_Hyla microcephala_ Cope
_Diagnosis._--Lateral dark stripe narrow, covering only upper edge of
tympanum, bordered above by narrow white stripe; dorsal pattern
consisting of pair of longitudinal brown lines and no interorbital bar
(eastern populations), or of irregular dark markings forming an X- or
)(-shaped mark in scapular region and an interorbital bar (western
populations).
_Content._--The populations inhabiting the Pacific lowlands of
southeastern Costa Rica eastward to Colombia are recognized herein as
_Hyla microcephala microcephala_ Cope; the populations in western
Costa Rica northward to Mexico are assigned to _Hyla microcephala
underwoodi_ Boulenger.
_Distribution._--Southern Veracruz and northern Oaxaca southeastward
through the Atlantic lowlands of Central America to north-central
Nicaragua, thence southeastward on the Pacific lowlands to eastern
Panama, and thence into the Cauca and Magdalena valleys (Caribbean
drainage) of Colombia (Fig. 1).
[Illustration: Fig. 1. Map showing locality records for _Hyla
microcephala_.]
Table 1.--Variation in Certain Measurements and Properties in the
Hyla microcephala Group. (All Data Based on Adult Males;
Mean and Standard Error of Mean Below Observed Range.)
========================================================================
Locality | N | Snout-vent | Tibia length |Foot length|
| | length | ------------ | --------- |
| | (S-V L) | S-V L | S-V L |
------------------------------------------------------------------------
| _H. m. microcephala_
|
Panama: Canal Zone | 25 | 21.5-24.1 | 50.2-56.0 | 40.9-46.6 |
| | 22.8+-0.20 | 52.9+-0.37 | 43.5+-0.28 |
| | | | |
Costa Rica: Golfito | 25 | 18.5-24.5 | 49.1-54.4 | 41.8-48.0 |
| | 22.4+-0.27 | 51.6+-0.26 | 45.1+-0.32 |
|
| _H. m. underwoodi_
| |
Nicaragua: La Cumplida | 25 | 23.0-25.6 | 51.0-55.7 | 41.3-46.5 |
| | 24.1+-0.19 | 52.9+-0.25 | 43.7+-0.25 |
| | | | |
Guatemala: Finca Chama | 25 | 21.8-25.0 | 51.0-57.2 | 41.2-47.8 |
| | 23.5+-0.16 | 54.3+-0.39 | 44.4+-0.30 |
| | | | |
Tabasco: Teapa | 25 | 22.7-25.8 | 48.0-54.5 | 40.7-46.8 |
| | 24.3+-0.14 | 51.5+-0.29 | 43.3+-0.25 |
| | | | |
Oaxaca: Donaji-Sarabia | 25 | 22.1-25.9 | 49.8-55.6 | 40.5-46.6 |
| | 23.8+-0.19 | 52.8+-0.33 | 43.4+-0.27 |
| | | | |
Veracruz: Alvarado | 25 | 21.9-25.4 | 49.6-54.4 | 40.7-47.5 |
| | 24.1+-0.17 | 51.1+-0.28 | 42.6+-0.34 |
|
| _H. robertmertensi_
|
Guatemala: La Trinidad | 21 | 21.8-24.6 | 47.1-52.8 | 40.9-51.3 |
| | 23.4+-0.15 | 49.9+-0.34 | 43.5+-0.17 |
| | | | |
Chiapas: Acacoyagua | 25 | 21.4-25.7 | 47.8-52.4 | 41.7-46.3 |
| | 24.1+-0.20 | 50.4+-0.45 | 43.9+-0.23 |
| | | | |
Oaxaca: Tapanatepec | 25 | 22.4-26.4 | 44.1-48.3 | 39.1-44.5 |
| | 24.7+-0.18 | 46.4+-0.23 | 41.7+-0.23 |
|
| _H. phlebodes_
|
Panama: Canal Zone | 25 | 19.6-23.2 | 49.1-56.9 | 41.9-47.1 |
| | 22.2+-0.16 | 52.8+-0.35 | 45.4+-0.26 |
| | | | |
Costa Rica: Turrialba | 25 | 19.7-23.6 | 47.4-55.7 | 38.1-46.4 |
| | 22.0+-0.18 | 51.1+-0.35 | 42.8+-0.38 |
|
| _H. sartori_
|
Guerrero: Tierra Colorada| 25 | 23.7-26.0 | 47.2-51.4 | 42.4-47.8 |
| | 24.8+-0.13 | 49.6+-0.23 | 45.2+-0.27 |
------------------------------------------------------------------------
Table 1. (continued)
===============================================================
Locality | Head length | Head width | Tympanum
| ----------- | ---------- | --------
| S-V L | S-V L | Eye
---------------------------------------------------------------
| _H. m. microcephala_
|
Panama: Canal Zone | 28.5-32.8 | 28.1-30.9 | 44.0-54.1
| 31.0+-0.22 | 29.4+-0.11 | 49.0+-0.55
|
Costa Rica: Golfito | 30.2-35.5 | 29.0-32.7 | 40.0-57.8
| 33.1+-0.25 | 30.8+-0.16 | 48.4+-1.10
|
| _H. m. underwoodi_
|
Nicaragua: La Cumplida | 29.7-33.5 | 28.9-31.8 | 42.3-60.0
| 31.6+-0.19 | 30.4+-0.17 | 49.3+-0.97
|
Guatemala: Finca Chama | 30.8-35.3 | 29.6-33.6 | 37.5-56.4
| 33.0+-0.16 | 31.3+-0.36 | 45.2+-0.89
|
Tabasco: Teapa | 29.5-33.0 | 28.7-31.8 | 40.7-53.8
| 31.7+-0.17 | 30.3+-0.16 | 45.5+-0.38
|
Oaxaca: Donaji-Sarabia | 30.4-34.8 | 28.9-32.6 | 37.0-54.1
| 32.8+-0.19 | 30.8+-0.17 | 45.1+-0.76
|
Veracruz: Alvarado | 29.9-33.8 | 29.1-32.9 | 40.7-53.8
| 31.4+-0.18 | 30.5+-0.17 | 46.6+-0.65
|
| _H. robertmertensi_
|
Guatemala: La Trinidad | 30.0-33.3 | 27.3-29.8 | 44.4-50.0
| 31.3+-0.20 | 28.5+-0.23 | 47.4+-0.46
| | |
Chiapas: Acacoyagua | 29.1-32.7 | 26.0-30.3 | 42.8-53.8
| 31.2+-0.29 | 28.1+-0.20 | 46.5+-0.50
| | |
Oaxaca: Tapanatepec | 26.1-30.4 | 25.4-28.1 | 45.8-58.3
| 28.4+-0.16 | 26.8+-0.14 | 52.9+-0.77
|
| _H. phlebodes_
|
Panama: Canal Zone | 33.6-37.4 | 32.3-36.0 | 37.9-46.4
| 34.8+-0.18 | 33.8+-0.18 | 41.6+-0.49
| | |
Costa Rica: Turrialba | 32.6-35.9 | 30.5-35.0 | 35.7-48.2
| 34.1+-0.16 | 32.9+-0.17 | 40.1+-0.53
|
| _H. sartori_
|
Guerrero: Tierra Colorada| 29.4-31.8 | 28.9-31.0 | 42.3-52.0
| 30.6+-0.13 | 30.0+-0.12 | 47.4+-0.59
---------------------------------------------------------------
_Hyla microcephala microcephala_ Cope
_Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281, February
11, 1886 [Syntypes.--USNM 13473 (2 specimens, now lost) from
Chiriqui, Panama; Mr. MacNeil collector]; Bull. U.S. Natl. Mus.,
32:14, 1887. Guenther, Biologia-Centrali Americana, Reptilia and
Batrachia, p. 265, June, 1901. Dunn, Occas. Papers Boston Soc.
