El Peten, Guatemala, and they average 52.5 mm. in snout-vent length
whereas males from Los Tuxtlas, Veracruz, average 50.6 mm. and those
from northern Oaxaca 50.3 mm. The smallest breeding male has a
snout-vent length of 44.6 mm. The average ratio of tibia length to
snout-vent length is 54.8 per cent in males from Piedras Negras,
and 56.4 and 56.3 per cent in males from Los Tuxtlas and Oaxaca,
respectively. The only other character showing noticeable geographic
variation is the size of the tympanum. The average ratio of the diameter
of the tympanum to the diameter of the eye is 76.3 per cent in males
from Piedras Negras, 71.8 from Oaxaca, and 66.9 from Los Tuxtlas.
The dorsal ground color of _Smilisca cyanosticta_ is pale green to tan
and the venter is creamy white. The dorsum is variously marked with dark
olive-green or dark brown spots or blotches (Pl. 6B). An interorbital
dark bar usually is present. The most extensive dark area is a V-shaped
mark in the occipital region with the anterior branches not reaching the
eyelids; this mark is continuous, by means of a narrow mid-dorsal mark,
with an inverted V-shaped mark in the sacral region. In many specimens
this dorsal marking is interrupted, resulting in irregular spots. In
some specimens the dorsum is nearly uniform pale green or tan with a few
small, dark spots. The hind limbs are marked by dark transverse bands,
usually three or four each on the thigh and shank, and two or three on
the tarsus. The webbing on the feet is brown. The loreal region is pale
green, bordered above by a narrow, dark brown canthal stripe extending
from the nostril to the orbit, which is bordered above by a narrow,
bronze-colored stripe extending from the nostril along the edge of the
eyelid to a point above the tympanum. The upper lip is white. A broad
dark brown mark extends posteriorly from the orbit and encompasses the
tympanum to a point above the insertion of the forelimb. The flanks are
dark brown with many pale blue, rounded spots, giving the impression of
a pale blue ground color with dark brown mottling enclosing spots. The
anterior and posterior surfaces of the thighs are dark brown with many
small pale blue spots. The inner surfaces of the shank and tarsus are
colored like the posterior surfaces of the thighs. Pale blue spots are
usually present on the proximal segments of the second and third toes. A
distinct white stripe is present on the outer edge of the tarsus and
fifth toe; on the tarsus the white stripe is bordered below by dark
brown. A white stripe also is present on the outer edge of the forearm
and fourth finger. The anal region is dark brown, bordered above by a
narrow transverse white stripe. The throat in breeding males is dark,
grayish brown with white flecks.
No geographic variation in the dorsal coloration is evident. Specimens
from the eastern part of the range (Piedras Negras and Chinaja,
Guatemala) have bold, dark reticulations on the flanks enclosing large
pale blue or pale green spots, which fade to tan in preservative.
Specimens from Oaxaca and Veracruz characteristically have finer dark
reticulations on the flanks enclosing smaller blue spots; in many of
these specimens the ventrolateral spots are smallest and are white.
All living adults are easily recognized by the presence of pale, usually
blue, spots on the flanks and thighs. Individuals under cover by day
have a tan dorsum with dark brown markings. A hiding individual at
Chinaja, Alta Verapaz, Guatemala (KU 55936), had a pale tan dorsum when
found; later the dorsal color changed to chocolate brown. A pale green
patch was present below the eye; the spots on the posterior surfaces of
the thighs were pale blue, and those on the flanks were yellowish green.
