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LAWRENCE, KANSAS

OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY The University Mtfe.Kansa^Q^ Lawrence, Kansas library

NUMBER 22, PAGES 1-27 FEB 1 j 'HNUARY 25, 1974

A SYSTEMATIC REVIE\Y|S{^^e> MARSUPIAL

FROGS (HYLIDAE: GAsfWfHECA)

OF THE ANDES OF ECUADOR

By William E. Duellman^

Many species of marsupial frogs, genus Gastrotheca, occur in the Andes and associated cordilleras from western \'enezuela soutli- ward to nortliern Argentina. Because of the paucity of specimens of many of the named taxa and confusing variation exhibited by some species, the taxonomy of the marsupial frogs has been chaotic. Duellman and Fritts (1972) reviewed the species occurring in the Andes to the south of the Huancabamba Depression in northern Peru and delimited sev(m speci(\s there as members of the Gastro- theca marsnpiata group. These frogs differ in cranial characters from the species in the Huancabamba Depression and northward, all of which were referred to the Gastrotheca argenteovirens group l)y Duellman and Fritts (1972).

The Ecuadorian species included in the latter group were: G. lojana Parker, 1932; G. monticola Barbour and Noble, 1920; G. phimbea (Boulenger, 1882); and G. riobambae (Fowler, 1913). Colombian and Venezuelan species assumed to belong to the same group were: G. argenteovirens (Boettger, 1892); G. aureomaculata Cochran and Coin, 1970; G. helenae Dunn, 1944; G. meclenii Coch- ran and Coin, 1970; G. mertensi Cochran and Coin, 1970; and G. nicefori Gaige, 1933. It now seems apparent that these frogs are treated best as two distinct species groups. The Ecuadorian species listed above and two species named in this paper can be referred to as the Gastrotheca plinnbea group, and the Colombian and Vene- zuelan species, as the Gastrotheca argenteovirens group. \Mien bet- ter known, G. wiUiamsoni Gaige, 1922, and an unnamed species re- lated to G. phnnbea from Antioquia and Cundinamarca, Colombia,

' Curator, Di\ ision of Herpetology, Museum of Natural History, University of Kansas.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

may be referable to the Gastrotheca argenteovirens and plumbea groups, respectively.

Thus, this paper is a review of the Gastrotheca plumbea group. The purpose of the preseut paper is to provide a synthesis of accu- mulated information and in so doing, 1) define the species, 2) as- sign names to taxonomically recognizable populations, and 3 ) sum- marize data on geographic variation, distribution, ecology, and life history. The results presented here, together with the synthesis of the southern Andean frogs by Duellman and Fritts (1972), will provide a basis for a future systematic assessment of the Gastro- theca argenteovirens group.

ACKNOWLEDGMENTS

For the loan of specimens or provisions of facilities in their re- spective institutions, I am grateful to F. Bernis, James E. Bohlke, F. W. Braestrup, Nelly Carrillo de Espinoza, Javier Castroviejo, Josef Eiselt, Alice G. C. Grandison, Jean Guibe, Konrad Klemmer, Alan E. Leviton, Robert F. Martin, Hymen Marx, Charles W. Myers, Giinther Peters, the late James A. Peters, Greta Vestergren, Charles F. Walker, and Ernest E. Williams. Much of the new material reported herein was collected by Thomas H. Fritts, John D. Lynch, and Linda Trueb; Arthur C. Echternacht, Bruce Nlac- Bryde, Richard R. Montanucci, and John E. Simmons also aided in the field work. I appreciate their efforts in my behalf.

Field work in Ecuador was supported in part by Watkins Mu- seum of Natural History Grants, The University of Kansas; travel to European museums was provided by a grant from the Penrose Fund (No. 5063) from the American Philosophical Society. I am indebted to Linda Trueb for critical review of my prose and to Thomas H. Fritts for his provocative comments during formulative phases of this work and critical review of the manuscript. My final acknowledgment is to Charles F. Walker, who has given freely his time and advice on marsupial frogs, and who offered valuable com- ments on the manuscript.

MATERIALS AND METHODS

I have examined 1125 preserved frogs (including type speci- mens of all taxa), 24 skeletons, and 50 lots of tadpoles referable to the species discussed in this paper, and I have studied each of the species in the field. Terminology follows that of Duellman (1970), except for two additional measurements: length of third finger is the distance from the proximal edge of the palmar tubercle to the tip of the finger; length of thumb is the distance from the proximal edge of the prepollical tubercle tt) the tip of the thumb. Through- out the text, specimens are listed by their catalogue numbers pre- ceded by the appropriate museum abbreviations, as follows:

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 3

AMNH American Museum of Natural History

ANSP Academy of Natural Sciences of Philadelphia

BMNH British Museum (Natural History)

CAS California Academy of Sciences

CAS-SU Stanford University Collection (in California Academy of

Sciences )

FMNH Field Museum of Natural History

FSM Florida State Museum

KU University of Kansas Museum of Natural History

MCZ Museum of Comparatixe Zoology, Hai-vard University

MJP Museo Javier Prado, Lima, Peru

MNCN Museo Nacional de Ciencias Naturales, Madrid

MNHN Museum National d'Histoire Naturelle, Paris

NHMW Naturhistorisches Museum, Wien

NHRM Naturhistoriska Riksmuseet, Stockholm

SMF Senckenbergische Museum, Frankfurt

TNHC Texas Natural History Collection

UMMZ University of Michigan Museum of Zoology

USNM United States National Museum

UZM Universitets Zoologiske Museum, Copenhagen

ZMB Zoologisches Museum, Berlin

THE CAST ROTH EC A PLUMBEA GROUP

Species comprising? this ^roiip ha\'e sliort to inoderately long legs, narrow to moderately wide heads, r(>lati\'el\' large hands, moderate to extensive exostosis of the dermal roofing l)')nes, broad frontoparietals expanded into lateral flanges, the frontoparietal fontanelle covered by the frontoparietals, and a long cultriform process of the parasphenoid. z\ll species have aquatic tadpoles.

In contrast, members of the Gasiroiheva or<ienteovirens group have long legs, broad heads, large hands, extensive exostosis of the dermal roofing bones and in some species eo-ossifieation and cas- ((uing, and at least in some species a short cultriform process. Inso- far as known, all species in this group have aquatic tadpoles, but at least in C. orp,enteovirens, the tadpoles hatch at an advanced stage, and the larxal period is short. Members of the Gastrotheca marsupiafa group ha\e short legs, narrow heads, small hands, no exostosis of the dermal roofing bones, small narrow frontoparietals not roofing the frontoparietal fontanelle, and a long cultriform process. Three species have aquatic tadpoles; in the other four development is completed in the maternal pouch.

ACCOUNTS OF SPECIES

In the following accounts, each taxon is diagnosed; the variation is discussed, and the distribution of each species is annotated. Per- tinent measurements and proportions are given in Table 1, and the frogs are illustrated in figures 1 and 2. Maximum snout-vent lengths are given in the diagnoses.

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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MARSUPIAL FROGS OF THE ANDES OF ECUADOR 5

Gastrotheca cavia new species

Holotype. — KU 148532, an adult female, 64.0 mm, from Isla Pequeiia, Laguna Cuicocha, Pro\'incia Imbabura, Ecuador, 2890 m; one of a series collected on 31 October 1971 by William E. Duell- man and John E. Simmons.

Paratopotypes.—KV 138216-20, 24 July 1970, Thomas H. and Patricia R. Fritts; 139137-9, 30 Januaiv 1971, William E. Duellman; 143094, 148530-1, 148533-36, 31 October 1971, William E. Duell- man and John E. Simmons; 148537-40, 30 January 1971, William E. Duellman.