Nat. Hist., 5:413, October 10, 1931; Occas. Papers Boston Soc.
Nat. Hist., 8:72, June 7, 1933. Stebbins and Hendrickson, Univ.
California Publ. Zool., 56:524, February 17, 1959. Fouquette,
Evolution, 14:484, December 16, 1960. Busack, Copeia, 2:371,
June 21, 1966.
? _Hyla cherrei_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894,
p. 195, 1894 [Holotype.--location unknown, apparently lost;
type-locality: "Alajuela, Costa Rica;" R. Alfaro collector].
Guenther, Biologia Centrali-Americana: Reptilia and Batrachia,
p. 264, June, 1901. Taylor, Univ. Kansas Sci. Bull., 35:846,
July 1, 1952.
_Hyla underwoodi_, Ruthven, Misc. Publ. Mus. Zool., Univ. Michigan,
8:55, September 15, 1922. Barbour, Proc. New England Zool. Club,
10:31, March 2, 1928.
_Hyla microcephala microcephala_, Smith, Herpetologica, 7:185,
December 31, 1951. Taylor, Univ. Kansas Sci. Bull., 39:23,
November 18, 1958.
_Diagnosis._--Brown lateral stripe narrow, extending from nostril
along canthus, along upper edge of tympanum to groin, bordered above
by narrow white line; pair of dark brown longitudinal lines on dorsum
extending to vent; shanks having dark longitudinal line or flecks, no
transverse bars; interorbital dark mark lacking.
_Description and Variation._--The color pattern is nearly constant. Of
103 males from the Canal Zone, all lack an interorbital dark bar, and
all have a dark longitudinal line on the dorsal surface of the shank
and a narrow lateral dark stripe, bordered above by a narrow white
line, extending to the groin. The longitudinal dark lines on the
dorsum are continuous to the groin in 95 specimens and fragmented in
two specimens. In two others the lines converge and fuse in the
scapular region, and in four specimens auxiliary, fragmented lines are
present dorsolaterally.
In all specimens from southeastern Costa Rica (Golfito, Palmar Sur,
and Villa Neilly) the pattern is constant, except that in about 10 per
cent of the specimens the longitudinal line on the dorsal surface of
the shank is replaced by a row of brown flecks.
Of the limited number of Colombian specimens examined, all are
patterned normally, except three from Sautata, Choco, three from
Curumani, and three from Arcataca, Magdalena, which have flecks on the
dorsal surfaces of the shanks, and one from Espinal, Tolima, which has
no markings on the shanks.
When active at night most individuals are pale yellowish tan dorsally;
the white dorsolateral line is noticeable, but the brown lateral
stripe, dorsal brown lines, and lines on shanks are so pale that often
they are barely discernible. By day the dorsum changes to tan or pale
reddish brown; the stripes are dark brown, and the dorsolateral stripe
that is white at night becomes creamy yellow (Pl. 13). Small brown
flecks are present on the dorsum of most individuals. The venter
always is white, and the iris is pale bronze with a brown tint
immediately anterior and posterior to the pupil. In breeding males the
vocal sac is pale yellow.
_Tadpoles._--Tadpoles of this species have been found in weed-choked
ponds in eastern Panama Province. The following description is based
on KU 104097, a specimen in developmental stage 34 (Gosner, 1960).
Total length, 20.5 mm.; body length, 8.2 mm.; body slightly wider
than deep; snout pointed; nostrils large, situated dorsally, much
closer to snout than eyes, directed anteriorly; eyes moderately
small, situated dorsolaterally and directed laterally; spiracle
sinistral, located just posteroventral to eye; anal tube dextral.
Tail xiphicercal; caudal musculature moderately deep, becoming
slender posteriorly, extending beyond caudal fin; fins deepest at
about one-third distance from body to tip of tail; dorsal fin
extending onto body, deeper than deepest part of caudal
musculature; ventral fin slightly shallower than musculature.
Mouth small, terminal, lacking teeth and fringing papillae, but
having finely serrate beaks. In preservative, top of head pale
brown; dark stripe from tip of snout through eye to posterior edge
of body, narrowing to thin line on proximal one-fourth of tail;
venter white; tail creamy tan with fine black flecks most numerous
posteriorly; posterior two-thirds of fins edged with black. In
life, top of head yellowish tan; lateral stripe brown; belly
white; anterior half of tail lacking pigment; posterior half deep
orange; iris pale bronze (Pl. 15).
_Remarks._--Evidence for intergradation of _Hyla microcephala_ with
_H. underwoodi_ is provided by four specimens [USC 818 (2), 6081-2]
from 6.1 kilometers northeast of the mouth of the Rio Tarcoles, and
nine specimens [USC 8254 (2), 8255, 8256 (4), 8258 (2)] from Parrita,
both in Puntarenas Province, Costa Rica. These localities lie about
two-thirds the distance from the northwesternmost locality for _H.
m. microcephala_ (Palmar Sur) to the southeasternmost locality for
_H. m. underwoodi_ (Barranca). Although in most aspects of coloration
the frogs are more nearly like _H. m. underwoodi_ than _H. m.
microcephala_, some specimens have longitudinal lines on their shanks,
such as are characteristic of _H. m. microcephala_. The dorsal pattern
varies from nearly complete longitudinal lines to broken lines, fused
into an X-shaped scapular mark or not.
As noted by Rivero (1961:135), _Hyla microcephala_ seems to be closely
related to _Hyla misera_ Werner, a species having a wide distribution
east of the Andes in South America. Despite the similarity in color
pattern, size, and structure, we are reluctant to place the two taxa
in the same species until data on coloration in life, mating calls,
and life history are available for _Hyla misera_ and compared with
those of _Hyla microcephala_.
The status of Cope's _Hyla cherrei_ is questionable. Since the type,
the only specimen ever referred to the species, apparently is lost,
the only extant information regarding the taxon is contained in the
original description (Cope, 1894). There the species was characterized
as having a narrow dorsolateral white stripe and lacking pigment on
the upper arms and thighs. These characteristics of the color pattern
combined with the statements "vomerine teeth few, opposite the middle
of the very large choanae" and "tympanic drum distinct, one half the
area of eye" serve to distinguish _H. cherrei_ from all other Costa
Rican hylids, except _H. m. microcephala_ and _H. m. underwoodi_. No
statements in the type description will definitely associate _cherrei_
with one or the other of these subspecies. Since it seems obvious that
_H. cherrei_ can be associated with _H. microcephala_, we prefer to
place the name in the synonymy of the nominate subspecies, thereby
preserving the commonly used name _H. underwoodi_ (Boulenger, 1899) as
a subspecies of _H. microcephala_.
_Distribution._--_Hyla microcephala microcephala_ inhabits coastal
lowlands from the area of Golfo Dulce (apparently absent from the
Osa Peninsula) in southeastern Costa Rica eastward in Panama,
including the Azuero Peninsula to northern Colombia and thence
southward in the valleys of the Rio Cauca and Rio Magdalena in
Colombia (Fig. 1). Except for the central area of the Canal Zone
the subspecies is unknown from the Caribbean drainage in Central
America, but in Colombia the subspecies occurs only in the
Caribbean drainage. In Central America this frog occurs mostly on
the coastal lowlands; the highest recorded elevation is 560 meters
at El Valle, Cocle, Panama. Throughout most of its range _Hyla
microcephala microcephala_ occurs in disturbed habitats--cut-over
forests, secondary growth, and pastureland. It does not seem to be
an inhabitant of either primary forest or of _Curatella_-savanna.
