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Herpetologica, 52(2), 1996, 282-289 C 1996 by The Herpetologists' League, Inc.

A NEW SPECIES OF PHOLIDOBOLUS (SQUAMATA: GYMNOPHTHALMIDAE) FROM THE HUANCABAMBA DEPRESSION OF NORTHERN PERU
TOD W. REEDER Department of Zoology and Texas Memorial Museum, The University of Texas, Austin, TX 78712-1064, USA
ABSTRACT: The occurrence of a previously undescribed species of Pholidobolus in the Huancabamba Depression represents the second species of the genus from Peru. The description of this species further increases the level of endemism of the Huancabamba Depression herpetofauna. The new species and P. annectens (and possibly P. anomalus) are proposed to represent a clade based of a single transparent palpebral disc and absence of a lateral on two synapomorphies-possession fold. The status of the poorly known P. anomalus is addressed. In addition, a single specimen of P. montium is reported from Colombia. This specimen seems to represent the first record of Pholidobolus from Colombia, even though P. montium long has been suspected to occur in extreme southern Colombia. A key is provided to the seven currently recognized species of Pholidobolus.

Key words: bia

Reptilia; Squamata; Gymnophthalmidae; Pholidobolus; New species; Peru; Colom-

THE microteiid lizard genus Pholidobolus is primarily known from the Andes and inter-Andeanvalleys of Ecuador, with five speciesoccurringat elevationsof 18004000 m (Hillis, 1985; Montanucci, 1973). A sixth species is known from southern Peru (Muller, 1923). In 1974, a Louisiana State University ornithology expedition made a transect across the Cordillera de Huancabamba in northern Peru, during which expedition members also secured a significant collection of amphibians and reptiles. Several new species of frogs and lizards have been described from this assortment. Included in this collection was a series of an undescribed species of microteiid lizard referable to the Andean genus Pholidobolus Peters 1862, as redefined by Montanucci (1973:31),by the possessionof the following external characters (osteological charactersnot examined):(1) snoutvent length ?66 mm, (2) tail accounting for '69% of total body length (64-71% in P. sp. nov.), (3) limbs pentadactyl, digits clawed, (4) body and tail cylindrical, neck not greatly constricted, (5) head distinctly flattened, (6) tympanum deeply recessed, (7) snout obtusely pointed, (8) nostrilpiercing nasal suture, (9) dorsalscales imbricate and weakly keeled to striated with some placoid (stronglykeeled in P. sp. nov.), (10)

dorsalscales in transverserows, (11) dorsal scales longer than wide, quadrangular to subhexagonal, (12) gular scales smooth, imbricate, becoming larger posteriorly, (13) two medial rows of widened gulars anterior to collar fold, (14) ventral scales smooth, imbricate, (15) ventral scales rectangular, in transverse and longitudinal rows, (16) tongue covered with imbricate, scale like papillae,and (17) hemipenes with minute, calcareous spines.
MATERIALS AND METHODS

I examined 94 preserved adult and juvenile specimens of Pholidobolusfrom Colombia, Ecuador, and Peru (Appendix I). Measurements were taken to the nearest millimeter using a ruler. Mean values for measurements and counts are reported as mean ? 1 SE. The drawings of the head and dorsalscalationwere made with a Wild M-7A microscope with camera-lucida attachment. Abbreviations for museum specimens follow those provided by Leviton et al. (1985).
SYSTEMATIC ACCOUNT

Pholidobolus huancabambae sp. nov. Holotype.-LSUMZ 27167 (Fig. 1), an adult male, from 33 km (by road) SW Huancabamba, elevation approximately

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FIG. 1.-Dorsal (upper),lateral (middle), and ventral (lower) aspects of head of holotype (LSUMZ 27167) of Pholidobolus huancabambae. Scale = 2 mm.