Diagnosis. — 1) Body robust, 58.5 mm in males, 67.2 in females; 2) snout round in dorsal view, rounded above and anteroventrally inclined in profile; 3) canthus rounded; 4) loreal region shallowly concave; 5) tympanum vertically elliptical; 6) suprat\'mpanic fold moderately heavy; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand low, round; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-third webbed; 12) tar- sal fold low, extending one-third length of tarsus; 13) inner meta- tarsal tubercle low, ovoid, visible from above; 14) outer metatarsal tubercle low, flat; 15) subarticular tubcTcles on foot large, round: 16) supernumerar\' tubercles on foot small, low; 17) discs round; 18) dorsal skin shagreened, tubercular in tympanic region; 19) dor- sum green or tan with or without small irregular black spots; narrow bronze dorsolateral stripe usually present; 20) facial area uniform green or tan; canthal and labial stripes absent; 21) flanks tan with numerous small black spots; 22) dorsal surfaces of limbs plain or with small black spots; 23) posterior surfaces of thighs cream with black mottling; 24) venter cream with black mottling; 25) sciua- mosal exostosed, in broad contact with maxillary; 26) temporal arcade complete in large indi\'iduals; 27) prevomers abutted medi- ally; 28) transverse processes on eighth presacral vertebra slightly inclined anteriorly.

GasfrotJwca cavia resembles G. riohamhac in having short legs, a narrow intt>rorbital distance, and a short snout. It difi^ers from G. riohamhae in having a distinct dorsolateral light stripe, small black flecks on the flanks, and dorsal markings comprised of small black spots. Gasirotheca riohamJ)ae lacks a dorsolateral light stripe and has large dark spots on the flanks and a dorsal pattern consist- ing of paired elongate dark marks; the dorsal pattern is shared with G. lojana and inonticola. Gastrotheca plumhea and psychrophila have unmarked dorsal surfaces and uniformly dark flanks and ven- ters. The venter in G. cavia is cream with small dark spots.

Variation. — \\''hereas some indi\'iduals have only a few black flecks on the posterior part of the dorsum, others have more flecks over the (Mitire dorsum; in more heavily flecked specimens, flecks on the shank tend to form transverse bars. In life, adults are green;

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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Fig. 1. A. Gasiwtheca cavia, 9, SVL 60.0 mm, KU 139139. B. G. lojana, $ , SVL 56.5 mm, KU 148549. C. G. monticola, 9 , 65.0 mm, KU 148568.

MARSUPIAL FROGS OF THE ANDES OF ECUADOR

Fig. 2. A. Gastrotheca phnnhea, i, S\h 55.3 mm, KU 142614. B. G. pstjchrophila, ?, S\'L 61.0 mm, KU 120650. C. G. liohamhae, $, 46.5 mm, KU 120725.

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

the flanks are cream or bronze witli black flecks, fused into reticii- kitions in some individuals. The groin and posterior surfaces of the thighs are pale blue with black flecks. The dorsolateral stripe, ex- tending from the posterior edge of the eye to the groin, usually is bronze and distinct. In some individuals, the stripe is narrow and fragmented by black flecks; in others it is cream, and in one female it is absent. The throat is pale yellow; the rest of the venter is dull grayish white with small black spots. The iris is dull bronze, heavily marked with brown spots and black flecks. Juveniles are pale green or bronze-tan; the flanks and hidden surfaces of the limbs are pale greenish yellow with black spots, and the venter is pale yellow with or without black reticulations. Although most males and all females are green, three males are tan.

Distribution. — This species is known definitely from only two localities in the Cordillera Occidental in northwestern Ecuador (Fig. 3). The species is abundant on two rocky islands in a crater lake, Laguna Cuicocha, at an elevation of 2890 m on the south slope of Volcan Cotacachi. The other locality. Hacienda San Nicolas is at an elevation of 2000 m on the Pacific slope of the Cordillera Occidental. Two specimens from "Western Ecuador" (BMNH 1860.6.16.124-125) and one from Ibarra (?) (BMNH 1898.4.28.156) are referred to this species.

Remarks. — Many adults were found in large terrestrial brome- liads in July, 1970, January and October, 1971. At the time of the last two visits, some adults were found beneath rocks. Brooding females were obtained in January and July, but not in October. Tadpoles were found in the lake surrounding the island in January, and at the same time two metamorphosing young were observed on reeds in the lake. Tadpoles have two upper and three lower rows of denticles.

Superficially, Gastrotheca cavia resembles G. argenteoviretis Boettger; I have examined the type of the latter (SMF 2676) and have compared living and preserved specimens of argenteovirens wath cavia. In comparison with G. cavia, G. argenteovirens has proportionately longer legs (ratio of tibia length to snout-vent length 0.488-0.514, x = 0.498 =b 0.010, N = 5 a $ ; 0.486-0.514, x = 0.501 ± 0.012, N = 4 9 ? ) and broader interorbital distance ( ratio of interorbital distance to head width 0.335-0.371 ± 0.014, x = 0.358, N = 56 6; 0.369-0.395 x = 0.383 ± 0.011, N = 4$ 9 ). The flanks and posterior surfaces of the thigh are mottled with dark blue, and the dorsum lacks black flecks. Furthermore, the cultri- form process of the parasphenoid is short, and in large individuals there is integumentary-cranial co-ossification.

Etymology. — The specific name is the same as the generic name for the guinea pig (Caviidae), called cui in Quecchua the domi-

MARSUPIAL FROGS OF THE ANDES OF ECUADOR

Fig. 3. Distributions of Gastrotheca cavia (triangles), G. 7nonticola (hexa- gons), G. plumbea (inverted triangles), and G. psijchrophila (circles).

nant Indian language in the Ecuadorian Andes. Cnicocha means

lake of the guinea pig.

Gastrotheca lojana Parker New combination

Gastrotheca marstipiata lojana Parker, 1932:25 [Holotype.— BMNH 1931.2.12.4 (RR 1947.2.31.13) from Loja, Pro\ incia Loja, Ecuador; C. Carrion col- lector] .

Diagnosis. — 1) Body depressed, 56.0 mm in males, 60.0 in fe- males; 2) snout round in dorsal aspect and in profile; 3) canthus rounded; 4) loreal region barely concave; 5) tympanum vertically elliptical; 6) supratympanic fold moderately heavy, angled postero- ventrally behind tympanum; 7) subarticular tubercles on hand

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

large, round; 8) supernumerary tubercles on hand small, subconi- cal; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one- half webbed; 12) tarsal fold low, extending one-half length of tar- sus; 13) inner metatarsal tubercle low, elliptical, visible from above; 14) outer metatarsal tubercle low, round; 15) subarticular tubercles on foot large, round; 16) supernumerary tubercles on foot low, round; 17) discs round; 18) dorsal skin shagreened; 19) dor- sum tan or green, with or without pair of elongate dark markings; narrow bronze or cream dorsolateral stripe present; 20) facial area green or tan; bronze canthal stripe and cream labial stripe present; 21) flanks dark brown with cream spots ventrally; 22) dorsal sur- faces of limbs green or tan, with or without narrow brown trans- verse bars; 23) posterior surfaces of tliighs heavily mottled with bluish purple; 24) chin and chest gray; belly pinkish bronze; 25) squamosal exostosed, in narrow contact with maxillary; 26) tem- poral arcade complete; 27) prevomers narrowly separated medial- ly; 28) transverse processes on eighth presacral vertebra transverse.