_Specimens examined._--522, as follows: +Costa Rica+: Puntarenas:
Golfito, KU 32172-207; 3 km. E Golfito, KU 86399, USC 2757-8;
Palmar Sur, KU 64591-608, USC 2650 (14), UU 3907-32; *1.5-2.5 km.
ESE Palmar Sur, KU 68293-7 (skeletons), 93957-62; Parrita, USC
8254 (2), 8255, 8256 (4), 8258 (2) [intergrades with _H. m.
underwoodi_]; 3 km. NW Piedras Blancas, KU 103689; 6.1 km. NE
mouth of Rio Tarcoles, USC 818 (2), 6081-2 [intergrades with _H.
m. underwoodi_]; Villa Neilly, USC 2651; *1-5 km. WNW Villa
Neilly, USC 6182-4, 8003 (4), 8031 (3), 8032; *10.5 km. WNW Villa
Neilly, KU 64609-27, 68398 (eggs).
+Panama+: Canal Zone: Albrook Air Base, TNHC 23389, 23497; Balboa,
ANSP 19555-6; *Fort Clayton, UIMNH 42008-12; *2.8 km. SW Fort
Kobbe, KU 96015-25; *Frijoles, MCZ 19208; *Bamboa, MCZ 21507; *8.3
km. N Gatun Locks, TNHC 23441; *Juan Diaz, MCZ 13747; *Juan Mina,
AMNH 55436-7, ANSP 21811-2, UMMZ 126734, 126735 (6), UU 3900-6;
*8-14 km. N Miraflores Locks, TNHC 23374-88, 23390-409, 23411-38,
23440, 23442-60, 23462-76; 23478-83, 23492, 23555-60, 23562-76;
*Rio Chagres, AMNH 55430, 55439; *Rio Cocoli, 3.5 km. N Miraflores
Locks, TNHC 23410; *Summit, ANSP 23365-71, FMNH 22966-9, KU
97783-87. Chiriqui: 5.5 km. E Concepcion, AMNH 69772; *14.4 km. E
Concepcion, AMNH 69773-8; 2 km. S David, AMNH 69779; *Progreso,
UMMZ 58252, 58253 (2), 58254, 58436; Rio Gariche, 8.3 km. ESE Paso
Canoas, KU 103065-8. Cocle: 1 km. SE El Cano, KU 103042-51; El
Valle de Anton, AMNH 59614-18 (10), 69785, ANSP 23502-5, KU
77201-14, MVZ 66578-83, UIMNH 46532. Colon: Cement Plant,
Transisthmian Highway, FMNH 60394-5. Darien: El Real, KU 80454-5,
103052-64, UMMZ 125036 (10), USNM 140567-8; Rio Canclon at Rio
Chucunaque, UMMZ 125035; *Rio Chucunaque, near Yavisa, AMNH 59523.
Los Santos: Tonosi, KU 101606-9. Panama: 5 km. S Bejuco, AMNH
69782; 3 km. W Chepo, KU 77172-4, 104097-8 (tadpoles); *6 km. WSW
Chepo, KU 77175; *Chico, Rio La Jagua, USNM 129070; *La Joya,
Cacora, ANSP 25129-33; Madden Dam, FMNH 67819; Nueva Gorgona, AMNH
69780-1; *1.6 km. W Nueva Gorgona, AMNH 69783-4; 1.5 km. W Pacora,
77176-200; *Rio La Laja, near Chame, ANSP 21845; *Rio Tapia, MCZ
10048; *Tapia, AMNH 18930, 18950, 18952-3; *18 km. E Tocumen, MVZ
78662.
+Colombia+: Choco: Sautata, Atrato, FMNH 74918 (2), 74919.
Magdalena: Aracataca, ANSP 19755-7; Curumani, MCZ 21465-74, UIMNH
28855; UMMZ 90168, USNM 118247; El Banco, Rio Magdalena, ANSP
25061; Fundacion, UMMZ 48281-2. Tolima: Espinal, MCZ 15068;
Mariquita, FMNH 81822-3. Valle: Sevilla, MCZ 13751-3.
_Hyla microcephala underwoodi_ Boulenger
_Hyla microcephala_ Boulenger, Proc. Zool. Soc. London, p. 481,
October 1, 1898 [Syntypes.--BMNH 94. 11. 1532-33 from Bebedero,
Guanacaste Province, Costa Rica; C. F. Underwood collector] (not
_Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281,
February 11, 1886, from Chiriqui, Panama).
_Hyla underwoodi_ Boulenger, Ann. Mag. Nat. Hist., ser. 7, 3:277,
April, 1899 (substitute name for _Hyla microcephala_ Boulenger,
preoccupied). Guenther, Biologia-Centrali Americana, Reptilia and
Batrachia, p. 278, September, 1901. Dunn and Emlen, Proc. Acad.
Nat. Sci. Philadelphia, 84:25, March 22, 1932. Stuart, Misc. Publ.
Mus. Zool., Univ. Michigan, 29:39, October 1, 1935. Taylor, Proc.
Biol. Soc. Washington, 50:44, April 21, 1937. Stuart, Occas.
Papers Mus. Zool., Univ. Michigan, 471:15, May 17, 1943. Taylor
and Smith, Proc. U. S. Natl. Mus., 95:586, June 30, 1945. Stuart,
Misc. Publ. Mus. Zool., Univ. Michigan, 69:35, June 12, 1948.
Smith and Taylor, Bull. U. S. Natl. Mus., 194:85, June 17, 1948;
Univ. Kansas Sci. Bull., 33:316, March 20, 1950. Stuart, Contr.
Lab. Vert. Biol., Univ. Michigan, 45:48, May, 1950. Taylor, Univ.
Kansas Sci. Bull., 35:891, July 1, 1952; Univ. Kansas Sci. Bull.,
39:25, November 18, 1958.
_Hyla phlebodes_, Cole and Barbour, Bull. Mus. Comp. Zool., 50:154,
November, 1906. Kellogg, Bull. U. S. Natl. Mus., 160:172,
March 31, 1932.
_Hyla microcephala martini_ Smith, Herpetologica, 7:187, December
31, 1951 [Holotype.--UIMNH 20965 from Encarnacion, Campeche,
Mexico; H. M. Smith collector]. Stuart, Contr. Lab. Vert. Biol.,
Univ. Michigan, 68:46, November, 1954. Fugler and Webb,
Herpetologica, 13:105, July 10, 1957. Stuart, Contr. Lab. Vert.
Biol., Univ. Michigan, 75:17, June, 1958. Neill and Allen, Publ.
Research Div., Ross Allen's Reptile Inst., 2:26, November 10,
1959. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13:62,
August 16, 1960. Stuart, Herpetologica, 17:74, July 11, 1961.
Hensley and Smith, Herpetologica, 18:70, April 9, 1962. Stuart,
Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, April 2, 1963.
Holman and Birkenholz, Herpetologica, 19:144, July 3, 1963.
Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 15:225, October 4,
1963; Univ. Kansas Publ., Mus. Nat. Hist., 15:588, June 22, 1965.
_Hyla microcephala underwoodi_, Smith, Herpetologica, 7:188,
December 31, 1951.