A calling male obtained 10 kilometers north-northwest of Chinaja (KU
55934) was reddish brown when found at night; later the dorsal color
changed to pale tan. A green patch below the eye was persistent. Usually
these frogs are green at night. The coloration of an adult male (KU
87201) from 11 kilometers north of Vista Hermosa, Oaxaca, Mexico, was
typical: "When calling dorsum pale green; later changed to dull
olive-green. Flanks dark brown with pale blue spots in axilla and groin
and bluish white flecks on mid-flank. Anterior and posterior surfaces of
thighs, inner surfaces of shanks, and median dorsal surfaces of tarsi
dark brown with blue spots. Canthal and postorbital stripes dark
chocolate brown; labial stripe creamy white. Forearm, tarsal, and anal
stripes pale cream-color. Throat dark brown with yellow flecks; belly
and ventral surfaces of limbs creamy buff; webs pinkish tan; iris deep
bronze, brown below pupil." (Duellman, field notes, June 24, 1964.)
Some individuals have both green and brown coloration in life. An
individual obtained at night on the south slope of Volcan San Martin,
Veracruz, Mexico, had a pale tan dorsum changing peripherally to pale
green. The dorsal markings were dark brown and dark olive-green.
In contrast to the color-changes noted above, the labial region below
the eye is always pale green, and pale spots are always present on the
flanks and thighs in adults. The iris is invariably golden or bronze
above and darker, usually brown, below. Minute black flecks are present
on the iris, and in some individuals these flecks are so numerous that
the eye appears gray.
Recently metamorphosed young have pale tan flanks, and the posterior
surfaces of the thighs are orange-yellow; pale spots are absent. A
juvenile (KU 55935) from Chinaja, Alta Verapaz, Guatemala, having a
snout-vent length of 35.0 mm. was pale yellowish tan above with
olive-green markings; the flanks were dark brown with pale blue spots,
and the anterior and posterior surfaces of the thighs were uniform
bright tomato red. A juvenile (UMMZ 121298), 18.6 mm. in snout-vent
length, from the southeast slope of Volcan San Martin, Veracruz, Mexico,
had pale tan flanks lacking blue spots, but had red thighs. Apparently
the ontogenetic changes in coloration proceed as follows: (1) flanks
pale tan and thighs orange-yellow, both lacking spots, (2) flanks pale
tan and thighs red, both lacking spots, (3) flanks dark brown with blue
spots and thighs red, lacking spots, and (4) flanks and thighs dark
brown, both having pale blue spots.
_Natural History._--_Smilisca cyanosticta_ inhabits humid tropical
forest and cloud forest from the lowlands to elevations of about 1200
meters in Los Tuxtlas and to about 900 meters in northern Oaxaca. In
these moist environments the frogs apparently are active throughout the
year. Active individuals have been obtained in January, July, and August
in Los Tuxtlas, Veracruz, in June and July in northern Oaxaca, in
February and March at Chinaja, Guatemala, and Taylor and Smith reported
(1945:589) activity in May at Piedras Negras, Guatemala. Calling males
were observed as follows; in a rain barrel 11 kilometers north of Vista
Hermosa, Oaxaca, Mexico, on June 23-28, 1964; in a quiet pool in a
stream 8 kilometers south of Yetla, Oaxaca, Mexico, in July, 1963 (Dale
L. Hoyt, personal communication); in and near springs flowing into a
stream at Dos Amates, Veracruz, Mexico, on August 4, 1959 (Douglas
Robinson, personal communication); and in a water-filled depression in a
log 10 kilometers west-northwest of Chinaja, Guatemala, on March 13,
1960. Taylor and Smith (1945:589) reported that individuals were found
at night on the ground at the edge of temporary pools at Piedras Negras,
Guatemala, on May 28-29, 1939. A clasping pair was found on a rock at
the edge of a small stream on the south slope of Volcan San Martin,
Veracruz, Mexico, on July 11, 1959 (Douglas Robinson, personal
communication).
Only one individual has been observed in a tree at night. In the
daytime, individuals were found in elephant ear plants (_Xanthosoma_) at
Chinaja, Guatemala.