7000 ft (-2133 m) (west of Cruz Blanca; below summit of Cordillera Huancabamba), Departamento Piura, Peru, collected on 28 November 1974 by RichardThomas. Paratypes. -LSUMZ 27168-81 and 27188, from 33 km (by road) SW Huancabamba, elevation approximately 68007000 ft (-2072-2133 m) (west of Cruz Blanca;below summit of CordilleraHuancabamba), Departamento Piura, Peru; LSUMZ 27182-87, approximately 15 km (road) E Canchaque, approximately 5700 ft (- 1737 m) (= near Puente Fierro), DepartamentoPiura, Peru; KU 181977, from 15 km E Canchaque, elevation 1850 m, Departamento Piura, Peru. Diagnosis.-This species of Pholidobolus can be distinguished from all other

members of the genus, except P. annectens (Parker, 1930) and P. anomalus (Muller, 1923), by the presence of a single transparent palpebral disc in the lower eyelid. Those species not possessing a transparent palpebral disc have a lower eyelid composed of 3-6 opaque scales. Pholidobolus huancabambae can be distinguished from P. annectens by the following characters. (1) Dorsal scales elongate (Fig. 2) (shorterin P. annectens); (2) dorsal scales strongly keeled (striated to weakly keeled in P. annectens); (3) presence of a distinct, white labial stripe, extending from the second or third supralabial to the forelimb (generally absent in P. annectens; some specimens possessing a faint trace of a labial stripe that terminates anteriorto the anteriormargin of ear opening); (4) 32-35 dorsal scales counted from base of occipital to posteriormargin of hind limb insertion (39-44 in P. annectens); and (5) 20-24 ventral scales counted from posterior margin of collar fold to anterior margin of preanal scales (25-27 in P. annectens). Pholidobolus huancabambae can be distinguished from P. anomalus by the following characters. (1) Dorsal scales strongly keeled (smooth anteriorly and weakly keeled posteriorly in P. anomalus); (2) usually two supraoculars (three supraocularsin P. anomalus); (3) femoral pores absent (femoral pores present in P. anomalus); and, (4) smallersize, mean snout-vent length (SVL) of 47 mm (Table 1) [mean SVL of 58 mm (n = 3) in P. anomalus]. Superficially, Pholidobolus huancabambae also closely resembles P. prefrontalis Montanucci 1973. However, P. huancabambae can be distinguished from P. prefrontalis by having a undivided, transparentpalpebraldisc in lower eyelid (composed of 3-6 opaque scales in P. prefrontalis) and usually lacking prefrontalscales. Small, roundish prefrontals that do not touch each other are present in 17%of P. huancabambae;prominentprefrontalsthat often touch usually are present in P. prefrontalis. Description of holotype. -Dorsal tongue plicae scale like; infralingualplicae present; head scales smooth; frontonasal

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FIG. 2.-(Upper) Pholidobolus huancabambae (LSUMZ27178; paratype) and (lower) P. annectens (KU 121190). Dorsal view showing the difference in shape of the dorsalscales between these two species.

Scale = 2 mm.

wider than long, shorterthan frontal;frontal longer than wide, pentagonal; prefrontals absent;frontoparietalspaired, pentagonal; interparietal longer than wide, heptagonal; parietals slightly smaller than interparietal, hexagonal, and positioned anterolaterally to interparietal; occipitals three, median scale smaller than laterals; supralabialsseven; infralabialsfive; nostril piercing nasal suture; loreal present,