_Diagnosis._--Brown lateral stripe narrow, extending to groin or
only to sacral region, bordered above by narrow white line; dorsal
pattern bold, consisting of X- or )(-shaped mark in scapular
region or pair of interconnected dark lines on back; interorbital
dark mark usually present; shanks usually having dark transverse
bars.
_Description and Variation._--The dorsal color pattern is highly
variable. The various permutations of the X-shaped scapular mark
and dark sacral marks differ proportionately in different samples.
The variation in color pattern in 12 samples is summarized in
Table 2. In samples from the southern part of the range (southern
Nicaragua and Guanacaste Province, Costa Rica) more (40-93%)
individuals have the lateral stripes extending to the groin than
in northern samples (0-42%) from southern Mexico and Guatemala.
Likewise, the percentage of specimens lacking bars on the shanks and
a dark interorbital bar is higher in the Costa Rican samples than
elsewhere in the range. The X- or )(-shaped scapular markings and
/\- or / \-shaped sacral markings are most prevalent in northern
samples, whereas to the south the dorsal markings are more commonly
arranged in a pattern of paired lines, which usually are discontinuous
and usually extend posteriorly only to the sacral region. Thus, the
color pattern in _H. m. underwoodi_ in the southern part of its range
shows trends towards the pattern characteristic of _H. m.
microcephala_. Intergrades between these two subspecies have
been discussed in the account of the nominate subspecies.
Table 2.--Variation in Color Pattern in Hyla microcephala underwoodi
==========================================================================
Population | N | Shanks || Interorbital || Dorsolateral |
| | || bar || stripe |
| |-------------||----------------||--------------|
| | Bars |Flecks|| Present| Absent|| Groin| Sacrum|
--------------------------------------------------------------------------
Oaxaca: | 27 | 22 | 5 || 27 | 0 || 0 | 27 |
Donaji-Sarabia | | | || | || | |
| | | || | || | |
Tabasco: | 55 | 46 | 9 || 55 | 0 || 0 | 55 |
Teapa-Villahermosa| | | || | || | |
| | | || | || | |
Guatemala: | 51 | 51 | 0 || 51 | 0 || 17 | 34 |
La Libertad | | | || | || | |
| | | || | || | |
Guatemala: | 32 | 32 | 0 || 32 | 0 || 0 | 32 |
Finca Chama | | | || | || | |
| | | || | || | |
Guatemala: | 31 | 31 | 0 || 31 | 0 || 14 | 17 |
Puerto Barrios | | | || | || | |
| | | || | || | |
Honduras: | 13 | 13 | 0 || 13 | 0 || 9 | 4 |
Lago Yojoa | | | || | || | |
| | | || | || | |
Nicaragua: | 56 | 44 | 12 || 54 | 2 || 13 | 43 |
La Cumplida | | | || | || | |
| | | || | || | |
Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 |
Tipitapa | | | || | || | |
| | | || | || | |
Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 |
Santo Thomas | | | || | || | |
| | | || | || | |
Costa Rica: | 12 | 0 | 12 || 6 | 6 || 7 | 5 |
Tenorio-Tilaran | | | || | || | |
| | | || | || | |
Costa Rica: | 38 | 21[A]| 15 || 34 | 4 || 25 | 13 |
Las Canas-Liberia | | | || | || | |
| | | || | || | |
Costa Rica: | 32 | 26 | 6 || 29 | 3 || 30 | 2 |
Esparta | | | || | || | |
--------------------------------------------------------------------------
==========================================================================
Population | Scapular markings || Sacral |
| || markings |
|----------------------------||----------------------|
| X | )( | ][ | Other || /\ | / \ | Other |
--------------------------------------------------------------------------
Oaxaca: | 23 | 4 | 0 | 0 || 7 | 6 | 14 |
Donaji-Sarabia | | | | || | | |
| | | | || | | |
Tabasco: | 53 | 2 | 0 | 0 || 19 | 11 | 23 |
Teapa-Villahermosa| | | | || | | |
| | | | || | | |
Guatemala: | 45 | 6 | 0 | 0 || 16 | 14 | 21 |
La Libertad | | | | || | | |
| | | | || | | |
Guatemala: | 32 | 0 | 0 | 0 || 26 | 2 | 4 |
Finca Chama | | | | || | | |
| | | | || | | |
Guatemala: | 23 | 0 | 4 | 4 || 6 | 4 | 21 |
Puerto Barrios | | | | || | | |
| | | | || | | |
Honduras: | 3 | 2 | 3 | 5 || 2 | 1 | 10 |
Lago Yojoa | | | | || | | |
| | | | || | | |
Nicaragua: | 11 | 35 | 8 | 2 || 0 | 19 | 37 |
La Cumplida | | | | || | | |
| | | | || | | |
Nicaragua: | 0 | 5 | 3 | 2 || 0 | 3 | 7 |
Tipitapa | | | | || | | |
| | | | || | | |
Nicaragua: | 3 | 0 | 7 | 0 || 0 | 5 | 5 |
Santo Thomas | | | | || | | |
| | | | || | | |
Costa Rica: | 0 | 0 | 12 | 0 || 0 | 0 | 12 |
Tenorio-Tilaran | | | | || | | |
| | | | || | | |
Costa Rica: | 0 | 11 | 19 | 8 || 0 | 0 | 38 |
Las Canas-Liberia | | | | || | | |
| | | | || | | |
Costa Rica: | 0 | 0 | 14 | 18 || 0 | 0 | 32 |
Esparta | | | | || | | |
--------------------------------------------------------------------------
[Footnote A: Longitudinal stripes present in two specimens.]
When this frog is active at night its dorsum is pale yellow; faint
flecks are present in some individuals. The white dorsolateral line
usually is evident in the tympanic region, but in many individuals a
dorsal pattern of lines and other marks is not evident. By day the
dorsum changes to yellowish tan or pale brown with dark brown or
reddish brown markings (Pl. 13). The venter is white, and the vocal
sac in breeding males is yellow. The iris is pale bronze with a brown
tint anterior and posterior to the pupil.
_Remarks._--_Hyla microcephala underwoodi_ has had a confused
nomenclatural history. The taxon was first named _Hyla microcephala_
by Boulenger (1898); this name was preoccupied by _Hyla microcephala_
Cope (1886). Cole and Barbour (1906) and Kellogg (1932) used the name
_Hyla phlebodes_ Stejneger (1906) for specimens of this frog from
Mexico. Dunn (1931, 1933, 1934) applied the name _Hyla underwoodi_ to
Panamanian specimens that we identify as _Hyla phlebodes_. Smith
(1951) named _Hyla microcephala martini_ from southern Mexico and
Guatemala and considered the northern populations to represent a
subspecies distinct from the Costa Rican _Hyla microcephala
underwoodi_, despite the fact the Stuart (1935:39) stated that
comparisons of specimens from El Peten, Guatemala, with the holotype
of _Hyla underwoodi_ showed only trivial differences.
Much of the confusion regarding the name _Hyla underwoodi_ stems from
the illustration given by Boulenger (1898:pl. 39, fig. 3) and
reproduced by Taylor (1952:892), which shows a frog having a unicolor
dorsum, dorsolateral white lines, and dark flanks. This pattern is in
marked contrast to the pattern seen in most preserved specimens, which
have the dorsum variously marked by dark brown lines or irregular
marks. Smith (1951:185), in his description of _Hyla microcephala
martini_ from southern Mexico, considered _H. underwoodi_ to be a
subspecies of _H. microcephala_ that lacked dorsal dark markings.