The breeding call consists of one or two moderately short notes that are
lower pitched than those of _S. baudini_, but higher pitched than those
of _S. phaeota_. Each note has a duration of 0.25 to 0.45 seconds and is
repeated at intervals of one-half minute to several minutes. Each note
is a vibrant "waunk," having 110 to 180 pulses per second and dominant
frequency of 1600 to 2100 cycles per second (Pl. 10B).
Apparently the eggs are deposited as loose clumps in the water. In eggs
in the yolk plug stage of development, the diameter of the embryo is
about 2.3 mm.; that of the outer envelope is 4.0 mm. Hatchling tadpoles
have total lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm.
The external gills are moderately long, slender, and filamentous; the
yolk sac is still moderately large. The body and anterior part of the
caudal musculature are dark brown; posteriorly the caudal musculature is
pale brown. The caudal fins are creamy tan. The oral discs are large and
ovoid. The growth of the tadpole is summarized in Table 10.
A typical tadpole in stage 30 of development (KU 87652 from 11 km. N
Vista Hermosa, Oaxaca, Mexico) can be described as follows:
Body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body
slightly wider than deep; snout rounded laterally, broadly ovoid
dorsally; eyes widely separated, directed dorsolaterally; nostril about
midway between eye and tip of snout; mouth anteroventral; spiracle
sinistral, slightly posterior to midpoint of body and slightly below
midline; anal tube dextral; caudal musculature slender, barely curved
upward distally; dorsal fin not extending onto body, depth of dorsal fin
slightly more than that of ventral fin on mid-length of tail; dorsal
part of body dark brown; ventral surfaces transparent, lacking pigment;
posterior edge of body pale cream-color; caudal musculature creamy white
with interconnected brown spots; caudal fins transparent with small
brown blotches on dorsal fin and posterior half of ventral fin; iris
coppery bronze in life (Fig. 12). Mouth small, median part of upper lip
bare; rest of mouth bordered by single row of bluntly rounded papillae;
lateral fold present; tooth rows 2/3; all tooth-rows approximately equal
in length; second upper row broadly interrupted medially; other rows
complete; upper beak moderately deep, forming broad arch with slender
lateral processes; lower beak slender, broadly V-shaped; both beaks
finely serrate (Fig. 15C).
All tadpoles having fully developed mouth parts have 2/3 tooth rows.
Little variation is noticeable in coloration. In many specimens the
posterior edge of the body is dark brown instead of pale cream-color.
Mottling is rather dense on the caudal fins in all specimens; in some
individuals pigment is concentrated along the anterior one-third of the
lateral groove. In life the body is dark brown with greenish gold flecks
ventrally; the caudal musculature is gray.
In each of two recently metamorphosed young the snout-vent length is
14.0 mm. Coloration of young in life (KU 87653 from 11 km. N Vista
Hermosa, Oaxaca, Mexico): "Dorsum pale tan with dark brown markings.
Thighs orange-yellow; labial stripe white; iris bronze" (Duellman, field
notes, July 10, 1964.)
_Remarks._--Smith (1953:150) named _cyanosticta_ as a subspecies of
_Hyla phaeota_. The differences in cranial characters and certain
external characters between _phaeota_ and _cyanosticta_ indicate that
they are distinct species. Furthermore, a gap of about 350 kilometers
exists between the known geographic ranges of the two kinds.
_Distribution._--_Smilisca cyanosticta_ inhabits wet forests on the
Atlantic slope of southern Mexico and northern Central America from
northern Oaxaca and southern Veracruz through northern Chiapas in Mexico
and into El Peten and northern Alta Verapaz in Guatemala (Fig. 2).
Apparently the range is discontinuous, for in southern Mexico the
species is found in cloud forest at elevations of 830 to 900 meters on
the northern slopes of the Sierra de Juarez. In the Sierra de Los
Tuxtlas in southern Veracruz the species is found in wet forest at
elevations of 300 to 1200 meters, but is absent in the intervening
lowlands characterized by drier forest. In the west forests of northern
Alta Verapaz and El Peten, Guatemala, the species is found at low
elevations.