Gastrotheca lojana resembles G. monticola and riobarnbae in usually having a pair of elongate dorsal markings. It differs from G. riobambae in having fine pale reticulations laterally or dark flanks, nearly uniformly dark posterior surfaces of the thighs, and a dorsolateral light stripe; G. riobambae has large spots on the flanks, mottled posterior surfaces of the thighs, and lacks a dorsolateral light stripe. Gastrotheca monticola has mottled posterior surface of the thighs and further differs from G. lojana in having a wider dorsolateral stripe and a cream venter with dark spots, whereas the venter in G. lojana is dark brown with creamy-wliite spots. The other Andean Gastrotheca in Ecuador lack paired dorsal markings.

Variation. — Variation in coloration can best be described by the following accounts of topotypic adult males ( colors in life ) :

KU 148549. — Tan above with greenish suffusion dorsolaterally; dorsolateral and labial stripes bronze; flanks and dorsal markings dark brown; upper surfaces of thighs bronze-tan; upper surfaces of shanks and posterior surfaces of thiglis dull green; anterior surfaces of thighs dark brown; throat brown; belly brown with white spots; ventral surfaces of thighs pinkish brown; ventral suiiaces of shanks bluish white.

KU 148550. — Tan above with brown markings; face mask and anterior flanks dark brown; labial stripe creamy bronze; anterior and posterior surfaces of thighs mottled dark brown and blue; throat gray-brown; belly brown with white spots.

KU 148551. — Dorsum dull green with dark brown markings; flanks are dark brown; anterior and posterior surfaces of thighs and inner surfaces of shanks mottled dull blue and dark brown; throat dark bronze brown; belly brown with cream spots.

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 11

KU 126073. — Dorsum pale greenish brown with dark brown markings; labial and dorsolateral stripes cream; flanks gray-brown; groin and posterior surfaces of thighs creamy brown with green flecks.

KU 142603. — Dorsum dull leaf green with no distinct markings; canthal, labial, and dorsolateral stripes bronze; flanks bronze- brown; groin and anterior and posterior surfaces of thighs purplish brown; throat pinkish bronze; belly same, becoming darker brown posteriorly, with white spots.

In all specimens the iris is bronze. Four specimens from Cha- chapoyas, Departamento Amazonas, Peru (KU 138238-41) are colored somewhat differently: "Adults with leaf green dorsum; one witli thin beige stripe from nostril through eye to inguinal re- gion and beige spots on flanks; all with white supralabial border; posterior thigh light leaf green; anterior thigh pale leaf green with few black flecks; venter yellow-beige; iris metallic orange. Juvenile with white patch at anus; dorsum tan with light brown blotches edged with dark brown; blotcli l^etween eyes and in\'erted U on body with few scattered small spots posteriori) ; broad rich light brown stripe from eye to midflank; groin gray-white with small black blotches; posterior thigh flesh-pink, leaf green distally; supra- labial area beige- white" (T. H. Fritts, field notes, 1 May 1970).

The dorsal markings are highly variable. Some individuals lack markings except for a few scattered spots. In most indi\'iduals a pair of broad longitudinal marks extend from the scapular region to the groin. In some indi\iduals tlie marks are fragmented into anterior and posterior components, whereas in others the marks are confluent anteriorly. If an interorbital mark is present, it may or may not be connected to the body markings.

Distri])iition. — This species occurs at moderate elevations in the Huancabamba Depression and associated interandean valleys in northern Peru and southern Ecuador, where it has been found from elevations of 2100 to 2350 m in both Atlantic and Pacific drainages (Fig. 4). A record from Zamora, Ecuador, at an elevation of 1000 m on the Amazonian slopes is highK' (luestionable; the specimen (BMNH 1933.6.24.45) is G. lojana, but the locality data probably are erroneous.

Remarks. — In the Loja valley in southern Ecuador this species is found most fre({uently in large Aiiave. where the frogs seek shel- ter at the bases of the leaves by day and call by night. Individuals also have been found under rocks and in marshy meadows. At Chachapoas, Pen'i, they were beneath rocks and clods of earth in a cultivated field. Tadpoles were found in a grassy irrigation ditch. They are uniformly black and have two upper and three lower rows of denticles.

12

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 4. Distributions of Gastrotheca lojana (triangles) and G. riohamhae ( circles ) .

One specimen (FSM 30080, an adult male having a snout-vent length of 48.5 mm) from 24 km WSW of Leimebamba, Departa- mento Amazonas, Peru, 3370 m tentatively is referred to G. lojana. This individual differs from all other G. lojana by having a dark brown dorsum with tan middorsal and dorsolateral stripes; the venter is mottled with dark gray. Futhermore, the locality is 1000 m higher than any other recorded for G. lojana.

Parker (1932:25) named lojana as a subspecies of Gastrotheca marsiipiata, a name which he applied to populations now known as G. riohamhae. As noted in the foregoing diagnosis, G. lojana dif- fers from G. riohamhae in a number of characters; there is no evi- dence for gene flow between the southern populations of G. rioham- hae and G. lojana.

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 13

Gastrotheca monticola Barbour and Noble

Gadrotheca monticola Barbour and Noble, 1920:426 [Holotype. — MCZ 5290

from Huancabamba, Departanienlo Piura, Pen'i; G. K. Noble collector]. Gastrotheca marsupiata monticola — Parker, 1932:25. Gastrotheca monticola monticola — Vellard, 1957:39.

Diagnosis. — 1) Body robust, 60.0 mm in males, 77.0 mm in fe- males; 2) snout round in dorsal view and profile; 3) canthus rounded; 4) loreal region barely eoncave; 5) tympanum nearly round, slightly higher than wide; 6) suprat\'mpanic fold moderately heaw, curved posteroventrally behind t)'mpanum; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand large, conical; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-half webbed; 12) tarsal fold tubercular, ex- tending full length of tarsus; 13) inner metatarsal tubercle elongate, visible from above; 14) outer metatarsal tubercle absent; 15) sub- articular tubercles on feet large, round; 16) supernumerary tuber- cles on hands small, round, flat; 17) discs round; IS) dorsal skin shagreened; 19) dorsum green or tan, usually with paired elongate dark markings; creamy bronze dorsolateral stripe present; 20) facial area green or tan; creamy bronze canthal stripe pesent; 21) flanks brown with cream and black flecks; groin blue; 22) dorsal surfaces of limbs green or tan with or without darker transverse bands; 23) posterior surfaces of thiglis blue; 24) tln-oat gray; chest and belly creamy gray with gray spots; 25) squamosal weakly exastosed, in moderately broad contact with maxillar\'; 26) temporal arcade complete in large indi\'iduals; 27) prevomers narrowly separated medially; 28) transverse processes on eighth presacral vertebra barely inclined anteriorly.

Gastrotheca monticola differs from G. phnnhea and psychw- phila in having a pale venter with black spots, instead of a uni- formly dark venter. Gastrotheca cavia resembles G. monticola in having a broad dorsolateral stripe and mottled posterior surfaces of the thighs, but G. cavia lacks paired longitudinal dark marks on the dorsum, characteristic of G. monticola, lojana, and riohamhae. The latter species differs from G. monticohi in ha\'ing large dark spots, instead of fine reticulations, on the flanks, and in lacking a dorsolateral stripe. Gastrotheca monticola is most like G. lojana, from which it differs in having a pale venter with dark spots ( dark with white spots in lojana), broader dorsolateral stripe, and more mottling on the flanks and thighs.

Variation. — The dorsum is green with paired longitudinal dark markings on the body; a large dark spot on the head, including the eyelids, is present in some individuals. The longitudinal marks extend from the eyelids or occipital region to the groin. In some indi\'iduals the marks are confluent anteriorh'. The marks are green, usually darker than the dorsal ground color, and outlined

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

with brown. The flanks arc brown with or without bronze flecks. The dorsolateral and labial stripes are bronze-tan to metallic cream. The groin and hidden surfaces of the thighs vary from pale green to blue with cream and black mottling. The venter is creamy tan to creamy yellow with brown flecks; the vocal sac is gray. The iris is pale bronze.

