Occasional papers of the Museum of Natural History, the University of Kansas

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

MCZ
OCCASIONAL PAPERS LIBRARY

of the APR ?.3 1990

MUSEUM OF NATURAL HISTORY
The University of Kansas ^l^^^xv
Lawrence, Kansas

NUMBER 134, PAGES 1-23 MARCH 8, 1990

SYSTEMATICS OF FROGS OF THE
HYLA LARINOPYGION GROUP

By

William E. Duellman' and David M. Hillis-

Stream-breeding hylid frogs of the genera Hyla. Plectrohyla, and
Ptychohyla are abundant and rich in species in the highlands of Mexico and
Central America (Duellman, 1970). In contrast, only Hyla is represented in
the Andes, and until recently comparatively few species have been known.
These include four species in southern Peru and Bolivia (//. armata, balzani,
callipleiira, and melanopleura) and 12 species in the Hyla hogotensis group
ranging from Venezuela to Peru, plus one species in Costa Rica and Panama
(Duellman, 1972, 1989). The other stream-breeding frogs in the Andes all
seem to represent a monophyletic group that has been referred to as the Hyla
larinopygion group.

The first of these frogs was discovered in 197 1 in the Cordillera Central in
Colombia and was named as Hyla larinopygion (Duellman, 1973). Another
species, Hyla lindae. was named from the Cordillera Oriental in Ecuador
(Duellman and Altig, 1 978). Twoother species were discovered subsequently —
Hyla pantosticta from the Cordillera Oriental in southern Colombia and
Ecuador (Duellman and Berger, 1982) and Hyla sarampiona from the
Cordillera Occidental of Colombia (Ruiz and Lynch, 1982). With the excep-
tion of H. larinopygion in the Cordillera Central in Colombia, frogs in this
group are poorly represented in collections. Perhaps this is because of their
proclivity to inhabit cliffs in the vicinity of waterfalls, where, even if males

'Curator, Division of Herpetology, Museum of Natural History, Dyche Hall, and
Professor, Department of Systematics and Ecology, Haworth Hall, The University of
Kansas, Lawrence, Kansas 66045-2454

■^Associate Professor, Department of Zoology. The Universitv of Texas. Austin,
Texas 78712-1064

2 OCCASIONAL PAPERS

are calling, their voices do not carry far, and the frogs are difficult to locate.
Our field studies in the Andes of Ecuador have revealed the existence of
three more unnamed species in \htHyla larinopygion group; furthermore, we
obtained frozen tissues of five species in the group. This material, plus a
paratype of a new species from the Cordillera Central of Colombia being
named by Pedro M. Ruiz, constitutes the basis for this paper, in which we (1)
define the Hyla larinopygion group, (2) describe three new species, (3)
provide a key to the species in the group, and (4) present an hypothesis of the
phylogenetic relationships among the species.

MATERIALS AND METHODS

The museum specimens examined are listed with the standardized acro-
nyms (Levi ton et al., 1985). Measurements and definitions of external
features and osteological characters follow Duellman (1970). Tadpoles were
staged according to Gosner (1960).

Tissues were collected for allozymic analysis from five species oHhcHyla
larinopygion group. In addition to those specimens listed in Appendix 1,
tissues were obtained from one specimen of//, larinopygion (Juan M. Renjifo
field number JMR 2072 from Finca La Carelia, Municipio de Salento,
Departamento de Quindio, Colombia; specimen to be deposited in collection
of INDERENA, Bogota, Colombia). Because outgroup relationships of the
Hyla larinopygion group are poorly understood, we selected representatives
from 11 Central and South American species groups of Hyla for use as
outgroups (Table 1).

Tissue samples (skeletal muscle and Uvers) were removed from freshly
killed frogs and frozen immediately in liquid nitrogen for transport to the
laboratory. Tissues were stored for up to three years at -80°C. Tissues were
homogenized in ground-glass grinding tubes and diluted 1 : 1 with 0.01 M tris-
0.001 M EDTA-0.001 M 2-mercaptoethanol, pH 7.5. Homogenates were
centrifuged at 7000 g for 3 min, and the supematants were refrozen at -80°C
prior to use.

Procedures of starch-gel electrophoresis, including buffer recipes and his-
tochemical staining protocols, were the same as described by Hillis (1985).
Eighteen genetic loci were scored among species (Table 2). Corrected Nei's
genetic distances were calculated as described by Hillis (1984). AUPGMA
phenogram of genetic distances was calculated (Sneath and Sokal, 1973) in
order to assess average genetic divergence among the species.

The most parsimonious cladogram was constructed from the allozymic
data using the procedure described by Hillis (1985). Specifically, any electro-
morphs that were found in the ingroup and in any of the outgroups were
considered to be symplesiomorphies. Electromorphs were ordered into

SYSTEMATICS OF HYLA LARINOPYGION GROUP

Table 1. Outgroups, collection localities, and informative loci.

Species

Species Group

Provenance

Informative Loci

H. alytolylax

bogotensis

Ecuador

EST, LDH-1, LDH-2,
MDH-2

H. carnifex

Columbiana

Ecuador

GPI

H. granosa

granosa

Peru

EST, PEPS, PGM

H. labialis

labialis

Colombia

AAT-2, IDH, MDH-2, ME

H. leali

minima

Peru

EST, GPI, IDH, PEP-A

H. leucophyllata

leucophyllata

Peru

AAT-2, IDH, ME, PEP-A

H. parviceps

parviceps

Peru

AAT-2, GPI, MDH-1,
PEP-A, PEP-S, PGM

H. pentheter

bistincta

Mexico

MDH-1, ME, PEP-A,
PEPS, PGM

H. picturata

geographica

Ecuador

IDH, MDH-1, PEP-A,
PEPS

H. rhodopepla

microcephala

Peru

AAT-2, GPI, IDH, PEP-A

H. rosenbergi

boans

Ecuador

AAT-2, IDH, ME, PEP-A,
PEP-S

transformation series following the taxonomic outgroup and functional
outgroup criteria of Watrous and Wheeler (1981), as expanded by Farris
(1982), Derived electromorphs were considered to be synapomorphies even
if they were not fixed within the lineage in which they first appeared.