_Specimens examined._--78, as follows: MEXICO: =Chiapas=: Monte Libano,
MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH
41588. =Oaxaca=: 11 km. N Vista Hermosa, KU 84918-20 (skeletons),
87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH
57199-201; 8 km. S Yetla, KU 87213, UMMZ 124838 (8). =Veracruz=: Coyame,
UMMZ 111459-60; between Coyame and Tebanco, UMMZ 121198; Dos Amates,
UMMZ 121297; between Laguna de Catemaco and Volcan San Martin, UMMZ
121200; Volcan San Martin, UIMNH 35403-4, 35408-12, UMMZ 118163; SE
slope Volcan San Martin, UMMZ 121199, 121295 (2), 121296, 121298.
GUATEMALA: =Alta Verapaz=: Chinaja, KU 55935-7, 55938 (skeleton). =El=
=Peten=: 10 km. NNW Chinaja (Alta Verapaz), KU 55934; Piedras Negras,
CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. S
Piedras Negras, CNHM 99008; Semicoch, USNM 35907.
[Illustration: FIG. 2. Map showing locality records for _Smilisca
cyanosticta_ (triangles) and _Smilisca phaeota_ (circles).]
=Smilisca phaeota= (Cope)
_Hyla phaeota_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 14
(9):358, 1862 [Holotype.--USNM 4347 from Turbo, Colombia; J. Cassin
collector]. Boulenger, Catalogue Batrachia Salientia in British
Museum, p. 402, Feb. 1, 1882. Werner, Sitzungsb. Akad. Wiss.
Muenchen, 27:215, 1897. Guenther, Biologia Centrali-Americana:
Reptilia and Batrachia, p. 269, Sept. 1901. Nieden, Das Tierreich,
Amphibia, Anura I, p. 261, June 1923. Dunn, Occas. Papers Boston
Soc. Nat. Hist., 5:413, Oct. 10, 1931. Gaige, Hartweg, and Stuart,
Occas. Papers Mus. Zool. Univ. Michigan, 357:4, Oct. 26, 1937.
Cooper, Copeia, 2:122, June 30, 1944. Breder, Bull. Amer. Mus. Nat.
Hist., 86(6):416, Aug. 26, 1946. Smith and Taylor, Bull. U. S. Natl.
Mus., 194:88, June 17, 1948; Univ. Kansas Sci. Bull, 33:364, March
20, 1950. Taylor, Univ. Kansas Sci. Bull., 35(1):837, July 1, 1952.
Brattstrom and Howell, Herpetologica, 10:117, Aug. 1, 1954. Goin,
Herpetologica, 14:120, July 23, 1958. Cochran, Bull. U. S. Natl.
Mus., 220:57, 1961.
_Hyla labialis_ Peters, Monats. Konigl. Akad. Wissen. Berlin, p.
463, 1863 [Holotype.--ZMB 4913 from "region of Bogota," Colombia];
Monats. Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873.
Boulenger, Catalogue Batrachia and Salientia in British Museum, p.
397, Feb. 1, 1882.
_Hyla baudini dolomedes_ Barbour, Occas. Papers Mus. Zool. Univ.
Michigan, 129:11, Jan. 25, 1923 [Holotype.--MCZ 8539 from Rio
Esnape, Sambu Valley, Darien, Panama; Barbour and Brooks
collectors]. Barbour and Loveridge, Bull. Mus. Comp. Zool. Harvard,
69:278, June, 1929.
_Hyla phaeota phaeota_, Smith, Herpetologica, 8:152, Jan. 30, 1953.
Minton and Smith, Herpetologica, 16:103, June 17, 1960.
_Smilisca phaeota_, Starrett, Copeia, 4:303, Dec. 30, 1960.