Comparison of specimens from Saraguro, Ecuador, with the type series from Huancabamba, Peru, revealed slight differences. The Peruvian specimens have less ventral spotting and more dark- pigment on the posterior surfaces of the thighs. One Peruvian specimen (UMMZ 57747A), a male, has a highly fragmented dor- sal pattern.

Distribution. — Gastrotheca monticola occurs at elevations of 1600-2500 m on the Pacific slopes and associated intermontane val- leys in northern Peru and southern Ecuador (Fig. 3). The Cordil- lera Occidental in that region is dissected by many broad, dry val- leys, so it is unlikely that the distribution of G. monticola is con- tinuous.

Remarks. — At Saraguro, Ecuador, adults were found beneath rocks in a pasture and in and along a vegetation-choked drainage ditch. At Giron, Ecuador, adults were in Agave plants by day. Tadpoles were found in the ditch at Saraguro. The body is dull green above and greenish silver below; the tail is tan with green lichenous markings. The tadpoles have two upper and three lower rows of denticles.

Parker (1932:25) considered G. monticola to be a subspecies of G. marsupiata (= riohamlyae). The differences noted in the diag- nosis obviate such an arrangement in the absence of evidence indi- cating genetic interchange. Cochran and Coin (1970:1(S5) used the combination Gastrotheca monticola argenteovirervs. l^oettger named argenteovirens in 1S92, whereas tlie name monticola dates from Barbour and Noble (1920). Moreover, the two taxa are con- siderably different and certainly not conspecific.

Gastrotheca plumbea (Boulenger)

Nototrcma plumheum Boulenger, 1882:417 [Holotype.— BMNH 78.1.25.22 (RR 1947.2..31.19) from Intac, Pro\incia Pichincha, Ecuador; Mr. Buckley collector].

Gastrotheca phimbeiim — Peters, 1955:346.

Diagnosis. — 1) Body robust, 62.3 mm in males, 68.0 mm in fe- males; 2) snout round in dorsal view, angular above and inclined anteroventrally in profile; 3) canthus rounded; 4) loreal region bare- ly concave; 5) tympanum vertically elliptical; 6) supratympanic fold weak; 7) subarticular tubercles on hand moderate, round; 8) super- numerary tubercles on hand small, round; 9) palmar tubercle bifid; 10) fingers webbed basally; 11) toes one-half webbed; 12) tarsal

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 15

fold round, extending one-third length of tarsus; 13) inner meta- tarsal tubercle elliptical, visible from above; 14) outer metatarsal tubercle absent; 15) subarticular tubercles on foot moderate, round; 16) supernumerary tubercles on foot small, conical; 17) discs round; 18) dorsal skin shagreened; 19) dorsum green with narrow bronze dorsolateral stripe; 20) facial area green with bronze canthal and labial stripes; 21) flanks brown; 22) dorsal surfaces of limbs green, mottled or not with tan; 23) posterior surfaces of thighs bronze-tan; 24) venter greenish yellow; 25) squamosal not exostosed, in moderately broad contact with maxillar\'; 26) tem- poral arcade incomplete; 27) prcvomers abutted medially; 28) transverse processes on eighth presacral vertebra strongly inclined anteriorly.

Gastrotheca phim])ea difi^ers from all other Andean Gastrofheca, except G. psychrophila, in having a uniformb' pigmented venter and a green iris. Like G. psycJtro})Jiiki, it also lacks dorsal mark- ings, but it diff^ers from G. psychrophila in having paler venter, a dorsolateral light stripe, a green, instead of brown dorsum, and a green, instead of copper, iris.

Variation. — The dorsum in\ariably is unmarked bright green to tannish green. The canthal, labial, dorsolateral, and anal stripes ar(> yellow. The loreal region, flanks, and posterior surfaces of the thighs are bronze, and the dorsal surfaces of the limbs are greenish bronze. The venter is \eIlowish tan with a greenish tint on the throat, and the ventral surfaces of the shanks arc blue. A diffuse blue spot is present in the grain; the iris is green.

Distribution. — Gastrotheca plnmhea is known from moderate elevations (1300-2350 m) on the Pacific slopes of the Cordillera Occidental in Ecuador (Fig. 3). Although the range of the species may extend northward into Colombia, it is doubtful if the species ranges into Peru; the dr\' \'alleys of the Huancabamba Depression probably arc a barrier to southward dispersal.

Remarks. — At Pilalo. Proxincia Cotopaxi. Ecuador, where there are remnants of cloud forest, individuals were found in bromeliads in trees and on a clift by day; males called from bromeliads at night.

Gastrotheca psychrophila new species

HoIotype.—KV 120760, an adult female, 61.0 mm, from the ridge between Loja and Zamora, 2850 m, 13-14 km E (by road) of Loja, Provincia Zamora-Chinchipe, Ecuador; obtained on 10 June 1968 b\' John D. Lynch.

Paratopotypes.—KV 120761, 10 June 1968, John D. Lvnch; 141586, 21 Mav 1971, Richard R. Montanucci; 142631-7, 21-23 July 1971, William E. Duellman.

Diagnosis. — 1) body depressed, 51.5 mm in males, 61.0 mm in

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

females; 2) snout pointed with vertical rostral keel, in profile rounded above and anteroventrally inclined; 3) canthus angular; 4) loreal region flat; 5) tympanum slightly higher than wide; 6) supratympanic fold heavy, curved downward behind tympanum; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand large, round; 9) palmar tubercle trifid; 10) fin- gers webbed basally; 11) toes slightly less than one-half webbed; 12) tarsal fold curved, two-thirds length of tarsus; 13) inner meta- tarsal tubercle elliptical, visible from above; 14) outer metatarsal tubercle low, round; 15) subarticular tubercles on foot large, round; 16) supernumerary tubercles on foot large, conical; 17) discs round; 18) dorsal skin shagreened; 19) dorsum imiform dark brown to grayish tan or dull green; 20) facial area colored like dorsum; bronze labial stripe in females; 21) flanks bluish black; 22) dorsal surfaces of limbs dark brown to grayish tan; 23) posterior surfaces of thighs bluish black; 24) venter grayish brown; 25) squamosal exostosed, in broad contact with maxillarv; 26) temporal arcade incomplete; 27) prevomers abutted medially; 28) transverse proc- esses on eighth presacral vertebra strongly inclined anteriorly.

Gastroiheca psijchrophUa diff"ers from all other Andean Gastro- theca, except G. phimheo, in having a dark venter. It also is like G. plumbea in lacking dorsal blotches, but G. psijchropluJa differs from G. plumhea in lacking a dorsolateral light stripe and in having a darker venter, usually a primarily brown dorsum (green in G. plumhea) and a copper iris (green in G. plumhea).

Variation. — Individuals of this species are capable of consider- able metachrosis. When frogs were found in bromeliads they were dark brownish black above and below; the flanks and posterior sur- faces of the thighs were dark bluish black. Later the dorsum changed to copper or 1:)ronze-tan with or without diffuse pale green blotclies or streaks. The flanks are orange-brown or dark brown. The axilla, groin, and hidden surfaces of the limbs are bluish gray or bluish purple. The lips are dull bronze, and the iris is copper with black flecks.

Distribution. — This species is known only from the ridge be- tween Loja and Zamora, Ecuador (Fig. 3). Most individuals have been found on the upper eastern slope between 2750 m and the crest at 2850 m.