Data accumulated in 1961 through field studies by the senior author at
the type locality, Bebedero, and other localities in Guanacaste and
Puntarenas provinces in Costa Rica provide a reasonable explanation of
the differences in color pattern. As noted in the preceding description
of this subspecies, at night the dorsal markings are not evident in
many living individuals, whereas by day the dorsal markings are
prominent. Most collectors prepare their specimens by day; consequently
the majority of specimens have a pronounced dorsal pattern. Of the
frogs collected in Costa Rica in 1961, some specimens were preserved
at night; others from the same series were preserved by day. The
differences are striking. In those preserved at night, dorsal markings
are faint, if present at all. Some specimens closely match the figure
given by Boulenger (1898).
It is extremely doubtful if the frog described and illustrated by
Boulenger could be associated with either _Hyla phlebodes_ or _H.
microcephala microcephala_. Individuals of the former species lack
a dorsolateral white line and always have some dorsal markings
evident at night; furthermore, _H. phlebodes_ is not known to occur
on the Pacific lowlands. _Hyla microcephala microcephala_ occurs
farther southeast. Since there is no reason to doubt the type locality
of _H. underwoodi_, since specimens from the area around the type
locality that have been preserved at night are like the holotype in
pattern, and since the characteristics of the populations of the frogs
in Guanacaste are the same as, or gradually blend into those of,
populations in northern Central America and southern Mexico, the
frogs from throughout the entire range can be referred to one taxon,
the earliest name for which is _Hyla underwoodi_ Boulenger, which
herein is considered to be a subspecies of _H. microcephala_ Cope.
_Distribution._--_Hyla microcephala underwoodi_ inhabits the
Atlantic slopes and lowlands from southern Veracruz and extreme
northern Oaxaca eastward across the base of the Yucatan Peninsula
(possibly the species is extant in the northern part of the
peninsula) to British Honduras and thence southeastward through
the Caribbean lowlands and interior valleys in Honduras to central
Nicaragua, where it apparently avoids the forested Caribbean
lowlands and the dry Pacific lowlands of northwestern Nicaragua,
but in the vicinity of Managua invades the Pacific lowlands and
continues southward into northwestern Costa Rica as far as the
Puntarenas Peninsula (Fig. 1). In Mexico and Guatemala the species
has not been taken at elevations of more than 350 meters, whereas
farther south it occurs at higher elevations--780 meters at
Silencio, Costa Rica, 830 meters on Montana de Guaimaca, Honduras,
960 meters at Finca Tepeyac, Nicaragua, and 1200 meters at Finca
Venecia, Nicaragua.
_Specimens examined._--1270, as follows: +Mexico+: Campeche: Balchacaj,
FMNH 100406, UIMNH 20944-6; Encarnacion, FMNH 27069-70, 75784,
MCZ 28360, 29637, UIMNH 20948-58, 20965, USNM 134264-5; Escarcega,
UMMZ 122999; *7.5 km. W Escarcega, KU 71229-43; Laguna Alvarado, 65 km.
S Xpujil, KU 75084-9; Pacaitun, Rio Candelaria, FMNH 83118-20;
*Tres Brazos, FMNH 113101-22, UIMNH 20947; 10 km. W Xpujil, KU 75082-3.
Chiapas: Palenque, UIMNH 47984, 49139-50, USNM 114973-8. Oaxaca: *5 km.
N Chiltepec, KU 87015-23; 3 km. N Donaji UMMZ 115249 (9); *3.7 km.
N Donaji, UMMZ 115250 (5); *43 km. N Matias Romero, UIMNH 42550-68;
*3.5 km. N Palomares, TNHC 25185, 25321-31, 25341-68; 4.6 km. N
Sarabia, UMMZ 115247 (2); *6.1 km. N Sarabia, UMMZ 115248 (11), *3 km.
N Tolocita, KU 39655; Tuxtepec, KU 87024-40. Tabasco: 24 km. N
Frontera, MCZ 35665-70; 0.8 km. E Rio Tonola, TNHC 25189; Teapa, UMMZ
119218 (4); *2.7 km. N Teapa, UMMZ 119216 (4); *10 km. N Teapa, UMMZ
119217 (6); *11.5 km. N Teapa, UMMZ 119219; *15.2 km. N Teapa, UMMZ
119220 (4); *17.6 km. N Teapa, UMMZ 119221 (12), 3.3 km. S Villahermosa,
UMMZ 119215 (12), *17.6 km. S Villahermosa, UMMZ 119214 (12).
Veracruz: 2.1 km. N Acayucan, UIMNH 42547-9; *6.4 km. NW Acayucan,
UMMZ 115254 (14); 1.6 km. ESE Alvarado, UMMZ 115258 (39); *2.4 km. ESE
Alvarado, UMMZ 115251 (2); *4.5 km. S Aquilera, UMMZ 115252 (21);
*8 km. SW Coatzacoalcos, UMMZ 119213 (10); 2.2 km. E Cosoleacaque,
UMMZ 119222 (26); 10 km. SE Hueyapan, UMMZ 115255; 0.8 km. S Lerdo de
Tejada, UMMZ 122778; *3.6 km. NE Minatitlan, TNHC 25150-2; 1.9 km. S
Naranja, UMMZ 115253 (3); 4.5 km. NE Novillero, UMMZ 115256; San Andres
Tuxtla, FMNH 113124-8, UIMNH 20942-3. Yucatan: Chichen-Itza, FMNH 36570,
MCZ 2463 (2).
+British Honduras+: Cayo: 6.2 km. S El Cayo, MCZ 37885-92. Stann Creek:
Stann Creek, FMNH 49068.
+Guatemala+: Alta Verapaz: 28.3 km. N Campur, KU 64578-90; Chinaja,
KU 57425; Cubilquitz, UMMZ 90887, 90888 (4); Finca Chama, UMMZ 90879
(13), 90880 (4), 90881, 90882 (28), 90883 (12), 90884 (46), 90885 (39),
90886 (20); *Finca Tinaja, BYU 16032; Panzos, UMMZ 90889 (2).
Chiquimula: Chiquimula, UMMZ 98113; 2 km. N Esquipulas, UMMZ 106844.
El peten: La Libertad, KU 57447-97, 59907-11 (skeletons), MCZ 21461,
UMMZ 75332 (13), 75333 (11), 75334 (14), 75335 (10); Piedras Negras,
FMNH 113123, UIMNH 20966; *5 km. S Piedras Negras, USNM 114951-72;
Tikal, UMMZ 117981 (2); Toocog, 15 km. SE La Libertad, KU 57426-46. El
Quiche: Finca Tesoro, UMMZ 89165 (5). Huehuetenango: Finca San Rafael,
16 km. SE Barillas, FMNH 40917-9. Izabal: Puerto Barrios, FMNH
20004-7; 8 km. S Puerto Barrios, KU 57507-37, 59991 (eggs), 59992
(tadpoles); Quirigua, CAS 69657-701; 2.5 km. NE Rio Blanco, KU 57539;
San Felipe, FMNH 35065. Zacapa: 14 km. ENE Mayuelas, KU 57502-6; 8 km.
ENE Rio Hondo, KU 57498-501.
+Honduras+: Atlantidad: La Ceiba, UMMZ 91948 (2), USNM 117593-600;
Lancetilla, MCZ 17981. Cortes: Lago Yojoa, AMNH 54917-9, 54957, 55134,
KU 64563-77. El Paraiso: Valle de Jamastran, AMNH 54807-12.