TAXONOMY

Frogs of the Hyla larinopygion group are characterized by (1) large size
(snout-vent length to 77.3 mm); (2) large hands and feet with large terminal
discs and little webbing; (3) skull well ossified with extensively ossified
sphenethmoid and small frontoparietal fontanelle (Fig. 1 A); (4) quadratoju-
gal present; (5) vomerine odontophores long, transverse, posterior to cho-
anae, abutting medially (Fig. 1 B); (6) prepoUical tubercle supported by broad,
elliptical, ossified prepoUex (Fig. IC); (7) tadpoles large (up to 80 mm total
length) with long, muscular tails; (8) larval mouth ventral with one or two
complete rows of marginal papillae and as many as eight upper and ten lower
rows of denticles.

The known members of the Hyla larinopygion group occur along high-
gradient streams in cloud forest at elevations of 1900-2700 m on the slopes

OCCASIONAL PAPERS

Table 2. Loci examined, Enzyme Commission Numbers, and Tissues
and Buffer Systems used.

Locus

Abbrev.

E.G.

No.

Tissue

Buffer
System

Aspartate aminotransferase- 1

AAT-1

2.6.1.1

muscle

Poulik

Aspartate aminotransferase-2

AAT-2

2.6.1.1

muscle

Poulik

Creatine kinase

CK

2.7.3.2

muscle

TC6.7

Esterase

EST

3.1.1.1

liver

Poulik

Glucose-6-phosphate isomerase

GPI

5.3.1.9

muscle

Poulik

Gly ceraldehy de-3 -phosphate

dehydrogenase

G-3-PD

1.2.1.12

liver

TC6.7

Glycerol -3 -phosphate

dehydrogenase (NAD+)

GPD

1.1.1.8

liver

TC6.7

Isocitrate dehydrogenase

IDH

1.1.1.42

muscle

TC6.7

Lactate dehydrogenase- 1

LDH-1

1.1.1.27

liver

TC6.7

Lactate dehydrogenase-!

LDH-2

1.1.1.27

liver

TC6.7

Malate dehydrogenase- 1

MDH-1

1.1.1.37

liver

TC6.7

Malate dehydrogenase-2

MDH-2

1.1.1.37

liver

TC6.7

Malate dehydrogenase

(NADP-H) ("malic enzyme")

ME

1.1.1.40

muscle

Poulik

Peptidase-A

PEP-A

3.4.11.13

liver

Poulik

Peptidase-B

PEP-B

3.4.11.13

liver

Poulik

Peptidase- S

PEPS

3.4.11.13

liver

Poulik

Phosphoglucomutase

PGM

5.4.2.2

muscle

Poulik

Triose-phosphate isomerase

TPI

5.3.1.1

muscle

TC6.7

of the Andes from southern Colombia to central Ecuador (Fig. 2). In addition,
H. larinopygion occurs at elevations of 2200-3300 m in the Cordillera
Central of Colombia north to the Departamento de Antioquia (J.D. Lynch,
pers. comm.).

Hyla pacha new species

Fig. 3

Holotype.— KU 202762, an adult male, from 11.2 km (by road) west-
southwest of Plan de Milagro (03°03'S, 78^08^), 2350 m, Provincia de
Morona-Santiago, Ecuador, one of a series collected on 12 March 1984 by
William E. Duellman and David M. Hillis.

SYSTEMATICS OF HYLA LARINOPYGION GROUP 5

Paratypes.— KU 202760-61 collected with the holotype; KU 202763
from 12.4 km (by road) west-southwest of Plan de Milagro, 2225 m, Provincia
de Morona-Santiago, Ecuador, obtained on 12 March 1984 by David M.
Hillis.

Diagnosis. — This large species (females to 66.5 mm snout-vent length)
differs from other members of the group by having a dark brown dorsum with
orange flecks (cream in preservative) and bold black and cream mottling on
the venter. Hyla pacha differs further from all species in the group, except H.
larinopygion and H. sp. A, by having small, triangular calcars. Hyla larino-
pygion has a uniformly brown dorsum and black and blue bars on the flanks,
whereas in H. sp. A the dorsum is brown with narrow, transverse black lines
and the flanks are brown with large cream spots. The only other species in the
group having orange on the dorsum are H. pantosticta and H. sarampiona.
The former has many distinct, small, round, orange spots on all dorsal
surfaces and the flanks and is unique in having areolate skin on the dorsum.
Hyla sarampiona has a pale olive dorsum with small orange spots; the venter
is black.

Description. — Body robust; head about as broad as long, as wide as body;
head width 31.6-33.1 (3c=32.3, n=4) percent of snout- vent length; head
length 32.1-34.3 (3c =32.9, n=4) percent of snout- vent length; length of snout
greater than diameter of eye; snout in dorsal view acutely rounded, in profile
bluntly rounded; canthus round; loreal region concave; lips not flared; nares
slightly protuberant, directed anterolaterally at point posterior to anterior

5 mm

Fig. 1. Osteology of Hyla lindae, KU 155477, female, 64.5 mm SVL: A. Dorsal view
of skull; B. Ventral view of anterior part of skull; C. Palmar view of right carpus.

OCCASIONAL PAPERS

79 78 77 76

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'=■ » ',V--^ ', >'i--

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▼ H pantosticia

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■*■ H ptychodactyla

♦ H sarampiona
+ W species A

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.(.'■;,^ ',^ ;.•.••-.:• 1500m

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79 78 77 76

Fig. 2. Distribution of species in the Hyla larinopygion group in Ecuador and southern
Colombia.