_Diagnosis._--Size large ([M] 65 mm., [F] 78 mm.); skull as long as
wide, lacking frontoparietal fontanelle; large supraorbital flanges
having straight edges and extending posterolaterally; large squamosal
not in contact with maxillary; tarsal fold moderately wide, full length
of tarsus; inner metatarsal tubercle moderately large, low, flat,
elliptical; hind limbs relatively long, tibia averaging more than 53 per
cent of snout-vent length; labial stripe silvery white; lips lacking
vertical bars; loreal region pale green; dark brown or black tympanic
mark dispersing into brown venated pattern on flanks; posterior surfaces
of thighs pale brown, with or without darker flecks or small
cream-colored flecks. (Foregoing combination of characters
distinguishing _S. phaeota_ from any other species in genus.)
Table 2.--Geographic Variation in Size and Proportions in Males of
Smilisca phaeota. (Means in Parentheses Below Observed Ranges;
Data Based on Ten Specimens From Each Locality.)
==================================================================
|Snout-vent |Head width/|Interorbital
Locality |length |snout-vent |distance/
| |length |head width
-----------------------------+-----------+-----------+------------
Bonanza, Nicaragua | 40.8-47.7 | 34.1-38.0 | 31.0-36.1
| (43.7) | (36.3) | (35.4)
-----------------------------+-----------+-----------+------------
Heredia Prov., Costa Rica | 46.3-59.0 | 32.5-36.0 | 30.5-39.6
| (51.7) | (35.0) | (34.7)
-----------------------------+-----------+-----------+------------
Puntarenas Prov., Costa Rica | 53.6-64.9 | 32.6-36.1 | 31.0-38.0
| (61.4) | (34.5) | (34.4)
-----------------------------+-----------+-----------+------------
Canal Zone, Panama | 52.4-65.5 | 33.5-37.6 | 31.3-37.2
| (56.5) | (35.6) | (34.7)
-----------------------------+-----------+-----------+------------
Rio Quesada, Colombia | 52.6-61.0 | 33.1-37.1 | 30.1-33.9
| (56.0) | (35.0) | (32.1)
------------------------------------------------------------------
_Description and Variation._--For the purposes of analyzing geographic
variation in size and proportions, measurements were taken on ten adult
males from each of five samples throughout the range of the species.
Aside from the data summarized in Table 2, the average ratio of tibia
length to snout-vent length is noticeably less in Colombian specimens
(53.4 per cent, as compared with 54.8 to 57.8 per cent in the other
samples) and the ratio of head length to snout-vent length is
noticeably less in Costa Rican specimens (33.5 per cent as compared with
34.9 to 35.1 per cent in the other samples). Also, specimens from
Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7
to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared
with means of 74.0 to 77.9 per cent in the other samples).
Two populations are distinctive as regards the size of adult males.
Specimens from the northern Caribbean lowlands of Nicaragua (Bonanza,
the northernmost locality for the species) are remarkably small. Males
having snout-vent lengths of between 40 and 43 mm. were breeding; the
largest male found had a snout-vent length of 47.7 mm. The other extreme
in size is attained in specimens from the Pacific lowlands of eastern
Costa Rica and western Panama, where most breeding males have snout-vent
lengths of more than 55 mm.; the largest male had a snout-vent length of
64.9 mm.
The rather striking differences in size among certain samples and the
minor differences in proportions among other samples show no geographic
trends. Instead, the variations apparently are random among the samples.
The data presented here possibly are the results of inadequate sampling,
but more likely reflect actual differences in the populations.