Francisco-Moranza: El Zamorano, AMNH 54873-81, KU 103223, UMMZ 123101;
Montana de Guaimaca, AMNH 54900-4 (8); Ranch San Diego, 19 km. SW
Guaimaca, AMNH 53939. Itibuca: Vieja Itibuca, AMNH 54912-3.
+Nicaragua+: Chontales: 3 km. SW Santo Tomas, KU 64770-9, 68308
(skeleton). Esteli: Finca Venecia, 7 km. N, 16 km. E Condega, KU
85296; 2.4 km. N Esteli, MCZ 28933-7. MANAGUA: 12-13 km. E Managua,
KU 85297-301; *10 km. SW Tipitapa, UMMZ 119977 (10). Matagalpa: *Finca
Tepeyac, 10.5 km. N, 9 km. E Matagalpa, KU 85302-3; Hacienda La
Cumplida, KU 64780-96, 68309-11 (skeletons), UMMZ 116482 (8), 116483
(23), 116484 (3), 116485 (5), 119984 (3). Rivas: *Finca Amayo, 13 km.
S, 14 km. E Rivas, KU 85304-7; 16 km. S Rivas, MCZ 29011-7; *20.5 km.
SE Rivas, KU 85308-10; 5 km. SE San Pablo, KU 43111-4.
+Costa Rica+: Guanacaste: Arenal, USC 6254 (2); *3 km. W Bagaces, USC
7019 (10); *3 km. NE Boca del Barranca, USC 8017 (21), *Finca San
Bosco, USC 6272 (6), 6276 (3); *Guayabo de Bagaces, USC 7022 (4), 7023
(3), 7025; 12 km. S La Cruz, USC 8091 (2); *Laguna Arenal, USC 6262;
*27 km. N Las Canas, USC 8171 (6); *16 km. E Las Canas, KU 102252-8;
16 km. SSE Las Canas, KU 65090-5; *20 km. SE Las Canas, KU 102251;
Liberia, KU 30827-39; *7.3 km. N Liberia, USC 8096 (4); *10 km. N
Liberia, USC 8085 (9); *7.5 km. SE Liberia, KU 65102-8, 68621-2
(skeletons); *14.7 km. S Liberia, USC 8238 (3); *4 km. W Liberia, KU
36847-57; 2 km. S Nicoya, USC 8230; *3-10 km. ESE Playa del Coco, USC
8012 (16), 8137 (14); *21.6 km. ESE Playa del Coco, USC 8138 (13);
*Penas Blancas, KU 102247-50; *Rio Bebedero, 5 km. S Bebedero, KU
65089; *Rio Higueron, USC 7168 (2); Santa Cruz, USC 8232 (2);
*Silencio, USC 6248; *Tenorio, KU 32313; Tilaran, KU 36858-60; *2 km.
E Tilaran, KU 86403, *5 km. NE Tilaran, KU 36840-6 USC 6269.
Puntarenas: Barranca, KU 32305-12, *5 km. WNW Barranca, UMMZ 119976
(2); *10 km. E Esparta, KU 86400-2; 1 km. WNW Esparta, KU 65101; *4 km.
WNW Esparta, KU 65088; *10 km. WNW Esparta, KU 65063-87, 68616-20
(skeletons); *12 km. WNW Esparta, KU 65096-100, USC 8251; 21.8 km.
W San Ramon, USC 8242 (15).
+Hyla robertmertensi+ Taylor
_Hyla robertmertensi_ Taylor, Proc. Biol. Soc. Washington, 50:43,
April 21, 1937 [Holotype.--CNHM 100096 (formerly EHT-HMS 2270)
from Tapachula, Chiapas, Mexico; H. M. Smith and E. H. Taylor
collectors]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:84,
June 17, 1948; Univ. Kansas Sci. Bull., 33:326, March 20, 1950.
Mertens. Senckenbergiana, 33:170, June 15, 1952;
Senckenbergischen Naturf. Gesell., 487:30, December 1, 1952.
Stuart, Contr. Lab. Vert. Biol., Univ. Michigan, 68:47,
November, 1954. Duellman, Univ. Kansas Publ., Mus. Nat. Hist.,
13:63, August 16, 1960. Duellman and Hoyt, Copeia, 1961 (2): 417,
December 19, 1961. Porter, Herpetologica, 18:168, October 17,
1962. Stuart, Misc. Publ. Mus. Zool., Univ. Michigan, 122:36,
April 2, 1963. Duellman and Trueb, Univ. Kansas Publ., Mus. Nat.
Hist., 17:348, July 14, 1966.
_Diagnosis._--Brown lateral stripe wide, including loreal region and
entire tympanum, extending to groin, bordered above by narrow white
line; dorsum unicolor or with pair of dark lines (or rows of dashes)
usually extending only to the sacral region; shanks having dark flecks,
no transverse bars; interorbital bar lacking.
_Description and Variation._--Males attain a maximum snout-vent length
of 26.4 mm. in Oaxaca, whereas in a sample from Acacoyagua, Chiapas,
the largest male has a snout-vent length of 25.7 mm., and from La
Trinidad, Guatemala, 24-6 mm. Specimens from the western part of the
range (eastern Oaxaca) have slightly smaller heads and proportionately
larger tympani than the more eastern populations (Table 1).
The color pattern shows little variation, except in the nature of the
dorsal markings. In a few specimens from throughout the range, but
especially in the eastern part of the range, the dorsum lacks markings
between the dorsolateral white lines. In most specimens the dorsal
pattern consists of flecks or dashes arranged in two parallel
longitudinal rows, and in some specimens the marks are fused into
parallel lines. Small brown flecks are present on the dorsal surfaces
of the shanks; in some specimens these flecks tend to form a
longitudinal stripe on the shank. An interorbital dark mark is
invariably absent.
When active at night _Hyla robertmertensi_ is pale yellow above with a
white dorsolateral line and pale brown lateral stripe; the dorsal
markings are faint. By day the dorsum is yellowish tan with brown
markings. The dorsolateral stripe is creamy white, and the lateral
stripe is dark brown (Pl. 14). The venter is white, and the iris is
dull bronze. In breeding males the vocal sac is yellow.
_Remarks._--Although this species superficially resembles _Hyla
microcephala microcephala_, the latter is easily distinguished by the
narrow brown lateral stripe, as compared with the much wider stripe
in _H. robertmertensi_. No other hylids in northern Central America
and southern Mexico can be confused with this species.
_Distribution._--_Hyla robertmertensi_ inhabits the Pacific slopes (to
elevations of 700 meters) and lowlands from eastern Oaxaca (east of
the Plains of Tehuantepec) southeastward to central El Salvador. The
species also occurs in the Cintalapa Valley (Atlantic drainage) in
southwestern Chiapas (Fig. 2.) The distribution seems to be limited on
the northwest and southeast by arid environments. The region in which
_Hyla robertmertensi_ lives is characterized by higher rainfall and
more luxuriant vegetation than occur on the Plains of Tehuantepec or
on the Pacific lowlands of eastern El Salvador and southern Honduras.
In addition to the localities listed below, Mertens (1952:30)
recorded the species from Hacienda Cuyan-Cuya, Depto. Sonsonate, El
Salvador.
[Illustration: Fig. 2. Map showing locality records for
_Hyla robertmertensi_.]