SYSTEMATICS OF HYLA LARINOPYGION GROUP

Fig. 3. Holotype oiHyla pacha. KU 202762, male. 58.0 mm SVL.

margin of lower jaw; intemarial region barely depressed; top of head flat;
interorbital distance noticeably wider than eyelid; eye moderately large,
prominent; tympanum vertically ovoid, separated from eye by distance
greater than twice length of tympanum; tympanic annulus distinct; supratym-
panic fold heavy, sigmoid, obscuringupper part of tympanum; post- tympanic
fold heavy beginning under supratympanic fold, extending to angle of jaw,
obscuring posterior edge of tympanum. Forearm robust; row of prominent
tubercles along ventrolateral edge of forearm; fingers long, bearing large,
round discs; width of disc on third finger twice length of tympanum; relative
lengths of fingers 1<2<4<3; fingers webbed basally; webbing formula for
outer fingers 112 — 3III3 — 3IV; subarticular tubercles large, round, none
bifid; supernumerary tubercles large, round, in two rows on proximal seg-
ments; palmar tubercle large, bifid; prepoUical tubercle large, elliptical;
nuptial excrescences absent. Hind limb moderately robust; tibia length
48.5-52.5 (^=50.2, n=4) percent of snout- vent length; foot length 47.5-50.3
(^=48.4, n=A) percent of snout- vent length; calcar present, small, triangular;
tarsal fold absent; inner metatarsal tubercle large, elliptical; outer metatarsal
tubercle moderately large, distinct, round; toes long, bearing discs only
slightly smaller than those on fingers; relative lengths of toes 1<2<3=5<4;

8 OCCASIONAL PAPERS

toes about one-half webbed; webbing formula 12" — l^UVA — IV-JBl' — 3'
IV21/2 — IViV; subarticular tubercles large, round; supernumerary tubercles
prominent, round, in two rows on proximal segments.

Skin on dorsum and flanks finely shagreened; skin on throat finely
granular; skin on chest, belly, and proximal posteroventral surfaces of thighs
coarsely granular; other surfaces smooth; anal opening directed posteroven-
trally at midlevel of thighs in males, posteriorly at upper level of thighs in
female; dermal supra-anal flap long, transverse in males, absent in female;
pair of vertically elongate tubercles below anal opening. Vomerine odonto-
phores long, transverse, abutting medially, behind level of large, ovoid
choanae, bearing 11-15 (^=13.3, /i=4) teeth; tongue broad, cordiform,
shallowly notched posteriorly, barely free behind; vocal slits along posterior
lingual margins of mandibles; vocal sac large, single, median, subgular.

Coloration in preservative: Dorsal surfaces of head, body, forearms, and
shanks black with scattered, minute cream flecks; flanks black with two or
three bold cream vertical or diagonal marks; all surfaces of thighs and upper
arms, inner surfaces of forearms and tarsi, and dorsal surfaces of hands and
feet black with cream bars; digits cream and black; all ventral surfaces black
with bold cream motthng (Fig. 4A).

Coloration in life: Dorsum dark brown with metallic orange flecks; venter,
flanks, and hidden surfaces of limbs brown and creamy white; iris olive-
brown.

Measurements: The measurements of three males (mm with means in
parentheses are followed by the measurements of one female: snout-vent
length 58.0-60.8 (59.2), 66.5; tibia length 29. 1-29.5 (29.3), 34.9; foot length
27.6-29.3 (28.3), 33.5; head width 18.9-19.9 (19.6), 21.5; head length
18.7-20.1 (19.3), 21.0; interorbital distance 5.9-6.5 (6.1), 6.2; intemarial
distance 4.9-5.4 (5.1), 5.4; eye-nostril distance 4.3^.9 (4.6), 4.8; diameter of
eye 5.5-6.0 (5.7), 6.0; diameter of tympanum 2.4-2.8 (2.6), 3.0.

Distribution and ecology. — This species is known from only two streams
at elevations of 2225 and 2350 m on the Amazonian slopes of the Cordillera
Oriental in central Ecuador. All individuals were active at night in cloud
forest. Males were calling from the ground adjacent to, a boulder in, and a tree
limb 2.5 m above, a cascading mountain stream. A female was on a frond of
a tree fern 10 m above the base of a waterfall. The call is a three-noted whistle
that is higher pitched than the three-noted call of H. lindae.

Etymology. — The specific epithet is a noun in apposition and is a
patyronym for Patricia (Pacha) A. Burrowes, who worked with us in the cloud
forests of Ecuador in 1984.

SYSTEMATICS OF HYLA LARINOPYGION GROUP

Fig. 4. Ventral color patterns in species in the Hyla larirwpygion group: A. H. pacha,
KU 202762; B. H. larinopygion. KU 144127; C. H. pantosticta. KU 190000; D. H.
species A, KU 189598; E. H. psarolaima, KU 164313. The ventral surfaces of//.
lindae, sarampiona, and ptychodactyla are unicolor, except that the latter has irregular
pale spots on the throat.

Hyla psarolaima new species
Fig. 5

Holotype. — KU 164313, an adult male, from 11 km (by road) east-
southeast of Papallacta ((X)°03'S, 78°08'W), 2660 m, Provincia de Napo,
Ecuador, obtained on 22 March 1975 by Linda Trueb.

Paratypes. — KU 164314-15, 164316 (skeleton) from the type locality,
collected on 23 March 1975 by William E. Duellman; KU 202727 from 26.2

10

OCCASIONAL PAPERS

km west-southwest of Plan de Milagro, 2640 m, Provincia de Morona-
Santiago, Ecuador, obtained on 12 March 1984 by William E. Duellman;
MCZ 1093 17-18 from between La Alegria and Sibundoy, 24 10 m, Provincia
de Napo, Ecuador, collected on 5 November 1984 by Robert Bleiweiss and
Juan Carlos Matheus.

Referred specimens. — Four juveniles, KU 169582-85 from 35 km east
of San Francisco, 1950 m, Departamento de Putumayo, Colombia, collected
on 26 September 1974 by William E. Duellman and Linda Trueb.

Diagnosis. — This species differs from all other members of the group by
having a creamy gray throat with brown and white flecks, gray venter with
brown and cream spots, grayish brown dorsum with small brown and minute
cream flecks, and a small tubercle on the heel. All other species in the group
have a uniformly black venter (//. lindae, sarampiona, and species A) or
boldly mottled black and cream or blue venters (//. larinopygion, pacha, and
pantosticta). Dermal appendages are absent on the heels of//, pantosticta and
sarampiona, whereas the other species in the group have small, triangular
calcars, instead of an elongate tubercle as in H. psarolaima.

Fig. 5. Paratype of Hyla psarolaima, KU 164314, male, 55.4 mm SVL.

