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Lynch.1982. Two New Species of Poison Dart Frogs - Edwardsi&ruizi
Lynch.1982. Two New Species of Poison Dart Frogs - Edwardsi&ruizi
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ABSTRACT: Two dendrobatids, provisionally assigned to Colostethus, are named from the
Cordillera Oriental of Colombia near Bogota, Colombia. Both species differ from other Colo-
stethus in lacking the vocal slits and in having elongate anal sheaths. One species is cavemic-
olous and occurs at elevations above 3000 m (caves within paramos).The other species occurs
along mountain streams in areas once supportinghigh altitude cloud forests (elevations ca. 2400-
2800 m).
Key words: Amphibia; Salientia; Dendrobatidae; Colostethus; Colombia; High altitude;
Troglodytic; Relationships
FIG. 1.-Anal sheath in Colostethus ruizi (a) posterior view, (b) lateral view, semi-diagrammatic,ICN
5419. Line equals 5 mm.
turales. The material from one site (Las support their generic separation from
Moyas caves) was obtained incidentally Colostethus, I am describing each as
to surveys on bats. The occurrence of frogs Colostethus pending systematic revision
in caves is uncommon and thus in 1980, of that paraphyletic genus.
with Dr. Alberto Cadena and Sr. Juan M. The following abbreviations are used
Renjifo, I collected adults, young and in the paper: E-N, eye to nostril distance;
tadpoles in the caves in the paramo HW, head width; IOD, interorbital dis-
northeast of Bogota. tance; SVL, snout-vent length; chord of
Two species of frogs are included in head length is the distance from the tip
the available samples. One occurs in of the snout to the angle of the jaw. All
caves in the p'aramos found on the chain specimens are deposited in the amphib-
of mountains immediately west of the ian collection of the Instituto de Ciencias
capital of Colombia. The other species is Naturales (ICN), Universidad Nacional de
known from streams near the Alto de San Colombia, Bogota.
Miguel on the old road from Bogota to
Fusagasuga (SE of Bogota). The two frogs Colostethus edwardsi sp. nov.
share a unique character-state (anal open- Holotype.-ICN 6376, an adult female
ing extended to mid-level of lower-level obtained from Cueva de Las Moyas, 0.7
of thighs by an anal sheath, Fig. 1), a pos- km SW Los Patios, vereda Las Mollas (=
sibly paedomorphic trait (absence of vo- Moyas?), municipio La Calera, Departa-
cal slits and sac), and a variety of unre- mento Cundinamarca, Colombia, 3030 m,
solved character-states (drab color pattern on 16 August 1980 by John D. Lynch and
lacking stripes; narrow digital pads with Juan Manuel Renjifo.
poorly developed scutes; basal toe web- Paratypes.-ICN 6377-90, 8561, 8563,
bing; smooth to feebly warty skin on the topotypes; ICN 8560, p'aramo Cruz Verde,
lower back; lack inner tarsal fold and tu- carretera a Choachi, municipio de
bercle; relatively large, metamorphosis at Choachi, Depto. Cundinamarca, Colom-
17-22 mm SVL, adult females 32.0-37.3 bia.
mm SVL). Referred specimens. ICN 2770, para-
The presence of an anal sheath is suf- mo Cruz Verde, camino a Choachi; ICN
ficient to unite these two species as a dis- 6391 (two tadpoles), 8562, 8564, topo-
tinct group within the Dendrobatidae. types.
Although further study of frogs of this Diagnosis.-A dendrobatid distin-
group and Colostethus will probably guished from all others except C. ruizi by
a 0o 7
FIG. 2. Plantarviews of left hind feet of (a) Colostethus edwardsi, ICN 6376, and (b) Colostethus ruizi,
ICN 5418. Line equals 2 mm.
the possession of an anal sheath opening round to ovoid in dorsal view, rounded
at mid-level of thighs and distinguished in lateral profile, nostrils small, not pro-
from C. ruizi in having a brown venter tuberant, directed dorsolaterally with
spotted with white, in having more ex- slight anterior vector; canthus rostralis
tensive toe webbing (Fig. 2), a shorter rounded, scarcely evident; loreal region
snout, and in having a nearly completely weakly concave, sloping abruptly to lips;
concealed tympanum. lips not flared; nostrils essentially at tip
Description.-Head narrower than to of snout; snout very short (Fig. 3); inter-
as wide as body, wider than long; snout orbital space flat, broader than width of
FIG 3. Profiles of heads of (a) Colostethus edwardsi, ICN 6376, and (b) Colostethus ruizi, ICN 5417.
Line equals 2 mm.
FIG.4.-Colostethus edwardsi, (left) ICN 6376, 37.3 mm SVL, (right) ICN 6377, 27.5 mm SVL.
