elevation of 2700 meters near San José de la Cumbre; specimens were collected there in July, 1955, and again in July, 1956. The site was visited in April, 1956, at which time the stream consisted of only a few puddles; no salamanders were found.
Ambystoma tigrinum velasci Dugès
Ambystoma velasci Dugès, La Naturaleza, ser. 2, 1:142, 1888.—Laguna Santa Isabel, near Guadalupe Hidalgo, Distrito Federal, México.
Ambystoma tigrinum velasci, Dunn, Copeia, no. 3:157, November 14, 1940.
Pátzcuaro (5); Tacícuaro (9).
Definite specific assignment of these specimens, all larvae, cannot be made at this time. They have shovel-shaped heads and laterally compressed bodies with the dorsal tail-fin extending anteriorly to the back of the head. The eyes are small. The body is pale tan with dark mottling on the tail and flanks. The average snout-vent length for nine specimens from Tacícuaro is 61.0 mm.
The larvae from Tacícuaro (UMMZ 89255) were collected by Dyfrig Forbes in October, 1939; those from Pátzcuaro, presumably Lago de Pátzcuaro (BMNH 1914.1.28-247-8 and CNHM 948), were collected by Hans Gadow and Seth Meek in 1908.
Pseudoeurycea belli (Gray)
Spelerpes belli Gray, Catalogue Batrachia Gradientia British Museum, p. 46, 1850.—México. Type locality restricted to 2 miles east of Río Frío, Puebla, México, by Smith and Taylor (1950a:341).
Pseudoeurycea bellii, Taylor, Univ. Kansas Sci. Bull., 30:209, June 12, 1944.
Axolotl (2); Carapan; Cerro Tancítaro (84); Macho de Agua; 22 km. W of Mil Cumbres; Opopeo; Pátzcuaro (8); Puerto Hondo (2): San José de la Cumbre; San Juan de Parangaricutiro (42); Uruapan (5); Zacapu (4).
This salamander seems to reach its greatest abundance in Michoacán in the Sierra de los Tarascos between Pátzcuaro and Tancítaro, where it is found at elevations from 1500 to 2900 meters. It is found less commonly in the eastern part of the Cordillera Volcánica in Michoacán, where it sometimes occurs in association with Pseudoeurycea robertsi.
On June 22 and 23, 1955, four clutches of eggs of this species were found beneath adobe bricks and rocks on the volcanic ash that has buried the village of San Juan de Parangaricutiro. The eggs were unstalked and separate, but adherent in clumps of three or four (Pl. 2, Fig. 1). The outer membranes were covered with fine particles of ash. The ash beneath the stones where the eggs were found was only slightly moist; one clump of eggs was partially desiccated. Three complete clutches have 20, 23, and 34 eggs; one clutch of 15 eggs was being eaten by beetles (Tenebrionidae: Eleodes sp.). The eggs vary in size from 4.6 to 6.5 mm. and average 5.3 mm. in diameter. They are unpigmented. Surrounding the embryo is a vitelline membrane, an inner, and an outer envelope (Fig. 1). In an average-sized egg having an embryo 4 mm. in length, the diameter of the outer membrane is 5.3 mm., the inner membrane 5.0 mm., and the vitelline membrane 4.6 mm. All of the eggs contained embryos in which the limb buds were developed; in about half of these the eyes were distinctly visible.
Fig. 1. Diagram of an egg of Pseudoeurycea belli from San Juan de Parangaricutiro, Michoacán. × 10.
The first heavy rain of the season occurred on the night of June 22, 1955. Thus, at least sometimes, Pseudoeurycea belli lays its eggs before the onset of the rainy season. A female having a snout-vent length of 110 mm., collected on June 22, 1955, contained 36 ovarian eggs having diameters from 3.0 to 3.5 mm. The fact that small juveniles were collected on the same date indicates that this salamander lays eggs over a period of several weeks in late spring and early summer.
The smallest juvenile examined has a snout-vent length of 17.0 mm. and a tail length of 7.5 mm. Twelve juveniles from the vicinity of San Juan de Parangaricutiro have an average snout-vent length of 19.4 mm. and an average tail length of 9.7 mm. In juveniles the adpressed limbs either touch or overlap by one intercostal space; in adults there are two or three intercostal spaces between adpressed toes. Therefore the greatest number of intercostal spaces between adpressed limbs is found in the largest specimens. A similar relationship between adpressed limbs (= length of limbs) and snout-vent length was shown for Plethodon richmondi by Duellman (1954a). The number of vomerine teeth is variable; the number of teeth seems to be closely correlated with the size of the salamander (Fig. 2). A similar correlation between the number of maxillary teeth and body length was reported for Chiropterotriton multidentatus by Rabb (1958). In 12 juvenile Pseudoeurycea belli there are 6-13 (8.8) vomerine teeth, and in 11 adults having snout-vent lengths greater than 90 mm. there are 39-49 (44.0) vomerine teeth. The coloration of the juveniles resembles that of the adults (Pl. 1).