SYSTEM ATICS OF HYLA LARINOPYGION GROUP 11

Description. — Body robust; head slightly longer than wide, as wide as
body; head width 3 1.2-34 .4 (3c=32.2,n=7) percent of snout- vent length; head
length 3 1 .9-34.4 (3c =32.7, n= 7) percent of snout- vent length; length of snout
greater than diameter of eye; snout in dorsal view acutely rounded, in profile
bluntly rounded, nearly truncate; canthus round; loreal region concave; lips
not flared; nares slightly protuberant laterally, directed anterolaterally at point
just posterior to anterior margin of lower jaw; intemarial area barely de-
pressed; top of head flat; interorbital distance greater than width of eyelid; eye
moderately large, prominent; tympanum broadly ovoid, higher than long,
separated from eye by distance 1.5 times length of tympanum; tympanic
annulus distinct; supratympanic fold heavy, curved from posterior comer of
eye to point above insertion of forelimb, obscuring upper part of tympanum;
post-tympanic fold absent.

Forearm moderately robust; row of low tubercles on ventrolateral edge of
forearm; fingers long, bearing moderately large, round discs; width of disc on
third finger slightly greater than length of tympanum; relative lengths of
fingers 1<2<4<3; fingers webbed basally; webbing formula for outer fingers
112* — 3*1113* — 31 V; subarticular tubercles large, round, none bifid; supernu-
merary tubercles moderately large, round, in two rows on proximal segments;
palmar tubercle large, flat, bifid; prepollical tubercle large, elliptical; nuptial
excrescences absent. Hind limb moderately robust; tibia length 46.8-53.2
(Y=48.7, n=l) percent of snout-vent length; foot length 45.7-48.7 (3c=47.0,
n=7) percent of snout-vent length; round, elongate, tubercle on heel; tarsal
folds absent; ventral surfaces of tarsus with many low, diffuse tubercles; inner
metatarsal tubercle moderately small, flat, elliptical; outer metatarsal tubercle
not distinguishable; toes long, bearing discs slightly smaller than those on
fingers; relative lengths of toes 1<2<3=5<4; toes about one-third webbed;
webbing formulaI2— 2*n2-— 302— 3IV3— 2V; subarticular tubercles laige,
round; supemumerary tubercles large, in single row on proximal segments,
continuous with tubercles on tarsus.

Skin on belly and proximal ventral surfaces of thighs weakly granular;
other surfaces smooth; anal opening directed posteroventrally at midlevel of
thighs in males, posteriorly at upper level of thighs in females; dermal supra-
anal flap long, transverse; pair of vertically elongate swellings ventrolateral
to anal opening. Vomerine odontophores long, transverse, abutting medially,
behind level of large, ovoid choanae, bearing 10-18 (:f=13.2, n=7) teeth;
tongue broadly cordiform, shallowly notched posteriorly, barely free behind;
vocal slits extending from midlateral base of tongue to angles of jaws; vocal
sac single, median, subgular.

Coloration in preservative: Dorsal surfaces of head, body, shanks, and
forearms brown with faintly darker brown flecks and scattered minute cream
flecks (two females with narrow, irregular, transverse tan lines); markings on
forearms and shanks in form of transverse bars; flanks dark brown with 3-5

12 OCCASIONAL PAPERS

vertical cream bars; all surfaces of upper arms and thighs, inner surfaces of
forearms and tarsi, and dorsal surfaces of inner two fingers and inner three
toes cream with black vertical bars; throat tan with small dark brown and
cream flecks (fine cream reticulation in one female and two longitudinal
cream bars in one female); belly grayish brown with diffuse dark brown spots
and cream flecks (Fig. 4E).

Coloration in life: Dorsum grayish brown with dark brown and minute
cream flecks (tan middorsal stripe in two specimens); flanks, upper arms,
thighs, ventral shanks, and dorsal surfaces of feet cream with dark brown bars;
throat, chest, and belly creamy gray with brown and cream flecks. Iris dull
bronze with fine black reticulations.

Measurements: The measurements of four males are followed by those of
three females (mm with means in parentheses): snout-vent length 53.6-55.6
(55.0), 58. 1-63.4 (60.2); tibia length 24.8-26.5 (25.5), 28.5-32.2 (30.6); foot
length 24.5-26.3 (25.5), 27.5-30.4 (28.7); head width 17.0-17.7 (17.5),
18.4-20.8(19.7); head length 17.1-18.2(17.8), 18.6-21.3 (20.0); interorbital
distance 4.5-5.2 (5.1), 5.3-6.2 (5.9); intemarial distance 4.4^.9 (4.7),
5.0-5.6 (5.3); eye-nostril distance 5.0-5.6 (5.3), 5.4-5.6 (5.5); diameter of
eye 5.0-5.6 (5.2),4.8-6.2 (5.4); diameter of tympanum 2.3-3.0 (2.8), 2.9-3.2
(3.0).

Distribution and ecology. — Hyla psarolaima is known from cloud for-
ests at elevations of 1950-2660 m on the Amazonian slopes of the Cordillera
Oriental in southern Colombia and Ecuador. Most individuals were found on
stems and branches of bushes near, or over, streams in cloud forest at night.
Two females were in terrestrial bromeliads by day.

Etymology. — The specific name is derived from the Grod^ psaros mean-
ing speckled and the Greek laimos meaning throat; the name is used in
reference to the brown and cream flecks on the throat.

Remarks. — The specimens from Colombia and Papallacta, Ecuador,
were referred to H. larinopygion by Duellman and Altig (1978).

Hyla ptychodactyla new species
Fig. 6

Holotype.— KU 209780, an adult male, from Pilalo (00°57'S, 79°02'W),
2320 m, Provincia de Cotopaxi, Ecuador, obtained in July 1987 by Giovanni
Onore.

Paratopotypes.— KU 209781, MHNG 18715, 18724 (2 specimens), all
collected by Giovanni Onore.