In the smaller individuals, the ventral two adult females, and one juvenile fe-
surfaces are cream with brown spots; in male (ICN 6377, 27.5 mm SVL).
slightly larger frogs, cream with brown Larvae.-(based on ICN 6391, a lot of
reticulation, and in the adults, brown with two free-swimming tadpoles, in devel-
cream spots. Concealed limb surfaces opment stage 25; Gosner, 1960). Body
mottled cream and brown (or brown and lengths 22.5 and 19.8 mm, total lengths
darker brown). 51.3 and 47.4 mm; body depth three-
In life, C. edwardsi is medium brown fourths greatest body width, deepest and
with dark brown spots above; largest in- broadest posteriorly; in dorsal view, body
dividuals have hints of yellowish dorso- an elongate ellipse with a rounded snout;
lateral stripes from eye to above arm; lips in profile snout round, nostrils directed
spotted white or pale yellow; hidden sur- anterolaterally, about midway between
faces of limbs dark olive to pale brown eyes and tip of snout; eyes small, dorso-
with brown spots; venter olive to brown lateral, broadly separated; spiracle sinis-
with yellow spots (throat more pale); tral; cloacal tube short, opening dextral;
slight orange wash on throat and under- caudal musculature deep, extending
sides of limbs; iris bright copper with nearly to tip of broadly rounded tail; cau-
black reticulations and black horizontal dal fins not extending onto body; dorsal
streak. and ventral fins about of equal depth, ex-
Measurements of holotype in mm.- cept on anterior 1/4of tail where dorsal fin
SVL 37.3, tibia 14.8, HW 12.3, head length is very low, each fin about 1/2 depth of
9.5, chord of head length 11.3, upper eye- caudal musculature at midlength of tail;
lid width 2.1, IOD 3.2, eye length 3.8, E- tail fins not tapering.
N 2.2. Mouth small, directed anteroventrally,
Variation.-Too few specimens are median 2/5 of upper lip lacking papillae;
available to assess differences in propor- rest of mouth bordered by two rows of
tions between sexes. Two adult males are small labial papillae; lips indented pos-
27.5-27.6 mm SVL, two gravid females terolaterally, beaks moderately robust,
are 30.8 and 37.3 mm SVL. Proportions bearing numerous sharp serrations; up-
are as follows: tibia/SVL 38.8-44.4 (x= per beak with long slender lateral pro-
40.9, n = 5) %; HW/SVL 33.0-34.4 (x = cesses; lower beak broadly V-shaped, two
33.4, n = 4) %; eyelid/IOD 60.7-87.5 (x = upper rows of denticles, inner row nar-
71.2, n = 4) %; E-N/eye 56.2-65.6 (x = rowly interrupted medially in larger tad-
60.4, n = 4) %; based on two adult males, pole, not interrupted in smaller individ-
ual; three lower rows of denticles, all Colostethus ruizi sp. nov.
complete, third lower row shorter than Holotype.-ICN 5419, an adult male
other two (Fig. 5). obtained "on the border of a stream,
In life, body brown above, throat and [along] the road to Fusagasuga, between
venter transparent, caudal musculature [Alto de] San Miguel and La Aguadita,"
pale olive or cream, heavily spotted with Departamento Cundinamarca, Colombia,
brown; caudal fins spotted with brown. on 1 December 1979 by J. Clavijo and A.
Ecological notes.-ICN 6376-91 were Fajardo.
captured in a cave within 30 m of its en- Paratypes.-ICN 5415-18, taken with
trance but all specimens encountered holotype; ICN 4838-39, Alto de San
were found in areas away from light. Ju- Miguel, Departamento Cundinamarca,
veniles were found beneath rocks in shal- Colombia, 2640 m.
low streams (2-3 cm deep) leading into Type-locality.-The collectors' notes
the larger stream (0.1-0.5 m deep) as well are imprecise ("carretera a Fusa, entre San
as swimming in the larger stream. The Miguel-La Aguadita"). Between the Alto
holotype was swimming in a pool 0.5 m de San Miguel and Aguadita, one finds
deep. Many tadpoles were seen in the three streams. One lies a few hundred
larger stream, especially in deeper pools meters above La Aguadita, a second lies
(0.3-0.5 m). Vladimir Corredor collected on the northeast edge of Aguadita. The
specimens on two occasions in the p'ara- third is the Quebrada Agua Bonita, 14.4
mo Cruz Verde. The frogs were first en- km (by road) SW Sibate, municipio Sil-
countered at the edges of fissures in the vania, 2410 m (between km 21-22, ca-
soil, but they attempted escape by hop- rretera Bogota-Fusagasuga), the first
ping into streams in the fissures. These stream below the Alto de San Miguel. At
fissures lead to caves too small to permit this stream, I saw Colostethus tadpoles'
entry of a collector. Reaching into the fis- in May and June 1981 but was unable to
sures resulted in captures of a juvenile catch examples. Quebrada Agua Bonita is
and an adult male. Although Sr. Corredor probably the source of the specimens ob-
pursued frogs seen at the surface, other tained by Clarijo and Fajardo.