Fig. 2. Correlation between the number of vomerine teeth and snout-vent length in 79 Pseudoeurycea belli from Michoacán.
The differences between this species and Pseudoeurycea gigantea are minor. Taylor (1939a) distinguished gigantea from belli by the larger size, fewer intercostal spaces between adpressed limbs, more vomerine teeth, and absence of occipital spots in gigantea. Taylor and Smith (1945) stated that in life the spots in gigantea are orange instead of red as in belli. Five specimens of Pseudoeurycea belli from Michoacán, including one juvenile, lack occipital spots. In the 34 living individuals that I have seen from Michoacán the spots varied from deep red to orange. Therefore, of the characters listed by Taylor (op. cit.) to diagnose Pseudoeurycea gigantea, only the over-all larger size and smaller number of intercostal spaces between adpressed limbs (= relatively longer limbs) are useful in separating Pseudoeurycea belli and gigantea.
Pseudoeurycea robertsi (Taylor)
Oedipus robertsi Taylor, Univ. Kansas Sci. Bull., 25:287, July 10, 1939.—Nevado de Toluca, México.
Pseudoeurycea robertsi Taylor, Univ. Kansas Sci. Bull., 30:209, June 12, 1944.
Atzimba (3); Macho de Agua (9); Puerto Lengua de Vaca (14).
Previously this species has been recorded only from the type locality. In July, 1956, individuals referable to this species were found at two sites in pine-fir forest immediately to the east of Macho de Agua and in pine-oak-fir forest at Atzimba. On August 20, 1958, a series was collected in pine-fir forest at Puerto Lengua de Vaca. These localities are between 2900 and 3000 meters in the Cordillera Volcánica in eastern Michoacán.
In life the coloration of these salamanders was highly variable. The belly and undersurfaces of the tail and hind limbs were pale gray, with or without silvery white flecks; the chin was a cream-color and flecked with silvery white in some specimens. The middorsal area was brown, orange-brown, or dull grayish yellow. The flanks and lateral surfaces of the tail were black with yellowish flecks or streaks on the flanks and yellowish or orange-brown flecks on the tail. The iris was golden brown. Measurements of eight males and two females are, respectively: snout-vent length, 42.5-56.0 (49.5), 54.0-60.0 (57.0); tail length, 42.0-56.0 (48.1), 52.0-55.0 (53.5). The smallest juvenile has a snout-vent length of 28.0 mm. and a tail length of 23.0 mm. Of the 26 available specimens, six have 12 costal grooves, and the others have 11.
In comparison with 36 topotypes, the specimens from Michoacán have a less striking dorsal color pattern; none has a well-defined dorsal reddish brown area or bold reddish mottling on the tail. Furthermore, the specimens from Michoacán have paler venters than do topotypic specimens.
Salientia
Rhinophrynus dorsalis Duméril and Bibron
Rhinophrynus dorsalis Duméril and Bibron, Erpétologie générale, vol. 8:758, 1841.—Veracruz, Veracruz, México.
Mouth of the Río Balsas (10).
These specimens (BMNH 1914.1.28.181-90) were collected by Gadow in 1908 and reported by him (1930:72): "Whilst this very sluggish termite-eating toad is common enough in the sweltering hot country of the state of Vera Cruz, up to an elevation of 1500 feet, it was unknown on the west side of the Isthmus until I found it in great numbers near the mouth of the Balsas River, in and near fresh-water pools, where it attracted attention by its loud peculiar voice during the pairing season in the month of July." Subsequently, Peters (1954:3) verified the identification of these specimens. Although torrential rains fell during the week in July, 1955, that I spent at Playa Azul near the mouth of the Río Balsas, the distinctive voice of Rhinophrynus was not heard. Elsewhere on the Pacific coast of México adult Rhinophrynus have been reported only from Tehuantepec and a few localities on the coastal lowlands of Chiapas. Taylor (1942b:37) found on the coast of Guerrero a tadpole that was referred to the genus Rhinophrynus by Orton (1943). In the summer of 1960 adults of Rhinophrynus were collected near Acapulco, Guerrero (Fouquette, in litt.). These recent collections verify the existence of the species along the Pacific lowlands of México at least as far north as Michoacán.
Scaphiopus hammondi multiplicatus Cope
Scaphiopus multiplicatus Cope, Proc. Acad. Nat. Sci. Philadelphia, 15:52, June 8, 1863.—Valley of México.
Scaphiopus hammondi multiplicatus, Kellogg, Bull. U. S. Natl. Mus., 160:22, March 31, 1932.