Diagnosis. — This species differs from all other members of the group by
having pale spots on the throat, mottled black and tan dorsum, a conical
tubercle on the heel, and a pale blue iris in life. All other species in the group
have either a uniformly black venter (//. lindae, sarampiona, and species A),

SYSTEMATICS OF HYLA LARINOPYGION GROUP

13

Fig. 6. Hyla ptychodactyla from Pilal6, Provincia de Cotopaxi, Ecuador. Photo
courtesy of Luis A. Coloma R.

boldly mottled black and cream or blue and cream venters (//. larinopygion,
pacha, and panto sticta), or gray venter with brown and cream spots (//.
psarolaima). Dermal appendages are absent on the heels of//, pantosticta and
sarampiona; an elongate tubercle is present on the heel in //. psarolaima, and
small, triangular calcars are present in the other species in the group, Hyla
ptychodactyla is unique among Andean hylids in having a pale blue iris in life.

Description. — Body robust; head as long as wide, about as wide as body;
head width 32.2-34.0 (3c=32.8, n=5) percent of snout-vent length; head
length 32.0-34.0 (^=33. 1, /i =5) percent of snout-vent length; length of snout
less than diameter of eye; snout in dorsal view truncate, in profile bluntly
rounded; canthus round; loreal region concave; lips not flared; nares barely
protuberant laterally, directed anterolaterally at level of anterior margin of
lower jaw; intemarial area slightly depressed; top of head flat; interorbital
distance greater than width of eyelid; eye moderately large, prominent;
tympanum nearly round; separated from eye by distance 1 .5 times length of
tympanum; tympanic annulus distinct; supratympanic fold heavy, extending
from posterior comer of eye to point above insertion of forelimb, obscuring
upper part of tympanum; post-tympanic fold moderately heavy, beginning
under supratympanic fold, extending to angle of jaw, obscuring posteroven-
tral edge of tympanum.

Forearm moderately robust; ulnar tubercles absent; fingers long, bearing
moderately large, round discs; width of disc on third finger slightly greater
than length of tympanum; relative lengths of fingers 1<2<4<3; fingers

14 OCCASIONAL PAPERS

webbed basally; webbing between Fingers I and n vestigial; webbing
formula for outer fingers 112— (3-3)in(3-3)— (2-2y2)IV; subarticular
tubercles large, round; supernumerary tubercles, large, subconical, in two
rows on proximal segments; palmar tubercle large, flat, bifid; prepollical
tubercle large, elliptical; nuptial excrescences absent. Hind limb moderately
robust; tibia length 46.9-49.0 ()c=47.8, n=5) percent of snout-vent length;
foot length 45.5^7.0 (^=46.2, n=5) percent of snout-vent length; heel
bearing blunt, conical tubercle; tarsal tubercles, tarsal fold, and outer meta-
tarsal tubercle absent; inner metatarsal tubercle moderately latge, flat, ovoid,
visible from above; toes long, bearing discs nearly as large as those on fingers;
relative lengths of toes 1<2<3<5<4; toes about two-thirds webbed; webbing
extending as lateral flaps onto bases of discs; webbing formula I(l-iy2) — (l"
-2)111— (2--2)ra(2-2)—(2y2-3)IV(2y2-3-)—(l-lV2)V; subarticular tu-
bercles large, round; supernumerary tubercles low, round, in single row on
proximal segments.

Skin on dorsum and flanks finely shagreened; skin on throat finely
granular; skin on belly and proximal ventral surfaces of thighs coarsely
granular; other surfaces smooth; anal opening directed posteroventrally at
midlevel of thighs in males, posteriorly at upper level of thighs in female;
dermal supra-anal flap long, transverse; pair of vertically elongate swellings
lateral to anal opening. Vomerine odontophores long, transverse, abutting
medially, behind level of large, round choanae, bearing 11-15 (y= 13. 1 , n =5)
teeth; tongue round, not free behind; vocal slits curved from lateral base of
tongue toward angles of jaws; vocal sac single, median, subgular.

Coloration in preservative: Dorsal surfaces of head and body dull reddish
brown and black; pattern on head and body of holotype consisting of irregular,
transverse black markings; female paratype dull brown with faint dark flecks;
other specimens having intermediate types of patterns, but one with faint,
narrow, pale middorsal line. Ranks uniform black (3 specimens) or with
reddish brown extensions of dorsal ground color (2 specimens); groin and
proximal surfaces of thighs black; restof thighs black with 1-3 transverse pale
bars on all but ventral surfaces. Forearms, shanks, tarsi, fingers, and toes with
black and reddish brown transverse marks- 2 or 3 black bars on forearms, 3
or 4 on shanks, 2 or 3 on tarsi. Venter black with large, irregular cream spots
on throat (also on chest of female).

Coloration in life: Dorsum orange-tan to reddish brown with black
markings (minute orange-tan flecks enclosed in black markings); venter
black with cream spots on throat; iris pale blue.

Measurements: The measurements of four males (mm with means in
parentheses) are followed by those of the single female: snout-vent length
65.0-67.5 (65.9), 77.3; tibia length 30.5-32.5 (31.7), 36.3; foot length
29.6-30.9 (30.5), 35.8; head width 21.2-23.0 (21.8), 24.9; head length
21.4-23.0 (22.0), 24.7; interorbital distance 8.0-8.2 (8.1), 8.7; intemarial

SYSTEMATICS OF HYLA LARINOPYGION GROUP 15

distance 4.9-5.8 (5.2), 6.0; eye-nostril 5.2-5.5 (5.3), 5.5; diameter of eye
6.2-6.4 (6.3), 6.5; diameter of tympanum 3.4-4.0 (3.6), 4.0.

Tadpoles. — Although the largest tadpole has a body length of 27.2 mm
and a total length of 78.4 mm, it has not yet developed hind limb buds. The
following description is based on KU 180360.

Body length 20.4 mm; total length 64.9 mm; body depressed, wider than
deep, ovoid, widest just posterior to eyes; snout bluntly rounded in dorsal
view and in profile; eyes small, widely separated, directed dorsolaterally;
nostrils about midway between eyes and snout; spiracle sinistral, its opening
directed posterodorsally at level of midline about two-thirds length of body;
cloacal tube short, dextral. Caudal musculature robust, gradually tapering to
pointed tip just short of tip of acutely rounded tail; dorsal and ventral caudal
fins about equal in depth; at midlength of tail depth of each fin slightly
shallower than depth of caudal musculature.