and touchingboththe frontonasal the first two supralabials; presubocular enlarged, two touchingloreal; supraoculars with the anteriormost firstbeingthe largest; superciliary enlarged, in contact with loreal; disc palpebral undividedand transparent; wider mostanterior postmental unpaired, than long, followed by two pairs of enlargedpostmentals touchingone another; twomedialrowsof widenedgulars, smooth; dorsalscaleselongularfold incomplete; gate, imbricate, arranged in transverse rows;dorsalsstronglykeeled, moderately and keeledon sidesof tail;dorsals ventrals separated thinbandof smallto granular by scalesanteriorly; lateralbody fold absent; ventrals smooth, juxtaposed,square to slightly wider than long; subcaudals smooth;limbs barely overlappingwhen adpressedagainst body; axillary region of scalesof forecomposed granular scales; limbsmooth weaklystriated, to mostlyimbricateexcept for small area of granular scales on ventral surface;subdigitallamellaeof forelimb one single,forming row, with a few lamellaebeing split into two scales;groin regioncomposedof smallto granularscales;scales on dorsal,ventral, and anteriorsurfacesof hind limb imbricate, thoseof the ventralsurfacesmooth, abruptly keeledtobecomingmoderately wards dorsal surface;scales of posterior surfaceof hindlimbsmooth, thoseof femoralregionsmallest (nearlygranular); subdigitallamellaeof hindlimbmostlysingle, with a few divided;femoraland preanal poresabsent;anteriorpreanalsfour; posterior preanalstwo, conspicuously larger than anteriorpreanals. Colorationof holotype.-In preservative (55%isopropanol, after formalinfixation), dorsumpale brown;dark brown middorsal stripepresentfromjustanterior to hind limbs,breakingup over proximal regionof tail; dorsolateral stripewhitish, terminating slightlyposterior forelimb of by gradinginto groundcolor, extending over anteriorly canthus rostralis (veryfaint, but discernible); distinctlabial stripe extending from second supralabial foreto limb; broad,dark brown stripe laterally; two lateral ocelli present between ear opening and forelimb,with one located

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TABLE 1.-Squamation

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and meristic data for Pholidobolushuancabambae.For characters1-5, the range is given (followed by the mean ? 1 SE). For characters6-8 only the number of individuals is given. "X" denotes characterstates for characters6-8 in the holotype.
Character Holotypecondition

1. Snout-vent length 2. Dorsalscalesbetween occipital and posterior margin of hind limb 3. Transversedorsal scale rows at midbody 4. Ventralscalesbetween collarfold and preanals

33-57 mm (47.1 mm ? 1.3) (n = 21) 32-35 (33.2 ? 0.2) (n = 23) 19-24 (20.2 ? 1.0)
(n = 23) 20-24 (20.6 ? 1.2) (n = 23)

47 33 22 23 8 X

5. Transverseventral scale rows at midbody 6. Supraoculars 2/2 3/2 2/3 3/3 7/7 7/6 6/7
8/7

7. Supralabials

8 (n = 23) 16 3 2 2 19 1 2
1

8. Infralabials

5/5
6/5 5/6

17
3 1

5/4 6/6

1 1

justabove forelimbinsertion;chin pale gray to bluish gray; belly blue-black; underside of tail with brown mottling. Variation. -Variation in squamation and meristic data are presented in Table 1. In 17% of the individuals, small prefrontals are present; however, they do not prevent contact between the frontal and frontonasal. The frontal is always longer than wide and ranges from pentagonal to heptagonal. The interparietal is pentagonal to heptagonal. One lateral occipital scale is divided in two individuals. Usually, there are seven supralabials; 13% of the specimens have only six on one side and one specimen has eight on one side. Usually, there are five infralabials;17%of the specimens have one additional infralabial on one side and one specimen has only four on the right side. An additional individual has six infralabialson each side. There are generally two supraoculars.However, 22% of the specimens possess an extra supraocular on one side (with the first and third supraocularsbeing subequal and the sec-

ond being the smallest) and 9% of the individuals have three supraocularson each side. The subdigital lamellae of the foreand hind limbs may be single or paired. The more distal lamellae are usuallysingle. Usually there are four anterior preanal scales and two enlarged posterior preanal scales. However, the lateral anterior preanal scales are slightly enlarged and elongated in 6% of the specimens. In 13% of the individuals, only two anterior preanal scales are present. Color in preservative.-Dorsum pale brown, with some specimens possessing a dark brown to black middorsal stripe, which often terminatesover the forelimbs. The middorsal stripe is also present just anterior to the hind limbs and breaks up over the proximal region of the tail. The dorsolateralstripe ranges from whitish to yellowish brown, sometimes edged with black, and terminates above or slightly beyond forelimb by grading into ground color. An anterior extension of the dorsolateral pale stripe is present in 45% of the