_Specimens examined._--490, as follows: +Mexico+: Chiapas: Acacoyagua,
USNM 114754-61; *2 km. W Acacoyagua, UMMZ 87843 (28), 87844 (50),
87845 (50), 87846 (45), 87847 (27), 87848 (3); 32 km. N Arriaga, KU
57619-24, 59917-8 (skeletons); Asuncion, FMNH 100413, 100501-4, UIMNH
26989-90, USNM 134267; *La Esperanza, USNM 114737-48, 114750-3, 17 km.
S Las Cruces, KU 57625-49, 59997 (eggs); 8.5 km. N Puerto Madero, UMMZ
119981 (2); *11.7 km. N Puerto Madero, UMMZ 119982; Tapachula, FMNH
100096, UIMNH 26987; *11 km. S Tapachula, KU 57605-18, 59916 (skeleton);
Tonola, FMNH 27073, 100505-10, UIMNH 26988. Oaxaca: Tapanatepec,
UMMZ 115245 (2), *1.6 km. E Tapanatepec, UMMZ 115244 (14); *4.3 km.
E Tapanatepec, UIMNH 38368-9; *7.5 km. W Tapanatepec, UMMZ 115246
(39); 12.8 km. W Tapanatepec, KU 65007-14; 7.2 km. WNW Zanatepec,
UMMZ 115243 (77); *13.6 km. WNW Zanatepec, TNHC 25213-22; 22.7 km.
WNW Zanatepec, TNHC 25203-9.
+Guatemala+: Jutiapa: Jutiapa, UMMZ 106848; La Trinidad, UMMZ
107733 (23). Retalhueleu: Casa Blanca, UMMZ 107732.
+El Salvador+: La Libertad: 16 km. NW Santa Tecla, KU 44112. San
Salvador: 21.9 km. N San Salvador, UMMZ 119983 (6).
+Hyla phlebodes+ Stejneger
_Hyla phlebodes_ Stejneger, Proc. U. S. Natl. Mus., 30:817, June 4,
1906 [Holotype.--USNM 2997 from "San Carlos," Costa Rica;
Burgdorf and Schild collectors]. Taylor, Proc. Biol. Soc.
Washington, 50:44, April 21, 1937; Univ. Kansas Sci. Bull.,
35:888, July 1, 1952; Univ. Kansas Sci. Bull., 39:25, November
18, 1958. Fouquette, Evolution, 14:484, December 16, 1960.
Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. Hist., 17:348,
July 14, 1966.
_Hyla underwoodi_, Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413,
October 10, 1931; Occas. Papers Boston Soc. Nat. Hist. 8:72,
June 7, 1933; Amer. Mus. Novitiates, 747.2, September 17, 1934,
Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool., Univ.
Michigan, 357:5, October 26, 1937. Breder, 1946, Bull. Amer. Mus.
Nat. Hist., 86:416, August 22, 1946.
_Diagnosis._--Dark brown lateral stripe, if present, usually extending
only to insertion of forearm, never posteriorly to sacral region;
white line above brown stripe absent or faint; dorsal pattern weak,
usually consisting of irregular dashes or interconnected lines;
interorbital dark mark present; shanks having weakly defined
transverse bars.
_Description and variation._--In the majority of specimens (70%) the
lateral dark stripe extends from the nostril to the eye and thence
above the tympanum to a point above the insertion of the arm; in 17
per cent the stripe extends to the mid-flank, whereas in 13 per cent
the stripe is absent. A narrow and faint white line is present on the
canthus in some specimens, but no distinct white stripe is present
above the lateral dark line posterior to the eye. An interorbital bar
and transverse marks on the shanks are invariably present. The dorsal
markings are variable, but in most specimens (92%) consist of either
an X- or )(-shaped mark in the scapular region; in the other specimens
the markings are irregular short lines or absent. Approximately equal
numbers of specimens have a transverse bar, chevron, or broken lines
in the sacral region, whereas about eight per cent of the specimens
lack markings in the sacral region.
When active at night, individuals are pale yellowish tan with faint
brown dorsal markings. By day they are tan with more distinct brown
markings (Pl. 14). The thighs are pale yellow; the belly is white. The
iris is pale creamy tan with brown flecks. In breeding males the vocal
sac is yellow.
_Tadpoles._--Tadpoles of this species have been found in an extensive
grassy pond at Puerto Viejo, Costa Rica. The following description is
based on KU 104099, a specimen in development stage 36 (Gosner, 1960).
Total length, 21.0 mm.; body length, 6.7 mm.; body slightly wider than
deep, snout pointed; nostrils large, directed anteriorly, situated
near end of snout; eyes small, situated dorsolaterally, directed
laterally; spiracle sinistral, located just posteroventral to eye;
anal tube dextral. Tail xiphicercal; caudal musculature moderately
deep, extending far beyond posterior edge of fins; fins deepest at
about midlength; dorsal fin extending onto body, slightly deeper than
caudal musculature; ventral fin slightly shallower than musculature.
Mouth small, terminal, lacking teeth and fringing papillae, but having
finely serrate beaks. In preservative top of head olive-tan with brown
flecks; dark stripe from snout through eye to posterior edge of body;
belly white, flecked with brown anteriorly; tail creamy tan with
grayish brown blotches. In life, dorsum of body reddish tan mottled
with darker brown; lateral stripe dark brown; belly white, mottled
with brown and black; caudal musculature heavily pigmented with
grayish tan; posterior tip of tail marked with dark gray; caudal fins
heavily blotched with grayish tan; iris orange-tan peripherally, red
centrally (Pl. 15).
_Remarks._--This species has been confused with _Hyla microcephala
underwoodi_ by many workers. Dunn (1931, 1933, 1934) and Breder (1946)
referred Panamanian specimens of _H. phlebodes_ to _H. underwoodi_;
likewise, Gaige, Hartweg, and Stuart (1937) made the same error. Cole
and Barbour (1906) and Kellog (1932) used the name _H. phlebodes_ for
Mexican specimens of _H. microcephala underwoodi_. The similarity in
color pattern of _H. microcephala underwoodi_ and _H. phlebodes_
easily accounts for the misapplication of names. Although both species
have nearly identical dorsal color patterns, that of _H. microcephala
underwoodi_ usually is bolder. Furthermore, in that species a narrow
white line usually is present above the well-defined lateral dark
stripe, whereas the lateral dark stripe is short and posterior to the
eye is not bordered above by a white line in _H. phlebodes_.
The type locality "San Carlos, Costa Rica" given by Stejneger
(1906:817) apparently refers to a region, the Llanuras de San Carlos,
in the northern part of Alajuela Province, Costa Rica.
[Illustration: Fig. 3. Map showing locality records for
_Hyla phlebodes_.]
_Distribution._--_Hyla phlebodes_ inhabits humid tropical forests from
southeastern Nicaragua southeastward on the Caribbean slopes and
lowlands to the Canal Zone in Panama, thence eastward in the Chucunaque
Basin of eastern Panama and onto the Pacific lowlands of Colombia
(Fig. 3). The species also reaches the Pacific slopes in the Arenal
Depression in northwestern Costa Rica and in the Panamanian isthmus,
where it occurs in humid forests on the Pacific slope of El Valle and
Cerro La Campana. Mostly the species is found at low elevations, but
it occurs at 600 meters at Turrialba and at 700 meters at Finca San
Bosco in Costa Rica.
_Specimens examined._--410, as follows: +Nicaragua+: Zelaya: Isla
Grande del Maiz, MCZ 14848; Rio Mico, El Recrero, UMMZ 79720 (6).