Mouth large, directed ventrally, completely bordered by single row of
small, pointed papillae; larger, conical papillae in lateral folds; beaks slender,
finely serrate; upper beak forming broad arch; lower beak broadly V-shaped;
seven upper and eight lower rows of denticles (one more upper and two more
lower rows in largest tadpole); innermost three upper rows, and innermost
lower row narrowly interrupted medially.

In preservative, body and caudal musculature brown; fins translucent. In
life, body gray flecked with black; caudal musculature finely reticulated with
brown; golden streak along top of caudal musculature edged with black stripe;
fins colorless except for fine brown reticulation; iris pale green.

Distribution and ecology. — The species is known only from the type
locality in disturbed cloud forest. Adults were found by day by tracing their
calls, a series of tick-like notes, to frogs under deep moss along seepage areas
by streams. The tadpoles were found at night in pools in a stream in a clearing.

Etymology. — The specific name is derived from the Greek ptychos
meaning fold or flap and the Greek dactylos meaning toe; the name alludes
to the dermal flaps on either sides of the toes that are extensions of the
webbing.

Remarks. — The type locality is a small village on the Pacific slopes of the
Andes; it is 78 km east of Quevedo on the road to Latacunga. The tadpoles of
this species were discovered by John D. Lynch in July 1970 and were
subsequently collected by him again in 1977, at which time he succeeded in
raising two individuals, one to Stage 42 with a snout- vent length of 32.9 mm
and a tail stub 33.4 mm long, and the other to metamorphosis with a snout-
vent length of 3 1 .0 mm.

Key to Adults of the Hyla larinopygion Group

1. Venter uniformly or predominately black 2

Venter mottled black and white or blue, or brownish gray with brown spots

16 OCCASIONAL PAPERS

and cream flecks 5

2. Throat uniformly black; no pale marks on flanks or hidden surfaces of

limbs H.lindae

Throat black with orange or cream flecks; hidden surfaces of limbs with
red spots or cream bars 3

3. Dorsum pale olive-brown with small orange spots; hidden surfaces of

limbs with red spots H. sarampiona

Dorsum brown with irregular black marks; hidden surfaces of limbs black
with pale bars 4

4. Digital discs dark; irregular, large pale spots on throat

H.ptychodactyla

Digital discs pale; small pale flecks on throat H. species A

5. Venter brownish gray with dark brown spots and cream flecks; dorsum

brown with small, dark brown and cream flecks H. psarolaima

Venter mottled dark brown or black and white or blue; dorsum uniform
brown or with orange (cream in preservative) spots or flecks 6

6. Dorsum uniform brown; flanks and hidden surfaces of thighs with bold

black and blue (cream in preservative) vertical bars H. larinopygion

Dorsum brown or black with orange (cream in preservative) spots or
flecks; flanks spotted, or barred 7

7. Dorsum brown with small orange flecks; throat and belly dark brown with
bold cream mottling; hidden surfaces of limbs black with narrow, vertical,

cream bars; discs on digits cream and black H. pacha

Dorsum and throat black with orange spots; belly black with white
mottling; hidden surfaces of limbs black with orange spots; discs on digits
yellow H.pantosticta

GENETIC DIVERGENCE AND
PHYLOGENETIC RELATIONSfflPS

Genetic divergences among the species in the Hyla larinopygion group
were considerable and ranged from 0.329 to 1.242 D (Table 3, Fig. 7).
Interestingly, the two most similar species genetically are the two described
herein from the Amazonian slopes of the Andes in Ecuador. We collected
these two species at elevations separated by only 290 meters on the same slope
in central Ecuador. However, they are quite distinct morphologically and
genetically.

The allozymic data were used to construct a proposed phylogeny of the
members of the group (Fig. 8). For this analysis, we identified symplesiomor-
phic electromorphs at 1 1 of the 1 8 loci by using a wide diversity of outgroups
(Tables 1,4). Variation at two of the loci (IDH and ME) suggest that //. lindae
is the sister species to the remaining taxa; this permitted the use of//, lindae

0.519

0.492

0.693

0.987

1.066

1.242

—

0.618

0.618

—

0.329

SYSTEMATICS OF HYLA LARINOPYGION GROUP 17

Table 3. Modified Nei's genetic distances (Hillis, 1984) among
species in Ihc Hyla larinopygion group.

Species H. lindae H. larinopygion H. psarolaima H. pacha

H. pantosticta 0.703

H. lindae —

H. larinopygion —

H. psarolaima —

as a functional outgroup (Watrous and Wheeler, 1981; Farris, 1982) to the
remaining species. Similarly, variation at the EST locus suggests that H.
lindae and H. pantosticta can be considered as functional outgroups to the
other three species. By using this information, it was possible to code
transformation series for all 18 loci, although more than one equally parsimo-
nious alternatives is possible for MDH-1 (this locus is diagnostic for each of
the species). The resulting cladogram (Fig. 8) contains no homoplasy.

Morphologically, H. psarolaima and H. larinopygion are the most similar
species in the group, a fact that resulted in confusion of the two species in the
past (Duellman and Al tig, 1 978). However, these taxa apparently are not sister
species; the electrophoretic data indicate that H. psarolaima is more closely
related to the morphologically distinct H. pacha than to the morphologically
similar//, larinopygion. These data suggest that//, pacha has diverged farther
from their common ancestor, both morphologically and allozymically, than
has //. psarolaima.

Although most of the morphological differences among species in this
group are autapomorphic, some morphological characters are consistent with
the phylogeny based on allozymes. Three of the species (//. larinopygion,
pacha, and psarolaima) have distinct vertical hght and dark bars on the their
flanks and limbs. These three species form a monophyletic group according
to the allozymic data, although this part of the tree is supported by only one
electromorphic change (Fig. S).Hyla lindae andpantosticta (also//, sarampi-
ona, for which allozymic data are not available) have pale discs on the digits,
which distinguish them from the branch of the tree containing//, larinopygion,
pacha, and psarolaima. Among the species for which allozymic data are
available, //. lindae is unique in having a plain, dark venter, a character shared
with //. sarampiona and //. sp. A. The available morphological data are
insufficient to determine the relationships of//, sarampiona and H. sp. A, but
they seem to be more like //. lindae than the other species in the group.
However, as noted for//, larinopygion and psarolaima, phenetic similarity of
color patterns may be misleading.