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individuals;when present, this anteriorextension is faint and often terminatesbefore reaching the tip of the snout. A distinct labial stripe is present, extending from second or third supralabialto forelimb. Some individuals possessa pale cream stripe below the broad brown lateral stripe, with the paler stripe extending from the forelimb to the hind limb. One or two lateral ocelli are present between ear opening and forelimb, one of which is always located justabove forelimb insertion.The chin and belly are pale gray to blue-black. Color in life.-(Richard Thomas, field notes), middorsum medium brown with a dark brown dorsolateral region. The dorsolateral stripe is cream to brownish buff and the labial stripe is cream. The throat and underside of tail are bluish gray. The venter is greenish gray to orangish brown. Etymology.-The specific name is Latin, an noun in the genitive singular case, and refers to the region of Peru (the Huancabamba Depression) for which all of the specimens of this species is known. This epithet also calls attention to the high degree of endemism of this region. Distribution and habitat.-The discovery of Pholidobolus huancabambae represents only the second species of Pholidobolus described from Peru. The first Peruvian species (P. anomalus) was describedfrom a single specimen from Cuzco over 70 yr ago (Muller, 1923). Pholidobolus huancabambae is geographically closer to all the Ecuadorian species of Pholidobolus than to the Peruvian P. anomalus. Pholidobolus huancabambae is known only from two localities in the HuancabambaDepression (=Huancabamba Deflection) of northern Peru, both on the Pacific slope of the Cordillera de Huancabamba. The two localities are separated by approximately 15 km. A map illustratingthe transect taken by The Louisiana State University (LSU)and The University of Kansas (KU) field parties across the Cordillera de Huancabamba is available in Duellman and Wild (1993:fig. 2). Specimensof Pholidobolus huancabambae were collected under rocks and trash piles in open fields that often lacked any herbaceous growth (R. Thomas, field

notes). The fields were surrounded by forest."Anotherspecimen was "subtropical found under a rock beside the road.
STATUS OF PHOLIDOBOLUS ANOMALUS (MULLER)

Pholidobolus anomalus was described by Muller (1923), based on a single specimen from Cuzco in southern Peru. The species remained known only from the holotype until rediscovered by Thomas H. Fritts (fide Montanucci, 1973). Examination by Fritts and Montanucci of the two new specimens of P. anomalus lead them to conclude that this species did not belong in Pholidobolus. As a result, Montanucci (1973) restricted the genus to the five Ecuadorianspecies. The data supportingthis conclusion and the proper generic allocation of anomalus were to be reported in a future paper by Fritts (fide Montanucci, 1973). Upon reexamination of the two new specimens of Pholidobolus anomalus (KU 134857-58), I do not concur with the conclusions of Fritts and Montanucci. Except for relative length of tail, the new KU specimens agree well with all the externalcharacters (except hemipenes, which were not observed; osteological characters not examined) described in the generic definition of Pholidobolus provided by Montanucci (1973). One specimen (KU 134858) possessesa tail accounting for 71% of the total body length, which is greater than that given in the generic definition (tail accounting for <69% of total body length). This appears to be a relatively trivial infraction of the generic definition. The other specimen (KU 134857) has a tail accounting for 68%of the total body length. I have not examined the holotype of P. anomalus. However, given in the type description,the holotype also agrees with the generic definition, except for the location of the nostril. Muller (1923) described the nostrilof the holotype as being in an upper divided nasal, implying a nasal with a horizontal suture. All other species of Pholidobolus and the two KU specimens of P. anomalus have the nostril situated in a vertical nasal suture. In the Ecuadorian species of Pholido-