Remarks. — The Loja-Zamora ridge is exceedingly wet; cold winds blow from the east. The vegetation near the crest consists of grasses and dense bushes. Large bromeliads are abundant on the ground and in the bushes. Adult G. psijchrophila were found in the bromeliads and imder rocks. Tadpoles were obtained from a grassy pond on tlic^ wc^st side of the ridge. Th(^ tadpoles are black and ha\e two upper and tlnce lower rows of denticles.

Etijmologij. — The specific name is from the Greek psycJiros,

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 17

meaning cold, and philos, meaning having an affinity for; the name is used in alhision to the chmate at the type locahty.

Gastrotheca riobambae (Fowler)

Hijla riohamhue Fowler, 1913:157 [Holotype. — ANSP 16161 from Riobamba, Provincia Chimborazo, Ecuador; S. N. Rlioads collector].

Htjla quitoe Fowler, 1913:159 [Holotype ANSP 1823S from Quito, Provincia Pinchincha, Ecuador; S. N. Rlioads collector].

Chorophihis oJivaceus Andersson, 1945:85 [Holotype. — NHRM 1965 from "Rio Napo, 400 m." ( =: ? Ranos, Provincia Tungurahua), Ecuador; William- Clarke Maclntyre collector].

Gastrotheca m[arsupiata] ecitadoriensis Vellard, 1957:43 [Nomen nuduni],

Gastrotheca riobambae — Duellman and Fritts, 1972:11.

Diagnosis. — 1) Body robust, 48.7 mm in males, 51.2 in females; 2) snout rounded in dorsal view and in profile; 3) canthus rounded; 4) loreal region shallowly concave; 5) tympanum round; 6) supra- tympanic fold weak, curved posteroventrally behind tympanmn; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand small, subconical; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-fourth webbed; 12) tarsal fold curved, extending full length of tarsus; 13) inner metatarsal tuber- cle elliptical, visible from above; 14) outer metatarsal tubercle ab- sent; 15) subarticular tubercles on foot ]arg(\ subconical; 16) super- numerary tubercles on foot small, round; 17) discs round; 18) dcn- sal skin shagreened, with scattered pustules, tubercular in tympanic region; 19) dorsum green or tan, usually with pair of large elongate dark spots; 20) facial area green or brown; canthal and labial stripes absent; 21) flanks green, blue, or tan with dark brown or black spots; 22) dorsal surfaces of limbs green or tan, usually with elongate dark mark on thigh and irregular blotclies or transverse bars on shank; 23) posterior surfaces of thighs dark brown with cream flecks; 24) venter cream with brown or gray mottling on chest and belly; 25) s(|uamosal exostosed in large individuals, in moderately broad contact with maxillary; 26) temporal arcade in- complete; 27) pre\'omers abutted or narrowly separated medially; 28) transverse processes on eighth presacral \'ertebra transverse or slightly inclined anteriorly.

Gastrotheca riohamlnie is like G. cavia in having short legs, a short snout, and a narrow interorbital distance, but it differs from G. cavia by having paired longitudinal dorsal marks, large spots on the flanks, and no dorsolateral light stripe. The dorsal c(^lor pat- terns of G. lojana and monticola are similar to that of G. riobambae, but both G. lojana and G. monticola have reticulated or plain flanks and dorsolateral light stripes. Gastrotheca plumhea and psij- chrophila have no dorsal markings and uniformly colored flanks; [he former has a dorsolateral light stripe.

Variation. — The preceding diagnosis is based principally on

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

topotypic material. Considerable variation, especially in coloration, obtains in this species. Five aspects of coloration are worthy of discussion (Table 2):

1. Dorsal ground color: The dorsum is either green or brown, varying to grayish tan in some individuals. The proportions of green to brown individuals is highly variable in local samples. At Papallacta on the high Amazonian slopes of the Cordillera Orien- tal, all individuals are green. The same is true at Guaranda on the Pacific slopes of the Cordillera Occidental. Approximately one-half of the frogs from the Cuenca Basin are green. At Riobamba, in the upper reaches of the Rio Pastaza drainage. 97 percent of the frogs are brown; at Baiios, lower in the Rio Pastaza valley, 89 percent are brown.

2. Dorsal markings: Most specimens have dark dorsal blotches. In green frogs these are usually darker green, but in some the blotches are brown. The same variation occurs in brown frogs, with the addition of green blotches that are dark brown peripherally and bordered or not by cream. The blotches usually are in the form of a pair of broad, irregular marks extending from the eyelid or occiput to the rump. In a few individuals the blotches are frag- mented into a row of spots; in others they are expanded so as to cover most of the dorsum. With the exception of the series from Riobamba, all large samples contain some individuals lacking dor- sal blotches. Plain individuals make up less than one-third of each sample, except that from Guaranda, in which one-half of the specimens lack blotches. Most Gastrotheca riohamhae lack definite dorsolateral light stripes, but these stripes are present in some speci- mens from Guaranda and the Cuenca Basin.

3. Thigh coloration: In most samples the posterior surfaces of the thighs are brown, gray, or tan (frequently with a green suffu- sion) with black flecks or small spots. In specimens from Riobamba and Guano the posterior surfaces of the thighs are tan or green with small cream flecks. The thighs are uniform dull blue in specimens from Papallacta and Biblian, bluish green from Cuenca, and blue with black flecks from Mulalo and Guaranda.

4. Flank coloration: The flanks are tan, green, gray, or blue, usually with black or dark brown spots. In specimens from Papal- lacta and Biblian the flanks are unifonn blue.

5. Ventral coloration: Specimens from Papallacta are uniform- ly gray below. In all other samples the belly is cream; the belly is marked with black, gray, or dark brown flecks, spots, blotches, or reticulations in all other samples, except those from the Cuenca Basin, in which the belly is uniformly cream.

In all spcx'imens th(^ iris is deep bronze to copper witli black reticulations; males in all samples have dark brown to gray vocal sacs.

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20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

There are some correlations between some of the aspects of coloration. Only three of the 37 plain specimens incoiporated in table 2 are brown. The color of the flanks generally is the same as the dorsum in green frogs, but in many brown frogs the flanks are green. Apparently the development of blue color on the flanks (most prevalent in the groin) is independent of dorsal color, but uniformly blue flanks are most prevalent in green frogs. The color of the flanks and posterior surfaces of the thighs usually are the same, but the markings on these surfaces are not necessarily the same.

When the variation in coloration is examined with respect to geography, two things are evident: 1) Samples containing only green frogs are from the outer slopes of the Andes ( Papallacta and Guaranda); all samples from the inner slopes and intcrandean val- leys contain both green and brown frogs. 2) Samples from the Cuenca Basin (Cuenca and Biblian) lack spots on the flanks, thighs, and venter. Other samples are more alike than any is to the four mentioned above. Because the population in the Cuenca Basin may be isolated genetically from more northern populations, it may represent a distinct taxon. Likewise, the populations on the outer Andean slopes may be isolated genetically from those in the inter- andean valleys. The populations at Guaranda and Papallacta are widely separated topographically with different phenons occurring in the intervening area.

If the variation in Gastrotheca riohamhae is examined with re- spect to genetic polymorphism, it can be conjectured that the poly- morphs at any given locality represent a balanced polymorphism resulting from selection for fitness to a particular environment (Levins, 1968). Data from the samples incoiporated in Table 2 were analyzed with respect to climatic variables (mean annual temperature, mean annual rainfall, minimum and maximum month- ly rainfall, and number of rainy days ) . Comparisons of percentages of green versus brown frogs, plain versus blotched frogs, and blue versus non-blue flanks and thighs with each of the climatic vari- ables resulted in no correlations.