18

OCCASIONAL PAPERS

•I

:»:

00

-

0 1

-

0.2

-

0.3

o

c
o

b

0.4
0.5

-

o

c
o

0.6
0.7

-

0 8

-

0.9

-

1.0

•V.

5:

I

<5

«3

0329

0 519

0.605

1.000

Fig. 7. Phenogram of modified Nei's genetic distances (Hillis, 1984) among five taxa
in the Hyla larinopygion group.

Differences exist in the configuration of the caudal fins and in the rows of
labial denticles in the tadpoles. The tadpole of//, lindae has relatively deep
caudal fins and six upper and eight lower rows of denticles. The tadpole of//.
pantosticta has very shallow fins and four upper and six lower rows of
denticles. Tadpoles from a stream near localities where //. pacha and
psarolaima were found are like those of //. lindae but have eight upper and
nine lower rows of denticles; these tadpoles may belong to either of those was

SYSTEMATICS OF HYLA LARINOPYGION GROUP 19

Table 4. Electromorphs of the Hyla larinopygion group.
Alleles found in both the ingroup and the outgroup are indicated under
the outgroup; if no alleles could be found in common between the
groups, an "x" is shown for the outgroup.

Lx)cus

larinopygion

lindae

pacha

pantosticta

psarolaima

Outgroup

AAT-1

e(l.OO)

d (1.00)

b (0.25)
c (0.75)

a (0.50)
d (0.50)

c (1.00)

X

AAT-2

a (0.50)
b (0.50)

b(l.OO)

b (1.00)

b (1.00)

b (1.00)

b

CK

a (1.00)

a (1.00)

a (1.00)

a (1.00)

a (1.00)

X

EST

b (1.00)

a (1.00)

b (1.00)

a (1.00)

b (1.00)

a

GPI

c (1.00)

a (0.83)
b (0.17)

d(l.OO)

c (1.00)

d(l.OO)

a

G-3-PD

a (1.00)

a (1.00)

a (1.00)

a (1.00)

a (1.00)

X

GPD

a (1.00)

a (1.00)

a (1.00)

a (1.00)

a (1.00)

X

IDH

b (1.00)

a (1.00)

b (1.00)

b (1.00)

b (1.00)

a

LDH-1

b(l.OO)

b (1.00)

b (1.00)

a (1.00)

b (1.00)

b

LDH-2

b (1.00)

a (1.00)

b (1.00)

b (1.00)

b (1.00)

b

MDH-1

b(l.OO)

c (1.00)

a (1.00)

e (1.00)

d(l.OO)

X

MDH-2

a (0.50)
e (0.50)

d(l.OO)

c (1.00)

b (1.00)

b (1.00)

d

ME

b (1.00)

a (1.00)

b (1.00)

b (1.00)

b (1.00)

a

PEP-A

a (1.00)

a (1.00)

a (1.00)

a (1.00)

a (1.00)

a

PEP-B

c(l.OO)

a (1.00)

b (1.00)

a (1.00)

a (1.00)

X

PEP-S

c (1.00)

c (1.00)

b(l.OO)

c (1.00)

a (1.00)

c

PGM

a (1.00)

b (0.33)
c (0.67)

a (1.00)

a (1.00)

a (1.00)

a

TPI

c (1.00)

a (0.83)
b (0.17)

b (1.00)

b(l.OO)

b (1.00)

X

species or be a variant of H. lindae. Tadpoles of H. ptychodactyla have
shallow fins and seven upper and eight lower rows of denticles.

Additional specimens of adults are needed for skeletal preparations, and
tadpoles of all of the taxa need to be identified before the relationships of all
of the species may be determined. These frogs seem to be extremely localized
in their distributions, and we suspect that several additional species await
discovery. Consequently, a biogeographic synthesis is premature. However,
the localized distributions in cloud forests in all three major Cordilleras in the

20

OCCASIONAL PAPERS

H. psarolaima H. pacha

H. larinopygion

u- -i. luinu-i X-dl

TP|b-c

pep-bq-c

H. lindae

)TPI

b-a

IMDH-I^'C
ILDH-2'''°

H. pantosticta

►ldh-1

IDH°-b

)GPI

a-b

► GPI

a-c

iTPl'^-b

'PEP-B''"^
GPDX-a

G-3-PD "-a
CKX-a

»AAT-1*-<^

OUTGROUP

Fig. 8. Cladogram of hypothesized phylogenetic relationships among five species in
the Hyla larinopygion group based on allozymic data. Solid circles mark the appear-
ance and fixation of new alleles (e.g., GPI'^'marks the evolution of allele c from allele
a at the GPI locus). Open circles mark the appearance of new alleles without fixation
(i.e., the ancestral allele is retained in a polymorphic state). Ancestral alleles repre-
sented by X could not be determined in the analysis.

northern Andes suggest that the pattern of speciation and biogeography may
support Duellman's (1982) hypothesis of alternate compression and expan-
r.lniirt forp.sts during the Oiiatemarv.

sion of cloud forests during the Quaternary.

ACKNOWLEDGMENTS

The collection of these elusive frogs was aided by the efforts of Patricia A.
Burrowes, John E. Simmons, and Linda Trueb. Field work in South America

SYSTEMATICS OF HYLA LARINOPYGION GROUP 21

supported by grants (DEB 74-02998 and 82-19388) from the National
Science Foundation to W. E. Duellman. Laboratory work was supported by
grants (BSR 86-57640 and 87-96293) from the National Science Foundation
to David M. Hillis. Collecting permits for Colombia were issued by Jorge I.
Hemandez-C. of the Instituto de los Recursos Naturales Renovables, and for
Ecuador by Sergio Figueroa and Abel Tobar V. of the Ministerio de Agricul-
tura y Ganaderia. We are grateful to Volker Mahnert, Jose E. Rosado, and
Pedro M. Ruiz-C. for the opportunity to examine material in their care, to Juan
M. Renjifo for the tissues of one species, to Luis A. Coloma R. and John D.
Lynch for information on the distribution and ecology of some species, to
Rafael de Sa for translating the resumen, and to Anne M. Musser for preparing
the illustrations.