frogs were seen with the aid of a head- Diagnosis.-Allied to C. edwardsi but
lamp deep in the fissures. Presumably the differing in having a dusky cream venter
frogs seen at the surface are conspecific without spots, less toe webbing (not en-
with those captured in the fissures. closing subarticular tubercle on second
The only other frogs found in Cueva de toe, reaching between subarticular tuber-
cles on third and fifth toes, see Fig. 2), a than second; thumbs of breeding males
longer snout (Fig. 3), and in having an not swollen, not bearing nuptial pads.
only partially concealed tympanum. No tubercles on knee, heel or tarsus;
Description.-Head narrower than to outer edge of tarsus lacking fold or tu-
as wide as body, wider than long; snout bercles; inner tarsal fold evident only as
ovoid in dorsal view, flat and rounded in slightly thickened area (no fold is pres-
lateral profile; nostrils small and slightly ent, at least not the sort of fold ending in
protuberant, directed dorsolaterally; can- a sickle-shaped tubercle typical of Colo-
thus rostralis rounded but evident, stethus); inner metatarsal tubercle low,
straight; loreal region weakly concave, twice as long as wide, three times size of
sloping abruptly to lips; lips not flared; rounded outer metatarsaltubercle; no su-
nostrils nearer to tip of snout than to eye; pernumerary plantar tubercles; subar-
snout short (Fig. 3); interorbitalspace flat, ticular tubercles round, low; digit tips
broader than width of upper eyelid; up- bearing slightly expanded pads on whose
per eyelids and skin of head bearing low dorsal surfaces one finds an ill-defined
warts, none enlarged; supratympanicfold pair of scutes (not separated by groove);
obscure, greatly thickened, obscuring lateral margins of toes bearing fringes
posterodorsal one-third of tympanum; which coalesce at bases of toes forming
tympanum round, directed anterolateral- basal webbing (Fig. 2); webbing formu-
ly (with slight posterior vector), separat- lae (following Savage and Heyer, 1967) I
ed from eye by distance equal to its 1-1+ II 1--2- III 12/3-22/3 IV 23/4- 11/2
length; postrictal tubercles not evident; V, webbing incised; no outer metatarsal
choanae large, longer than wide (tear- fold; when hind legs are flexed and held
drop-shaped), not concealed by palatal at right angles to sagittal plane, heels do
shelf of maxillary arch; no vomerine not touch.
odontophores or teeth; tongue twice as Dark brown above flecked with even
long as wide, its posterior border feebly darker brown or black; slightly paler
notched, posterior two-fifths not adherent stripe from eye to above insertion of up-
to floor of mouth; males lack vocal slits per arm (remnant of dorsolateral stripe?);
and vocal sac; premaxillae and maxillae similar stripes around snout; a few pale
bearing numerous medially curved spots along lower lip to base of arm;throat
pointed teeth. brown; venter (including undersides of
Skin of dorsum and upper surfaces of arms and legs) dirty cream; undersides of
limbs finely shagreened; round warts su- hands and feet, concealed surfaces of
perimposed on shagreened dorsum on limbs, lateral margins of forearms, and
lower back of adult male holotype but not posterior margins of undersides of thighs
on adult females or other adult male; no dark brown.
folds on dorsum; skin of ventral surfaces Measurements of holotype in mm.-
smooth; anal opening extended in a SVL 35.2, tibia 14.6, HW 11.7, head length
sheath opening at mid-level of thighs; no 9.3, chord of head length 11.2, upper eye-
enlarged warts in vicinity of anal open- lid width 2.5, IOD 2.9, tympanum length
ing; forearm robust, lacking ulnar fold or 1.1, eye length 3.8, E-N 2.4.
tubercles; palmar tubercle round, twice Variation.-Both males lack obvious
size of oval thenar; no supernumerary patterns. They have vague partial dorso-
palmar tubercles; subarticular tubercles lateral stripes (anterior 1/3 of body), fleck-
round, not pungent; sharp lateral keels ing or mottling on the dorsum, darker
on fingers but no fringes; third finger of flanks than dorsum and no limb bands.