+Costa Rica+: Alajuela: 12.4 km. N Florencia, MVZ 76108-10, USC 2628;
*Las Playuelas, 11 km. S Los Chiles, USC 7216; Los Chiles, USC 7217,
7219; 3 km. NE Muelle de Arenal, USC 2644 (2); *"San Carlos," USNM
29970. Cartago: Chitaria, KU 103690; *1.6 km. E Rio Reventazon Bridge,
east of Turrialba, UMMZ 119978 (2); *Tunnel Camp, near Peralta,
KU 32456, 32458-69, 41098 (skeleton); Turrialba, FMNH 101794, 103188-9,
KU 25725-9, 32439-48, 41095-7 (skeletons), 64797-827, 68300-2
(skeletons), 68403 (eggs), 68404 (tadpoles), MCZ 29224-5, 29310-2,
UMMZ 119979 (6), USC 31, 256 (2), 458 (2), 580, 594, 599 (7), 7074 (2),
USNM 29933. Guanacaste: Arenal, USC 6254; *Finca San Bosco, USC 62724,
6276 (3), Guayabo de Bagaces, USC 7022 (3), 7023; *Laguna Arenal,
USC 6262 (4); 3 km. NE Tilaran, USC 524; *5 km. NE Tilaran, USC 6269;
*6 km. NE Tilaran, UMMZ 122653 (6), S-2680 (skeleton), USC 523 (8).
Heredia: Puerto Viejo, KU 64828-63, 68303-7 (skeletons), 68405-6
(tadpoles), 104099-100 (tadpoles); *1.5 km. N Puerto Viejo, KU 64871;
*1 km. S Puerto Viejo, KU 86432-40; *4.2 km. W Puerto Viejo, KU
64864-5; *5.9 km. W Puerto Viejo, KU 64866-70; *7.5 km. W Puerto Viejo,
KU 86431. Limon: Batan, UMMZ 119980 (2); La Castilla, ANSP 23707;
Puerto +Limon+, KU 32449-55.
+Panama+: Bocas del Toro: 3.2 km. NW Almirante, KU 96026; Cayo de
Agua, KU 96027-31; Fish Creek, KU 96032-4. Canal Zone: Barro Colorado
Island, AMNH 69790, ANSP 23244-50; FMNH 13380, 22972-4; Juan Mina,
AMNH 55429, UU 3899; *8.6-13.8 km. N Miraflores Locks, TNHC 23439,
23477, 23484-8, 23491, 23494-9, 23501-2, 23504-8, 23510-17, 23519-30,
23532-8, 23541-54, 23561. *Rio Chagres, AMNH 55431-4; Rio Cocoli,
3.5 km. N Miraflores Locks, TNHC 23461, 23489-90, 23493, 23500, 23503,
23509, 23518, 23531, 23539-40; *Summit, ANSP 23361, KU 97788; *Three
Rivers Plantation, SU 2130. Cocle: El Valle de Anton, AMNH 55435,
69786-9, ANSP 23506-9. Colon: Achiote, KU 77215-78; Ciricito, CAS
71499-500, 71505-6. Darien: Rio Canclon at Rio Chucunaque, UMMZ 126733;
Rio Chucunaque, near Yavisa, AMNH 51783. Panama: Cero La Campana, FMNH
67847-50.
+Colombia+: Choco: Andagoya, FMNH 81856; Boca de Raspadura, AMNH
13570-8.
+Hyla sartori+ Smith
_Hyla underwoodi_ (in part), Smith and Taylor, Bull. U. S. Natl.
Mus., 194:85, June 17, 1948.
_Hyla microcephala sartori_ Smith, Herpetologica, 7:186,
December 31, 1951 [Holotype.--UIMNH 20934 from 1 mile north of
Organos, south of El Treinte, Guerrero, Mexico; H. M. Smith and
E. H. Taylor collectors]. Duellman, Univ. Kansas Publ., Mus. Nat.
Hist., 15:124, December 20, 1961. Porter, Herpetologica, 18:168,
October 17, 1962. Davis and Dixon, Herpetologica, 20:230,
January 25, 1965. Duellman, Univ. Kansas Publ. Mus. Nat. Hist.,
15:652, December 30, 1965.
_Diagnosis._--Dorsum tan with broad dark brown chevrons or transverse
bars; shanks marked with two or three broad transverse bars;
dorsolateral stripes absent.
_Description and variation._--No noticeable geographic variation is
apparent in either structural features or coloration in this species.
All specimens lack a dorsolateral dark stripe and white line, although
a dark line is present on the canthus and dissipates in the loreal
region. A broad interorbital brown bar is present in all specimens.
The color pattern on the dorsum invariably consists of a broad, dark,
chevron-shaped mark in the scapular region and a broad dark chevron or
transverse bar in the sacral region. The shanks invariably have two or
three dark brown transverse bars.
When active at night individuals are yellowish tan above with chocolate
brown markings (Pl. 14). The belly is white, and the thighs are pale
yellowish tan. The iris is dark bronze-color. In breeding males the
vocal sac is yellow. By day some individuals were observed to change
to creamy gray with distinct darker markings.
_Remarks._--Although tadpoles of this species have not been found,
observations on the breeding sites indicate that the tadpoles probably
develop in ponds. Except for calling males observed around a pool in a
stream-bed 11.8 kilometers west-northwest of Tierra Colorada, Guerrero,
all breeding congregations have been found at temporary ponds.
Smith (1951:186) named _Hyla sartori_ as a subspecies of _Hyla
microcephala_. This subspecific relationship seemed reasonable until
analysis of the mating calls showed that the call of _H. sartori_ is
more nearly like that of _H. phlebodes_ than that of _H. microcephala_.
The broad hiatus separating the ranges of _H. microcephala_ and _H.
sartori_ is additional evidence for considering _H. sartori_ as a
distinct species.
[Illustration: Fig. 4. Map showing locality records for
_Hyla sartori_.]
_Distribution._--_Hyla sartori_ occurs in mesophytic forests to
elevations of about 300 meters on the Pacific slopes of southern Mexico
from southwestern Jalisco to south-central Oaxaca (Fig. 4). The lack of
specimens from Colima and Michoacan probably reflects inadequate
collecting instead of the absence of the species there. On the basis of
available habitat the species would be expected to occur in Nayarit,
but extensive collecting there has failed to reveal its presence. The
semi-arid Plains of Tehuantepec apparently limit the distribution to
the east.
_Specimens examined._--190, as follows: +Mexico+: Guerrero: 5 km. E
Acapulco, AMNH 54611-2; 23.2 km. N Acapulco, UIMNH 26404-7; Colonia
Buenas Aires, 23 km. E Tecpan de Galeana, UMMZ 119223 (7); *El
Limoncito, FMNH 75785, 100390-402, 104631, 104633, UMMZ 117250, USNM
134266; El Treinte, FMNH 100403, UIMNH 20935-7; Laguna Coyuca, AMNH
59686; La Venta, MCZ 29635; *Morjonares, UIMNH 26392-402; 1.6 km.
N Organos, FMNH 100404-5, UIMNH 20933-4; 19.2 km. S Petaquillas,
UIMNH 26408; 6.1 km. E. Tecpan de Galeana, TNHC 23396-408; *11.2 km.
N Tierra Colorada, UIMNH 26403; 11.8 km. WNW Tierra Colorada, UMMZ
119225 (51), S-2677-9 (skeletons); Zacualpan, UMMZ 119224 (6). Jalisco:
6.4 km. NE La Resolana, KU 67853-69; 24 km NE La Resolana, KU 67870-3.
Oaxaca: 3 km. N Pochutla, KU 57539; 13.4 km. N Pochutla, UMMZ
123495 (40).