RESUMEN

El grupo de especies de Hyla larinopygion contiene ocho especies cono-
cidas que habitan el bosque nebuloso de las laderas de los Andes de Ecuador
y Colombia, a alturas entre 1900 y 3300 m. Se describen tres nuevas especies,
H. pacha y H. psarolaima de las laderas amazonicas de los Andes ecuatori-
anos y H. ptychodactla de las laderas pacificas de los Andes ecuatorianos. Se
obtuvo informacion aloenzimatica de cinco especies y mediante electroforesis
en geles de almidon se registraron 18 loci diferentes. Basados en los electro-
morfos, el cladograma mas parsimonio indica a H. lindae como la especie
hermana a las demas especies y tambien a grupo a H. larinopygion, psaro-
laima, y pacha en un mismo clade que tiene a H. pantosticta como especie
hermana. El cladograma tambien es soportado por caracteristicas del patron
de coloracion.

LITERATURE CITED

Duellman, W.E. 1970. The hylid frogs of Middle America. Monogr. Mus. Nat. Hist.

Univ. Kansas 1:1-753.
Duellman, W.E. 1972. A review of the neotropical frogs of the Hyla bistincta group.

Occas. Pap. Mus. Nat. Hist. Univ. Kansas 11:1-31.
Duellman, W.E. 1973. Descriptions of new hylid frogs from Colombia and Ecuador.

Herpetologica 29:219-227.
Duellman, W.E. 1982. Compresion climatica cuatemaria en los Andes: efectos sobre

la especiacidn. Pp. 1 77-201 in P. J. S alinas (ed.). Zoologica Neotropical, Actas VIII

Congreso Latino americano de Zoologia. Merida, Venezuela.
Duellman, W.E. 1989. New species of hylid frogs from the Andes of Colombia and

Ecuador. Occas. Pap. Mus. Nat. HisL Univ. Kansas 131:1-12.
Duellman, W.E., and R. Altig. 1978. New species of tree frogs (family Hylidae)

22 OCCASIONAL PAPERS

from the Andes of Colombia and Ecuador. Herpetologica 34:177-185.
DuELLMAN, W.E., AND T.J. Berger. 1982. A new species of Andean treefrog (Hyli-

dae). Herpetologica 38:456-460.
Farris, J.S. 1982. Outgroups and parsimony. Syst. Zool. 31:328-334.
GosNER, K.L. 1960. A simplified table for staging anuran embryos and larvae with

notes on identification. Herpetologica 16:183-190.
HiLUS, D.M. 1984. Misuse and modification of Nei's genetic distance. Syst. Zool.

33:238-240.
HiLLJS, D.M. 1985. Evolutionary genetics of the Andean lizard genus Pholidobolus

(Sauria: Gymnophthalmidae): phylogeny, biogeography, and a comparison of tree

construction techniques. Syst. Zool. 34:109-126.
Leviton, A.E., R.H. Gibbs, Jr., E. Heal, and C.E. Dawson. 1985. Standards in

herpetology and ichthyology: Part I. Standard symbolic codes for institutional

resource collections in herpetology and ichthyology. Copeia 1985:802-832.
Rufz-C, P.M., AND J.D. Lynch. 1982. Dos nuevas especies de Hyla (Amphibia:

Anura) de Colombia, con aportes al conocimiento de Hyla bogotensis. Caldasia

8:647-671.
Sneath, P.H.A., AND R.R. Sokal. 1973. Numerical taxonomy: the principles and

practice of numerical classification. W. H. Freeman and Co., San Francisco.
Watrous, L.E., AND Q.D. Wheei^r. 1981. The outgroup comparison method of

character analysis. Syst. Zool. 30:1-11.

APPENDIX 1: SPECIMENS EXAMINED

Tissues for allozymic analysis were obtained from those numbers desig-
nated by an asterisk.

Hyla larinopygion. — COLOMBIA: Cauca; Quebrada Santa Teresa, 2200
m, KU 144127-28. [Also see Materials and Methods*].

Hyla lindae.— ECUADOR: Morona-Santiago; 21.6 km WSW Plan de
Milagro, 2350 m, KU 202728*-29*; 23.3 km WNW Plan de Milagro, 2125
m, KU 202730-31*; 202966 (tadpoles). Napo: 11 km ESE Papallacta, 1660
m, KU 155475, 155477 (skeleton), 155523 (tadpoles), 164402, 166210 (tad-
poles), 171188 (tadpoles).

//y/fl;7ac/ia.— ECUADOR: Morona-Santiago: 1 1 .2 km WSW Plan de Mi-
lagro, 2350 m, KU 202760*-61*, 202762; 12.4 km WSW Plan de Milagro,
2225 m, KU 202763.

Hyla pantosticta.— COLOMBIA: Narino: La Victoria, 2700 m, KU
140385. ECUADOR: Carchi: 0.5 km S Santa Barbara, KU 190000-01. Napo:
1 km E Santa Barbara, 2520 m, KU 202733*; 12 km E Santa Barbara, 2550
m, KU 202965 (tadpole); 18 km E Santa Barbara, KU 202732.*

Hyla psarolaima. — COLOMBIA: Putumayo: 35 km SE San Francisco,
1950 m,KU 169582-85. ECUADOR: Morona-SanUago: 26.2 km WSW Plan
de Milagro, 2640 m, KU 202727.* Napo: Between La Alegria and Sibundoy,

SYSTEMATICS OF HYLA LARINOPYGION GROUP 23

2410 m, MCZ 109317-18; 11 km ESE Papallacta, 2660 m, KU 164313-15,
164316 (skeleton).

Hyla ptychodacty la, —ECUADOR: Cotopaxi: Pilalo, 2320 m, KU
132553-54 (tadpoles), 180360 (tadpoles), 180361-62 (young), 209780-81,
MHNG 18715, 18724 (2).

Hyla sp. A, — COLOMBIA: Cauca: Paez, 34.5 km between Belalcazarand
Tacueyo, 2400 m, KU 189598.

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