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bolus and P. huancabambae,there is a dorsolateral pale stripe, continuing anteriorly over the canthus rostralis (faint and not extending to the tip of the snout or absent in some individuals). Both KU specimens of P. anomalus have a faint, but discernible, extensionof the dorsolateral pale stripe over the canthus rostralis (terminating at anterior end of first superciliary in KU 134858). Such a feature was also reported in the holotype description of P. anomalus (Muller, 1923). An extension of the dorsolateral pale stripe over the canthus rostralis appears to be a unique trait of Pholidobolus. Therefore, the presence of a dorsolateral pale stripe over the canthus rostralisand good agreement with the generic definition (Montanucci, 1973:31) leads me to conclude that P. anomalus is properly allocated to the genus Pholidobolus. The two KU specimens of P. anomalus, when compared to the type description, were found to agree in most details of squamationand coloration.However, there were also some notable differences. The KU specimens differ from the holotype in the following characters. (1) Prefrontals prominent and in contact, preventing contact of frontonasaland frontal (prefrontals small and widely separated, allowing contact of frontonasaland frontalin holotype); (2) first supraocularlargest (second supraocular largest in holotype); (3) second supraocular contacts superciliary border (second supraocular does not contact superciliary border in holotype); (4) nostril in vertical nasal suture (nostril in upper divided nasal in holotype); and (5) single transparent palpebral disc in the lower eyelid (opaque palpebraldisc, divided into two scales in holotype). These differences may reflect the existence of two different taxa. Alternatively,the differences may be the result of individual variation. Prefrontal size is known to vary within other species that possess them (i.e., P. affinis, P. huancabambae, P. macbrydei, and P. prefrontalis). Also, the presumed difference in the condition of the palpebral disc (transparent versus opaque) may be the result of differing interpretationsby Muller and myself. Nonetheless, because of the apparent differences between the KU

specimens and the type description, a reexamination of the holotype is warranted. Any potential changes in the taxonomic status of the KU specimens should wait until direct comparisonof these specimens can be made to the holotype.
DISCUSSION

Expeditions to the Cordillera de Huancabamba by KU in 1970, 1979, and 1991 and by Louisiana State University (LSU) in 1974 resulted in notable herpetological collections. Nine new anuran species have been describedfrom these series (reviewed in Duellman and Wild, 1993): four Eleutherodactylus (Leptodactylidae) (Duellman and Wild, 1993), two Gastrotheca (Hylidae) (Duellman and Trueb, 1988; Trueb and Duellman, 1978), two Phrynopus (Leptodactylidae)(Cannatella,1984; Lynch, 1975), and one Phyllonastes (Leptodactylidae) (Lynch, 1986). In addition, Cadle (1991) described five new species of lizards of the genus Stenocercus (Tropiduridae) from the Huancabamba Depression. While the LSU and 1979 KU field parties secured type material used in the descriptionsof three of the new species of Stenocercus, none of these new species is restricted to the Cordillera de Huancabamba.The HuancabambaDepressionwas proposed to be a major area of endemism within the Andean herpetofauna (Duellman, 1979; Duellman and Wild, 1993). The descriptionof this new microteiid species, as well as other recent frog and lizard species, from the Huancabamba Depression lends further support to this hypothesis. Using proteinelectrophoresis infer the to phylogenetic relationshipsamong the five Ecuadorianspecies of Pholidobolus, Hillis (1985) concluded that the species with the most derived characterswas P. annectens. This conclusion was corroboratedfurther by mapping seven qualitative morphological characters from Montanucci (1973) onto the phylogeny (Hillis, 1985). Two of these characters-single transparent palpebral disc and the absence of a lateral fold-were autapomorphic in P. annectens. Pholidobolus huancabambae also possessesthese two character states. If the