Jameson and Pequegnat (1971) demonstrated that similar color polymorphism in Hijla regilla is correlated with seasonal and micro- ecological differences in vegetation color. The samples of G. rio- hamhae containing only green frogs are from areas where the vege- tation is in leaf and green throughout the year. The population containing the highest percentage of brown frogs (Riobamba) is from an area having sparse deciduous vegetation. At these and other localities, there was no planned sampling at different seasons; at those localities sampled at different seasons there is no significant difference in the frequency of different morphs in the samples.

Distribution. — Gastrotheca riohamhae has a broad geographic

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 21

and altitudinal range in Ecuador (Fig. 4). It occurs on the upper Pacific slopes of the Cordillera Occidental ( > 2600 m ) and Ama- zonian slopes of the Cordillera Oriental ( > 1800 m ) , and in inter- andean valleys ( > 2300 m ) as far south as the slopes of Cerro Tinajillas in Provincia Azuay. The species occurs at elevations of 3860 m in the Paramo de Apagua, 3960 m at Paso de Guamani, and 4135 m on Volcan Antisana. The species occurs in extreme southern Colombia, but specimens resembling G. riobambae from Bogota, Departamento Cundinamarca, and San Pedro, Departamento An- tioquia, apparently are not conspecific.

Remarks. — Gastrotheca riobambae occurs in a variety of habi- tats ranging from wet montane meadows to dry rocky hillsides. The species frequents ruderal situations — drainage and irrigation ditches, Agave, and corn fields. On cloudy or rainy days individuals are active, and males commonly call by day. Despite low tempera- tures (as low as 2-4° C), adults are most active at night. Tadpoles develop in still water. In large ponds the tadpoles aggregate in shallow water, but upon the slightest disturbance, they rapidly flee to deep water.

As noted in the preceding discussion of variation, some popula- tions currently assigned to Gastrotheca riobambae may be spe- cifically distinct. I suggest tliat biochemical and karyological in- vestigations might be fruitful approaches to the taxonomy of this complex.

DISCUSSION

Apparent evolutionary trends in the Andean marsupial frogs are confusing. Members of the Gastrotheca nuirsupiata group are the most terrestrial and live in what seem to be suboptimal environ- mental conditions — dry interandean valleys. On the other hand, members of the Gastrotheca argenteovhens group and some mem- bers of the GastrotJieca plumbea group are arboreal and live in what seem to be more optimal anuran environments — cloud forest and wet paramo.

Within the GastrotJieca plumbea group, two species (phimbea and psijchrophila) inhabit cool moist environments. Gastrotheca phimbea lives in arboreal and terrestrial bromeliads in cloud forest, and G. psijchrophila inhabits terrestrial bromeliads in wind-swept subparamo ( Fig. 5). The other species in the Gastrotheca phimbea group principally inhabit drier interandean valleys and Pacific slopes of the Andes. In these areas the frogs live in paramo. Agave, and cultivated fields; G. cavia inhabits bromeliads in scrubby sub- paramo (Fig. 6).

Among the members of the Gastrotheca phimbea group, G. riobambae is most like members of the GastrotJieca marsupiata

00.

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 5. Type locality of Gastrotheca psychroplula, ridge east of Loja Ecua- dor. Note terrestrial bromeliads in foreground.

group in having a relatively narrow head, least developed lateral flanges on the frontoparietals, and relatively small hands. The color pattern of G. riobombae is similar to that of G. peruana, the north- ernmost species in the Gastrotlieca maisupiata group. I consider the Gastrotheca phunbea group to have been derived from the Gastrotlieca marsiipiata group and G. riobambae to be the most primitive member of the Gastrotheca phimbea group. All other members of the group have more extensive cranial ossification and exostosis.

The major phyletic line in the Gastrotlieca phunbea group has dorsolateral light stripes and moderately long snouts. Two members (G. lojana and monticola) of this line retain the paired dorsal blotches of G. riobambae, whereas the dorsum is plain in G. phunbea, a species most like members of the Colombian Gastro- theca argenteovirens group. The nearly unicolor G. psijchrophila and the flecked G. cavia, each apparently represent independent derivatives from a G. riobam])ae-likc stock.

Vuilleumier (1971) documented Pleistocene changes in the flora and avifauna in the Andes. Her summary of geological, climatic, and biogeographic evidence demonstrates two glaciations in the Ecuadorian Andes. During glacial periods snow line was lowered as much as 700 m, and temperatures were depressed 4-11° C. The patterns of speciation and distribution of the Gastrotlieca phunbea group are compatible with Vuilleumier's palcobiogeographic hy-

MARSUPIAL FROGS OF THE ANDES OF ECUADOR

23

%%:\m

Fig. 6. Type locality of Gastrotheca cavia, Isla Pequena, Laj^una Cuicocha, Provincia Imhabura, Ecuador. Note bromeliads in scruhhy trees in middle of picture.

potliesis. GastrotJicca lojaiid and inonticohi arc rclictiial popula- tions of waiin-(lr\- interglacial periods now isolated in lower and drier areas than other menibers of tlie group. During climatic de- pression in glacial periods, populations were isolated in interandean basins and the outer slopes of the Andes; thus, G. riohamhiie, plumJ)eo, and psijchrophila differentiated in the interniontane val- l(>\s. Pacific slopes, and Amazonian slopes, rcspccti\-ely. A r\o- hamhac-likv stock apparenth' was isolated from more southern populations by uninhabitable environments in the Nudo de Mo- janda during a glacial period and differentiated into G. cavia. Con- ceivably, the differentiation of the six species occurred at the time of the first glacial period. If so, the differentiation within G. rio- Immhac and southward migration of G. lojana and monticola into northern Peru could be coincidental with the second glaciation. This proposed speciation model is similar to that suggested by Montanucci ( 1973 ) for the Andean microteiid genus PhoUdoJ)olus, a group of lizards inhabiting the same areas as Gastrotheca.

SUMMARY

The marsupial frogs of the genus Gastrotheca inhabiting the Andes and interandean valleys of Ecuador form a phylogenetie unit {Gastrotheca p]um])ca group) that is intermediate between the more southern GastrotJicca marsitpiata group and the more

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

northern Gastrothecu ar^enteovirens group. The Gastrotheca phimbea group is characterized by a supraor])ital flange on the frontoparietals and extensive exostosis, but no co-ossification, of the cranial roofing bones. Apparently all species in the group have free-swimming tadpoles.

The Gastrofheca phim])ea group contains six species: G. lojaua Parker, G. monticola Barbour and Noble, G. phimI)eo (Boulenger) and G. riohamhae (Fowler). In addition, two new species are named herein: G. ccwia from Laguna Cuicocha, Provincia Im- babura, Ecuador, and G. psycJiwphUa from the ridge east of Loja, Ecuador. Gastrotheca riohamhae is highly variable; some popula- tions may represent distinct species.

The Gastrotheca phimhea group seems to have been derived from the Gastrotheca marsupiata group, and G. riohamhae prob- ably is the most primitive member of the group. Speciation within the group evidently occurred through isolation of populations due to climatic fluctuation during the Pleistocene.

RESUMEN

Las ranas marsupiales del genero Gastrotheca cjue habitan los Andes y valles interandinos del Ecuador forman una unidad filo- genetica (el grupo Gastrotheca phimhea) que es un grupo inter- mediano entre el grupo Gastrotheca marsupiata del sur y el grupo Gastrotheca argenteovirens del norte. El grupo Gastrotheca plumhea se caracteriza por tener una protuberencia supraorbital en los frontoparietales y prominencias extensas pero no co-ossifica- cion de los huesos que forman la cubierta craneal. Parece que todas las especies del grupo tienen renacuajos acuaticos.