The dorsal ground color of _Smilisca phaeota_ is pale green to tan; the
venter is creamy white. The dorsum is variously marked with dark
olive-green or dark brown spots or blotches (Pl. 6C). A dark
interorbital bar is usually present. Usually a large dark dorsal mark
extends from the occiput to the sacral region, but in many individuals
this blotch is replaced by two or three dark marks. The dorsal markings
are irregular in shape and do not tend to form transverse bands or
longitudinal bars. The hind limbs are marked by dark transverse bands,
usually four or five on the thigh, five or six on the shank, and four on
the tarsus. Two or three narrow bands are usually present on the
proximal part of the fourth toe. The webbing on the feet is brown. The
loreal region is pale green, bordered above by a narrow dark brown
canthal stripe extending from the nostril to the orbit. The upper lip is
silvery white. A broad dark brown or black mark extends posteriorly from
the orbit, encompassing the tympanum, to a point above the insertion of
the forelimb. The flanks are pale green or pale tan and marked with a
fine dark brown or black venation. The anterior surfaces of the thighs
usually are pale brown or grayish tan, sometimes having small,
indistinct darker flecks. The posterior surfaces of the thighs are
similarly colored, but in most specimens small but distinct dark flecks
are present; in some specimens small cream-colored spots are also
present on the posterior surfaces of the thighs. A distinct, narrow
creamy white anal stripe usually is present. A distinct white stripe is
present on the outer edge of the tarsus and fifth toe; on the tarsus the
white stripe is bordered below by dark brown. A white stripe also is
present on the outer edge of the forearm and fourth finger. In breeding
males the throat is dark gray.
Little geographic variation in color or pattern is evident. Few, if any,
specimens from the Pacific lowlands of South America are green in life.
(We have seen no living individuals from South America.) Some living
individuals from Costa Rica and all those seen alive from Nicaragua have
a tint of pale blue on the flanks. In some specimens the dorsal pattern
is so faint as to be barely discernible, whereas in most specimens the
pattern is bold.
The coloration in the living frogs is highly variable due to extreme
metachrosis. Individuals of this species are capable of changing the
dorsal coloration from green to brown in a short period of time. Both
green and brown individuals have been found active at night. Usually
those individuals found hiding by day are brown. One individual from
Finca La Sumbadora, Panama (now KU 91914), was kept alive in the
laboratory for nearly one month. This individual usually was pale green
with tan dorsal markings at night and tan with pale green markings by
day. On occasion the pale green dorsal markings were boldly outlined by
bright dark green.
In living individuals from throughout the range of the species the iris
is a bronze color, darkest medially with fine black reticulations.
_Natural History._--_Smilisca phaeota_ inhabits humid lowland tropical
forest and seldom ascends the foothills to more than 1,000 meters. The
rather equable climatic conditions, especially more or less evenly
distributed rainfall throughout the year, permit this frog to be active
most of the year. Dunn (1931:413) reported males calling on Barro
Colorado Island, Panama, in February and in July, and Breder (1946:416)
noted calling individuals in the Chucanaque drainage of Darien, Panama
in January, March, July, August and October and in Costa Rica in April
through August inclusively. Calling males were found at Bonanza,
Nicaragua in March and in July.
At all times of year the usual daytime retreats for these frogs are near
water; the frogs have been found in elephant ear plants (_Xanthosoma_)
and in bromeliads; occasional individuals have been found sitting on
shaded branches of bushes and trees. None has been observed on the
ground or beneath ground-cover by day.
The length of the breeding season cannot be determined definitely. The
earliest date on which eggs have been found is May 23; Gaige, Hartweg,
and Stuart (1937:5) reported a gravid female taken at El Recreo,
Nicaragua, in September, and we have a gravid female taken at Almirante,
Panama, in March.
Males usually call from secluded spots at the edge of water. All calling
males that we observed were on the ground within a few centimeters of
the water. The males usually are hidden beneath an overhanging leaf or
some other cover; they definitely do not sit in the open like _Smilisca
baudini_. Most calling males and clasping pairs have been found at the
edges of small pools or shallow ditches, although occasional individuals
are found at the edges of large ponds or streams.
The breeding call consists of one or two moderately short, low-pitched
notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20
seconds to several minutes. Each note is a low, vibrant "wauk," having
100 to 130 pulses per second and a dominant frequency of 330 to 420
cycles per second (Pl. 10C).