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polarities of these two characters are correct, then the autapomorphies proposed for P. annectens (Hillis, 1985) may be synapomorphies uniting P. huancabambae and the geographically proximate P. annectens as sister species. Pholidobolus anomalus also possessesa single transparent palpebral disc and is polymorphic for lateral fold condition (lateral fold absent in KU 134857; lateral fold present in KU 134858); therefore, P. anomalus may belong to an exclusive clade containing P. annectens and P. huancabambae. However, P. anomalus also exhibits the putatively primitive condition (i.e., three supraoculars, prefrontals present, femoral pores present,and lateralgranulespresent) in four of the seven characters listed by Hillis (1985), suggesting a relatively basal position for P. anomalus. In addition to describing a new species of Pholidobolus from northern Peru and reporting on the status of P. anomalus, I take this opportunityto reportupon a specimen of Pholidobolus from Colombia in the American Museum of Natural History. The specimen (AMNH 131232) was collected at "Narinio: San Luis, Finca La Quinta, near Ipiales" on 2 October 1975 and can be identified as P. montium (Peters, 1862). Pholidobolus montium is the northern most species of Pholidobolus in Ecuador,and Montanucci (1973) expected this species to be discovered eventually in extreme southern Colombia. However, I am unaware of any reports of this species having ever been taken previously in Colombia. Therefore, AMNH 131232 may represent the first record of Pholidobolus from Colombia.
KEY TO THE SPECIES OF PHOLIDOBOLUS

3.

4.

5. 6.

7.

anteriordorsal Usuallytwo supraoculars; 3 scales striated to strongly keeled -----------------.... Dorsal scales elongate (Fig. 2), strongly keeled .... ...... P. huancabambae Dorsal scales not elongate (Fig. 2), striP. ated to weakly keeled ..-.....----- annectens Three supraoculars;prefrontals present P. ....... ...... affinis ......................... Two supraoculars, subequal;prefrontals 5 present or absent..... Prefrontalspresent ..............6.........6 Prefrontalsabsent.... 7 Dorsolateralpale stripe distinct, extending to tip of snout; femoral pores absent in both sexes; head not distinctly broaderin males than in females;sides of neck and tail lacking red stripe in . .... .. P. prefrontalis males Dorsolateralpale stripe distinct, not extending to tip of snout; femoral pores usuallypresentin males;head distinctly broader in males than in females; sides of neck and tail with red stripe in males.. .P. macbrydei Dorsolateralpale stripe distinct, not extending to tip of snout; femoral pores usuallypresentin males;head distinctly broader in males than in females; sides of neck and tail with red stripe in males.P. macbrydei Dorsolateralpale stripe distinct, extending to tip of snout; femoral pores absent in both sexes; head not distinctly broaderin males than in females;sides of neck and tail lacking red stripe in . males P. montium

Acknowledgments.-Ithankthe followingindividualsfor loanof specimens (institutional abbreviationsas in Levitonet al., 1985):W. Duellman and J. Simmons (KU)and D. A. Rossman I (LSUMZ). thankD. Cannatella, Fritts, Frost, Hillis,D. T. D. D. R. and Kizirian, Montanucci, D. Rossman comfor ments andcriticisms various on of drafts thismanuof script.
LITERATURE CITED
CADLE,

The seven species of Pholidobolus can be identified by the following key, which is modified from Montanucci (1973) to include P. huancabambae and P. anomalus: 1.

2.

J. E. 1991. Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northernPeru: New Species and comments on relationshipsand distributionpatterns. Proc. Acad. Nat. Sci. Philadelphia 143:1-96. CANNATELLA, D. C. 1984. Two new species of the Lower eyelidhavinga singletransparent leptodactylid frog genus Phrynopus, with comdisc palpebral 2 ments on the phylogeny of the genus. Occas. Pap. Lowereyelid having3-6 opaquescales Mus. Nat. Hist. Univ. Kansas113:1-16. ........................................................................................4.................. W. E. 1979. The herpetofaunaof the ... 4 DUELLMAN, Threesupraoculars; anterior dorsal scales Andes.Pp. 371-459. In W. E. Duellman (Ed.), The smooth .. P. anomalus South American Herpetofauna:Its Origin, Evolu-