El grupo Gastrotheca phtm])ea tiene seis especies: G. h)jana Parker, G. monticola Barbour and Noble, G. plumhea (Boulenger), y G. rio])am])ae (Fowler). Ademas dos especies nuevas son nomi- nadas aqui: G. cavia de la Laguna Cuicocha, Provincia Imbabura, Ecuador, y G. psijchrophiki de la cordillera al este de Loja, Ecua- dor. Gastrotheca riohamhae es muy variable; algunas poblaciones pueden representar especies distintas.

El grupo Gastrotheca phimhea parece dirivarse del grupo Gas- trotheca marsupiata, y probablemente G. riohamhae es la especie mas primitiva de este grupo. La diferenciacion en el grupo ocurre evidentemente como resultado del aislamiento de poblaciones de- bido las fluctuaciones climaticas durante el Pleistoceno.

SPECIMENS EXAMINED

GiistrntJu'ca cavia

"Western Ecuador," BMNH 1860.6.16.124-125. Iiulialnira: Hacienda San Nicolas, 2000 m, UMMZ 92269, 92278-9, 92289-98; Ibarra, 2300 m, BMNH

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 25

(898.4.28.156; locality?); Laguna Cuicocha, 2890 m, KU 1.38216-20, 1.39136- 9, 139439 (tadpoles), 139440 (young), 143094, 14.35.37 (tadpoles), 148530- 42, 148543-4 (skeletons), 148545-7 (tadpoles), 148548 (young).

Gastrotheca lojana

Loja: Celica, 2130 ni, BMNH 1931.11.3.3-4; Loja, 21.50 m, BMNH 1931. 2.12.10-13, 19.33.6.24.18-44, 19.35. 11.3.26-.32, 1947.2.31.6-18, KU 120673-4; 2 km N Loja, 2100 m, KU 142846 (tadpoles); 5 km N Loja, 21.50 m, 1382.35- 6, 138237 (skeleton); 2 km E Loja, 220 m, KU 120675; 9 km E Loja, 2660 m, KU 121.387 (tadpoles); 2 km S Loja, CAS 93898; 3 km W Loja, 2150 m, KU 1.38233; 5.5 km W Loja, 23.30 m, KU 142603-8, 148549-51; 10 km W Loja, 2500 m, KU 1,382.34.

Zainora-Chinchipc: Zamora, 1000 m, BMNH 1933.624.45 (locality?).

PERU: Amazonas: Chachapoyas, 2340 m, KU 138238-41; 24 km WSW Leimebamba, 3.370 m, FSM 47216 (ID?). Cajamarca: Cajamarca, MJP 204. Fiura: Ayabaca, MJP 702 (2).

Gastrotheca monticola

Azuaij: Giron, 2240-2500 m, KU 138401-3. Loja: Saragino, 2500 m, KU 1.38404-9, 1.38410 (skeleton), 1.38769 (tadpoles), 141565, 142609-13, 142847 (tadpoles), 14856.3-8, 148.569-70 (.skeletons), 148.571 (tadpoles).

PERU: Cajamarca: Bella\ista, BMNH 1947.2.22.47-8, 1947.2.25.77-8; Querocotilla, MCZ ,5328-20. Piura: Iluancabamba, AMNH 7551, MCZ 5290- 3, 5296-7, .5299-,300, 5.302, 5304-7, 5,309, 5312-15, 5317. .5319, 5328-30, SMK 2677, UMMZ 55747.

Gastrotheca i)lunil)ca

Azuai/: Molleturo, 2,350 m, ZMB 300.57. Carclii: Atal, near San Gabriel, UMMZ 8.3655. Cotopaxi: Pilalo, 2320 m. KU 132413-22, 1.32423 (.skeleton), 142614. Pichincha: Intac, 1200 m. BMNH 1947.2.31.19.

Gast rotheca psych roph ila

Loja: 10 km E Loja, 2.570 ni, KU 1428.55 (tadpoles). Zamora-Chmchipc: 13-15 km E. Loja, KU 120760-2, 141.585 (skeleton), 141586, 141.59.5, 142631-

7.

GastrotJieca rlohanilyac

Province Unknown: No .specific locality, NLMIN 965, 6227-9 (8), 9.595; Andes, BMNH 58.7.25.21, .58.7.25.23, 58.7.25.25, ,58.7.25.27-8, 58.7.25.31-3; Western Ecuador, BMNH 60.6.16.17, 60.6.16.127. Azitay: Bestion, AMNH 13967; Cuenca, 2.540 m. CAS 85172, KU 120676-723, 129797-8, SMF 2669- 75, USNM 617.57-60, USNM-JAP 2345, 2347-8, 23.50; 6 km N Cuenca, AMNH 71588-600; 9 km N Cuenca, CAS 85339-40, 93884-94; 18 km N Cuenca, CAS-SU 21851; 4 km E Cuenca, 2540 m, KU 138587-613, 138622-3 (.skeletons), 138773 (tadpoles); 8 km SW Cuenca, ANLNH 71601-2; 8.8 km NW Cuenca, 2620 ni, KU 141583-4, 141594 (tadpoles); 9 km S Cumbe, 3300 m, KU 1,32536 (tadpoles); 10 km S Cumbe, ,3,350 m, KU 132392; 28.6 km S Cumbe, 3190 m, KU 1428,53 (tadpoles); 0.8 km S Cutcbil, 2535 m, KU 141582; 2.1 km S Cutchil, 2720 m, KU 141.572; 3.5 km S Cutchil, 2785 m, KU 141579-81; 8.5 km S Cutchil, KU 141577-8; Lago de Sarogucho, 20 km W Cuenca, CAS 94114; Laguna de Zurucuchu, 3200 m, KU 121.388 (tad- poles); Narihuina, MNHN 06-283; Rio Matadero, 8 km E. Cuenca, CAS-SU 21845-6; Rio Matadero. 9 km E Cuenca, CAS 94217 (tadpoles), 94218-22, CAS-SU 21847-8; Rio Matadero, 12 km E Cuenca, KU 129779-96; Sinicay, 2560 m, AMNH 17451-7, 17459-63, 17465-8, 17552, 17567. Bolivar: Guaran- da, 2640 m, KU 132403-12, 1.32531 (tadpoles), 132540 (young); 27.3 km