male not swollen; tips of digits swollen ICN 4838 and 5418 are yellow-brown
forming rounded pads, pads only slightly with small brown spots. Both have limb
wider than digit; tops of digital pads bars (oblique, as wide as interspaces on
bearing obscure scutes; first finger longer shanks). Both have short dorsolateral
stripes. The other two females (5416-17) and is thus subject to cautious interpre-
are colored like the males except no dor- tation. The absence of the subgular vocal
solateral stripes are apparent. The ventral sac and vocal slits is unique with Colo-
surfaces of ICN 4838 are pale. The throat stethus and nearly so within the Dendro-
is dusted with brown contrasting with the batidae, but Myers (1982:5) recently
cream venter. ICN 4839 is a recently reported the absence of vocal slits in
metamorphosed young (17.2 mm SVL). Dendrobates reticulatus and variable
The dark flecking of the dorsum is only presence in D. quinquevittatus. No fea-
feebly indicated and there is a pale retic- ture is known that unites the 60 plus
ulation across the breast. species now placed in Colostethus, and
Too few specimens are available to as- ratherthan encumber future research with
sess differences in proportions between a possibly needless name, I am assigning
sexes. Two adult males are 34.4 and 35.2 the two species to Colostethus.
mm SVL, three adult females are 32.0, Colostethus edwardsi and C. ruizi are
34.8 and 38.5 mm SVL. Proportions are more similar to one another than either
as follows: tibia/SVL 37.9-46.2 (i = 41.4, is to any other dendrobatid species
n = 6) %; HW/SVL 32.2-35.2 (x = 33.5, known. This observation is insufficient to
n = 6) %; eyelid/IOD 62.5-86.2 (i = 73.4, support a hypothesis that the two phe-
n = 3) %; tympanum/eye 28.9-36.8 (x = notypically distinct forms are subspecies
33.2, n = 6) %; E-N/eye 54.8/69.7 (x = of a single species. Because interpopu-
62.3, n = 6) %;based on two adult males, lational variation is not bridged by intra-
three adult females, and one juvenile fe- populational variation, the geographical-
male (ICN 5418, 29.8 mm SVL). ly and ecologically isolated populations
Ecological notes.-ICN 5415-19 were are most parsimoniously represented by
taken in December beneath rocks along a hypothesis that these are distinct
a stream.The two adult females are spent. species. Extensive collecting in the Cor-
ICN 4838, a gravid female, was obtained dillera Oriental is necessary in order to
in March 1978. accumulate sufficient evidence to dis-
Etymology.-Named for my friend and criminate between the possible hypoth-
colleague, Pedro Ruiz, who found the first eses.
specimens.
Other frogs found at the QuebradaAgua
LITERATURE CITED
Bonita and/or vicinity of Alto de San
Miguel include Atelopus subornatus, EDWARDS, S. R. 1974. A Phenetic Analysis of the
Colostethus sp., Eleutherodactylus bo- Genus Colostethus (Anura:Dendrobatidae).Ph.D.
Dissertation, University of Kansas.
gotensis, E. sp. and Hyla bogotensis. GOSNER, K. L. 1960. A simplified table for staging
Distribution.-Known only from the anuran embryos and larvae with notes on iden-
vicinity of the Alto de San Miguel (ap- tification. Herpetologica 16:183-190.
prox. 4?25' N, 74?18' W), in cloud forests LYNCH, J. D. 1971. Evolutionary relationships,
osteology, and zoogeography of leptodactyloid
on the western slopes of the Cordillera frogs. Univ. Kansas Mus. Nat. Hist. Misc. Publ.
Oriental, Departamento Cundinamarca, 53:1-238.
Colombia, 2410-2469 m. MYERS, C. W. 1982. Spotted poison frogs: de-
scriptions of three new Dendrobates from west-
DISCUSSION ern Amazonia, and resurrection of a lost species
from "Chiriqui." Amer. Mus. Novitates 2721:1-
Colostethus edwardsi and C. ruizi are 23.
peculiar within the family Dendrobati- MYERS, C. W., J. W. DALY, AND B. MALKIN. 1978.
dae for at least two reasons: no other A dangerously toxic new frog (Phyllobates) used
member of the family has an anal sheath by Embera indians of western Colombia with dis-
and all other dendrobatids have inner tar- cussion of blowgun fabrication and dart poison-
ing. Amer. Mus. Nat. Hist. Bull. 161:307-366.
sal folds and/or tubercles-one of which NOBLE, G. K. 1931. The Biology of the Amphibia.
refers to the absence of a characteristic McGraw-Hill Book Co., New York.
ABSTRACT: Ptychohyla panchoi is named from the upper tropical moist forest on the Sierra
de las Minas in Guatemala. A phylogenetic arrangement of the species of Ptychohyla shows P.
panchoi to be intermediate between species in the P. euthysanota group and the P. schmidtorum
group.
Key words: Amphibia; Anura; Hylidae; Ptychohyla panchoi sp. nov.; Guatemala