Angahuan (5); Cuitzeo (4); Cuseño Station (2); Jiquilpan (9); Morelia (7); Pátzcuaro (3); Quiroga; Tarécuaro; Uruapan (24); Zacapu.
This small toad has been found at elevations between 1500 and 2500 meters on the Mexican Plateau and associated mountain ranges; it occurs in mesquite-grassland and in pine forests. Calling males and females laden with eggs have been collected in the rainy season in the months of July and August. The call is a medium-pitched snore. In living individuals the dorsal ground color varies from pale brown to gray with dark brown or olive-brown markings. In many individuals the tips of the small dorsal pustules are red.
Bufo coccifer Cope
Bufo coccifer Cope, Proc. Acad. Nat. Sci. Philadelphia, 18:130, 1866—Arriba, Costa Rica.
Apatzingán (27); Lombardia; Nueva Italia (5).
In life the dorsal color pattern consists of a yellowish tan ground color with dark brown spots; the middorsal stripe is deep yellow or cream color. The venter is a dusty cream color, and the iris is pale gold. Males have dark brown horny nuptial tuberosities on the thumb. The following measurements are of 21 males and four females, respectively: snout-vent length, 43.5-51.7 (48.1), 55.6-62.6 (59.1); tibia length, 16.6-18.8 (17.6), 18.8-20.3 (19.3); head width, 16.7-19.7 (18.4), 20.6-22.2 (21.4); head length, 13.8-16.6 (14.8), 16.5-18.2 (17.3).
The specimens from the Tepalcatepec Valley differ slightly from specimens from southeastern México and Central America. Those from Michoacán have low and narrow cranial crests; in about one-half of the specimens the occipital crest exists only as a row of tubercles, and in some the postorbital and suborbital crests are barely discernible. Specimens from the southern part of the range, Costa Rica and Nicaragua, have much higher and thicker cranial crests; in these the occipital crest is well defined and extends posteriorly to a point back of the anterior edge of the parotid gland; the postorbital and suborbital crests are well marked. Of 48 specimens from Esquipulas, Guatemala, all have high crests, but these are not so well developed as in ten specimens from Matagalpa, Nicaragua, and three from various localities in Costa Rica. Six specimens from Tehuantepec, Oaxaca, have cranial crests that are lower than those in specimens from Guatemala. In three of the specimens from Tehuantepec the occipital crests are reduced to a series of tubercles. Of six specimens from Agua del Obispo, Guerrero, four have poorly developed occipital crests. These observations suggest the presence of a cline in the development of the cranial crests; specimens have higher crests in the southern part of the range than in the northern part.
In México Bufo coccifer has been collected only in semi-xeric habitats, but to the south, from Guatemala to Costa Rica, it has been found in more upland and humid habitats. Southern specimens are darker than those from the north, a possible correlation with the differences in habitat.
These toads probably range throughout the Tepalcatepec Valley, but they are unknown from the coast of Michoacán. Breeding choruses were found after heavy rains on June 24, 1955, and on August 2, 1956. The first was in a muddy ditch; the second was in a flooded grassy field. The call is a high-pitched, but not loud, "whirrr." Males were calling from the edge of the water or from clumps of grass in the water. Clasping pairs were in the water; amplexus is axillary.
Bufo compactilis compactilis Wiegmann
Bufo compactilis Wiegmann, Isis von Oken, 26:661, 1833.—México. Type locality restricted to Xochimilco, Distrito Federal, México, by Smith and Taylor (1950a:330).
Bufo compactilis compactilis, Smith, Herpetologica, 4:7, September 17, 1947.
Cuitzeo (2); Emiliano Zapata (20); Jiquilpan (5); La Palma (5); Morelia; Tupátaro.
The southwestern terminus of the range of this species is on the Mexican Plateau in Michoacán. 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, México. 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, México.
Aguililla; Apatzingán (3); Barranca de Bejuco; Capirio; Charapendo; Chichihuas; Coahuayana (2); Coalcomán (7); Cofradía (2); 25 km. S of Cuatro Caminos; El Sabino (10); Huahua, La Playa (13); Ojos de Agua de San Telmo; Ostula; Playa Azul (2); Pómaro (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 México 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 México 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, México.
Barranca de Bejuco; Coahuayana (11); El Diezmo (2); La Placita (9); La Orilla (12); Motín del Oro; Ostula (9); Playa Azul (5); Pómaro (15); Salitre de Estopilas; San Pedro Naranjestila.
In Michoacán this species is confined to elevations of less than 1000 meters on the coast and foothills of the Sierra de Coalcomán. 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%) |
| Izúcar, 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 Michoacán | 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%) |
| Tonolá, 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.
Fig. 3. Adult male of Bufo perplexus from Apatzingán, Michoacán. × 1.5.
Fig. 4. Adult male of Bufo marmoreus from Pómaro, Michoacán. × 1.5.