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

E Guaranda, 3800 m, KU 142850 (tadpoles); 2.5 km S Guaranda, 2650 m, KU 142616-9, 142851 (tadpoles), 142852, 148573; Guaranda-Riobamba road at Chimborazo border, 3700 m, KU 132541, 132542 (young). Canar: 3 km S Azogues, 2500 m, KU 138624-7, 138774 (tadpoles); Biblian, 2620 m, KU 141570-1, 141573, 142620-4, 147113; 8 km NW Biblian, 3420 m, KU 132537 (tadpoles); Canar, NHMW 6476, 6480, 2.8 km S Canar, 3150 m, KU 141574- 6; km 94, Guayaquil-Cuenca railroad, CAS 93899-900. Carchi: El Carmelo (El Pun), 2750 m, USNM-JAP 4946-7; Quebrada de Piedras, 20 km S Tulcan, 3400 m, KU 118120 (tadpoles); Tulcan, 3000 m, KU 117978-9, 118119 (tad- poles). Chimborazo: No specific locality, BMNH 1932.10.2.86; Guamote, USNM 33863; 1 km S Guano, 2500 m, KU 132354-89, 148581, 148582-4 (skeletons), 148592; Hacienda Alao, 15 km SE Pungala, 3100 m, KU 1.32543 (tadpoles); Laguna de Colta, 15 km SE Riobamba, 3400 m, NSNM-TAP 1728, 1730-3; Riobamba, 2780 m, ANSP 16161, KU 120732, MNHN 02-62(2), 02-350(2); 10 km N Riobamba, 2730 m, KU 138547-73, 138574-6 (skeletons); 15 km E Riobamba, 2600 m, KU 120725-31, 120758-9, 121389 (tadpoles); Rosario, NHMW 6485; San Juan, 3160 ni, KU 142615; 10 km W San Juan, 3160 m, KU 120724; 20 km SW Santa Rosa, 3700 m, KU 132348-9; Urbina Railroad Station, 3609 m, KU 1.32350-3; Volcan Chimborazo, USNM 103268- 74. Cotopaxi: El Porvenir, 2 km W Campamento Mariscal Sucre, 3620 m, KU 124167 (tadpoles); Guilo, 8 km E Pilalo, 3500 m, KU 1.325.38 (tadpoles); Laguna de Limpios, N base Volcan Cotopaxi, 3890 m, KU 122593; 4 km S Latacunga, USNM 164337; 6 km S, 7 km E Latacunga, 2750 m, KU 127082-4; Mulalo, 2980 m, KU 141566, 146263-4, 146749-61, 146762-3 (skeletons); Paramo de Apagua, 3860 m, KU 132390-1, 132535 (tadpoles); 24.3 km E Pilalo, .3750 m, KU 142848 (tadpoles); 11.3 km W Pujili, 3500 m, KU 141567; 38.3 km W Pujili, 3350 m, KU 142849 (tadpoles); Rio Pita, N base Cerro Ingaloma, 3780 m, KU 122594-9. Imhahura: Ibarra, 2300 m, AMNH 10569-71, BMNH 98.4.28.152-4, NHMW 6482-6; N slope Nudo de Mojanda, 3650 m, KU 132393; Nudo de Mojanda, 4 km S San Pablo, 3050 m, KU 132.394; Otovalo, 2.550 m, KU 68708, 138587-613, MNCN 333; Quebrada San Miguel, 1 km N Otovalo. 2560 m, KU 117980; Quiroga, 2500 m, KU 1138577-86, 148585-6 (tadpoles). Napo: Laguna Papallacta, .3350 m, KU 109169 (tadpoles); Papallacta, 31.30 m, KU 14.3095-102, 143538-40 (tad- poles), 14.3541, 148574-7; Rio Napo, 400 m, MNRM 1965 (locality?); Santa Barbara, 2625 m, USNM-JAP 4479, 4507; 1 km NW Santa Barbara, 2625 m, USNM-JAP 4487, 4491-3; 1 km SW Santa Barbara, 2625 m, USNM-JAP 4567; Volcan Antisana, 4135 m, AMNH 20127. Pastaza: Mera, 1140 m, AMNH .52852 (locality?). Pichincha: Intac, 1200 m, BMNH 78.1.25.20, FMNH 3607, NHMW 6481 (6) (locality?); Llave Pongo, AMNH 20140; Machachi, 2950 m, SMF 2667-8, UMMZ 47216; Paso de Guamani, 20 km E Pifo, 3960 m, KU 111626, 112316-7 (tadpoles), 127081, 1271.34 (tadpoles); W slope Paso de Guamani, 3940 m, KU 109170 (tadpoles), 109334-5; Quito, 2840 m, AMNH 20438-41, 20447-50, 20471-90, 60631; ANSP 182.35, CAS- SU 2274, 114.36-7, KU 94403, 111613-25, 112313-5 (tadpoles), 148416-28, 148578-80, 148587-91 (tadpoles), 148593-9, NMCN 156 (2), 1.58 (6), MNHN 34 (2), 1662 (2), 4878 (4), USNM 57804, USNM-JAP 1,570-2, 1574, 1576-7, 1.579, 1584, 1586-7, 1593, 1595, 1620-4, 1666, 1669, 1686, 2248-50, 2254, 2487, 2506-8; UZM 1474, 1477-8, 14424-93; between Rio Arturo and Taldadas, NE Cayambe, 3450 m, CAS-SU 8281; Rio Chiche, Valle de los Chillos, 2535 m, KU 152147-8; Santo Domingo de los Colorados, 500 m, AMNH 20147 (locality?). Tunmrahua: Ambato, 2700 m, KU 1207,3.3-40, 121390 (tadpoles), USNM 164302; Bailos, 1800 m, CAS-SU 5082, FMNH 28091-2, 173661-80, KU 99123, 99124-9 (skeletons), 991.30- 84, UIMNH 6.5,539-675, USNM-JAP 5834-6, 6010-12, 6014, 6019-20; Chambo Grande, 7.6 km SE Pelileo, 2340 m, KU 141,568-9, 142625-6, 146261-2; 10 km W Cotalo, 3300 m, KU 132400-2; 1 km W Juan Benigno Vela, 3080 m.

MARSUPIAL FROGS OF THE ANDES OF ECUADOR 27

KU 132395-6; Llanganati, near Rio Jorge, 3000 m, CAS-SU 17426-7; 4 km N Mocha, 3140 ni, KU 120757; 10 km SW Mocha, 3700 m, KU 120741-56; Pelileo, 2600 m, MNHN 03/211; 3 km SSW San Miguelito, 2620 m, KU 132399; 12 km SW Santa Rosa, 3400 m, KU 132397-8.

COLOMBIA: Ciindhmmarca: Bogota, BNLNH 1919.3.6.37 (locality?). Naiino: Cuaspud, TNHC 40564-5.

LITERATURE CITED

Anderssox, L. C.

1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke-Maclntyre and Rolf Blomberg. Arkiv Zool., 37A (2):l-88.

Barbour, T. and G. K. Noble

1920. Some amphibians from northwestern Peiii, with a revision of the genera Phyllobates and Telmatobius. Bull. Mus. Comp. Zool., 63:395-427.

BOULENGER, G. A.

1882. Catalogue of the Batrachia Salientia s. Ecuadata in tlie collection of the British Museum, ed. 2 London, xvi + 503 pp.

Cochran, D. M. and C. J. Coin

1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288: xii + 655 pp.

DuELLMAxV, W. E.

1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, l:xi -|- 753 pp.

DuELLMAN, W. E. and T. H. Fritts

1792. A taxonomic re\ iew of the southern Andean marsupial frogs ( Hy- lidae: Gastrotheca } . Occas. Papers Mus. Nat. Hist. Univ. Kansas, 9:1-37.

Fowler, H. W.

1913. Amphibians and reptiles from Ecuador, Venezuela, and Yucatan. Proc. Acad. Nat. Sci. Philadelphia, 55:153-176.

Jameson, D. L. and S. Peqltegnat

1971. Estimation of relative viabilit> and fecundity of color polymor- phisms in anurans. Evolution, 25:180-194.

Levins, R.

1968. Evolution in changing en\ ironments. Princeton Univ. Press, Princeton, ix + 120 pp.

Montannuci, R. R.

1973. Systematics and exolution of the Andean lizard genus PlioUdobohts (Sauria: Teiidae). Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 59:1-52.

Parker, H. W.

1932. Some new or rare reptiles and amphibians from soutlieni Ecuador. Ann. Mag. Nat. Hist., (10) 9:21-26. Peters, J. A.

1955. Herpetological tvpe localities in Ecuador. Rev. Ecuatoriana Ent. Parasit., 2 (3-4) :335-352. Vellard, J.

1957. Estudios sobre batracios andinos lY. El genero Gastrotheca. Mem. Mus. Hist. Nat. Ja\ier Prado, 5:1-47.

VUILLEUMIER, B. S.

1971. Pleistocene changes in the fauna and flora of South America. Sci- ence, 173 (3999):771-780.

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