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tion, and Dispersal. Univ. KansasMus. Nat. Hist. TRUEB, L., AND W. E. DUELLMAN. 1978. An extraordinary new casque-headed marsupial frog Monogr.7:1-485. (Hylidae: Gastrotheca).Copeia 1978:498-503. DUELLMAN, W. E., AND L. TRUEB. 1988. Cryptic species of hylid marsupial frogs in Peru. J. HerAccepted: 9 February 1995 petol. 22:159-179. Associate Editor: Linda Trueb DUELLMAN, W. E., AND E. R. WILD. 1993. Anuran de Huancabamba, amphibiansfrom the Cordillera northern Peru: Systematics, ecology, and biogeography.Occas. Pap. Mus. Nat. Hist. Univ. Kansas 157:53. HILLIs, D. M. 1985. Evolutionary genetics of the Andean lizard genus Pholidobolus (Sauria:GymAPPENDIX I nophthalmidae):Phylogeny, biogeography, and a Specimens Examined comparison of tree constructiontechniques. Syst. Pholidobolus affinis.-ECUADOR: Chimborazo: Zool. 34:109-126. 15 km E Riobamba,2600 m, KU 121155-60. LEVITON, A. E., R. H. GIBBS,JR., E. HEAL, AND C. Pholidobolus E. DAWSON. 1985. Standardsin herpetologyand annectens.-ECUADOR: Loja: 2 km ichthyology: Part I. Standard symbolic codes for E Loja, 2200 m, KU 121181-90. Pholidobolus anomalus.-PERU: Cuzco: Cuzco, institutional resourcecollectionsin herpetologyand Machu Pichu, 2000 m, KU 134857-58. ichthyology. Copeia 1985:802-832. Pholidobolus macbrydei. -ECUADOR: Azuay: LYNCH, J. D. 1975. A review of the Andean leptodactylidfrog genus Phrynopus.Occas. Pap. Mus. Contrayerbas,W of Cuenca, AMNH 23466. Caniar: 15 mi N Cafiar,AMNH 24345. Morona Santiago: 8 Nat. Hist. Univ. Kansas35:51. . 1986. New species of minute leptodactylid km S Cutchil,3040 m, KU 121249-55. Zamora-Chinfrogs from the Andes of Ecuadorand Peru.J. Her- chipe: Sabarilla(=Sabanilla?),Rio Zamora, AMNH 18309. petol. 20:423-431. MONTANUCCI, R. R. 1973. Systematicsand evoluPholidobolus montium.-COLOMBIA: Narinio: tion of the Andeanlizard genus Pholidobolus(Sau- San Luis, Finca La Quinta, near Ipiales, AMNH ria: Teiidae). Misc. Publ. Mus. Nat. Hist. Univ. 131232.ECUADOR: Imbabura: Ibarra,AMNH5282, Kansas 59:1-52. 13486-9; Lago Cuicocha, KU 142874-79; San Pablo vic. HostariaCusin, AMNH 124045. Napo-Pastaza: MULLER, L. 1923. Neue oder seltene reptilien und batrachierder ZoologischenSammlung des bayr. Banos, Rio Pastaza, AMNH 60611-14. Pichincha: Staates.Zool. Anz. 57:49-61. Quito, AMNH 28771-82; 3 km N Ot6n, AMNH PARKER, H. W. 1930. Two new reptilesfrom south118883-4. ern Ecuador.Ann. Mag. Nat. Hist. 10:568-571. Pholidobolusprefrontalis. -ECUADOR: Azuay:6 PETERS, W. H. C. 1862. Uber Cercosauraund die km N Cuenca, nr. Rio Matadero,AMNH 91827-32. mit dieser Gattung verwandten Eidechsen aus Siu- Chimborazo:3.3 km S Tixan, 2885 m, KU 141094damerica. Abh. Akad. Wiss. Berlin 1862:165-225. 99.

DATE OF PUBLICATION

Herpetologica, Vol. 52, No. 1, was mailed 1 March 1996.