¡ournal 01Herpetology,Vol. 36, No. 3, pp. 349-355, 2002Copyright 2002 Society lar the Study 01 Amphibians and Reptiles

A New Species of Echinosaura (Gymnophthalmidae) from Ecuadorand Colombia with Comments on Other Members of the Genus and

Teuchocercus keyi

THOMAS H. FRITTS/,2 ANA ALMENDÁRIZ,3 AND SISSI SAMEC4

'USo Geological Survey, Midcontinent Ecological Science Cente/; 4512 McMurry Avenue,Fort Collins, Colorado 80525, USA; E-mail: frittstom@cs.com

3Escuela Politécnica Nacional, Departamento de Ciencias Biológicas, Apartado 2759, Quito, Ecuador

4Naturhistorisches Museum Wien, Burgring 7, A-1O14, Wien, Austria

ABSTRACT.-A new species of the genus Echinosaura is described from the Pacific drainages of northernEcuador and adjacent Colombia. Evaluation of variation among congeners justifies their recognition asdistinct species instead of subspecies as previously proposed. The new lizard occurs in close proximity toEchinosaura horrida and Teuchocercus keyi.

RESUMEN.-Se describe una nueva especie de lagartija del género Echinosaura de los drenajes Pacificosen el norte de Ecuador y el sur de Colombia. Evaluación de variación entre las formas nombradas en elgenero se justifica reconocimiento como especies distintas, y no como subespecies. La especie nueva ocurrecerca de las distribuciones de Echinosaura horrida y Tellchocerclls keyi.

Lizards of the genus Echinosaura are distrib-uted from Panama to the Pacific drainages ofEcuador. Four Domínate forros have been de-scribed, but only a single species with threesubspecies was recognized by Uzzell (1965).Subsequent to Uzzell's review, Teuchocercuskeyi,presumably related to Echinosaura, was namedby Fritts and Smith (1969). Although consider-able material exists from Ecuador assignable toa single taxon (Echinosaura horrida), other laxa ofthis group remain sparsely documented. Thepresent review was stimulated by our attemptsto identify specimens of Echinosaura from the ex-treme northern regions of Esmeraldas and Car-chi Provinces in Ecuador and adjacent afeas ofColombia. We interpret these lizards as an un-described species of the genus Echinosaura. Ad-ditional material of Echinosaura and Teuchocercusis now available to allow reexamination of mor-phological variation in the groups and a reeval-uation of taxonomic and distributional relation-ships. The cephalic scutellation of this group isextremely variable both within and among laxa.This variation must be interpreted cautiously inassessing relationships within these lizards.

MATERIALS AND METHODS

We have examined nearly all specimens of thegenera Teuchocercus and Echinosaura available(Appendix 1) in the collections of the NationalMuseum of Natural History (USNM), AmericanMuseum of Natural History (AMNH), EscuelaPolitecnica Nacional, Ecuador (EPN), Museum

2 Corresponding Author.

of Comparative Zoology, Harvard (MCZ), Uni-versity of Kansas Museum of Natural History(KU), Academy of Natural Sciences, Philadel-phia (ANSP), and the Naturhistorisches Muse-um Wien (NMW). We have alBo used data sum-marized by Uzzell (1965) to supplement sam-pIes available to uso

In an attempt to clarify the comparison of themembers of this group and other poorly knownmicroteiid groups, we have adopted a terminol-ogy for scutellation patterned after the defini-tions used by Peters (1964), which is somewhatat variance with that of Uzzell (1965), Ofteda](1974), and other recent workers (e.g., Harris,1994; Kizirian, 1996). Acknowledging that thecephalic scutellation of lizards in the generaEchinosaura and Teuchocercus is variable and de-viales from the typical gymnophthalmid pat-terns, we have attempted to avoid confusion byproviding explicit descriptions to supplementllames of scales if the identity of scales is notclear. Although many gymophthalmid lizardshave only one scale or an additional pair ofscales (typically called frontonasals and pre-frontals, respectively) between the rostral andtIle' frontal, Teuchocercus and Echinosaura speci-mens reported here have three series of scalesin this regían warranting careful description ofscale positions and homologies. We have usedthe following definitions of the cephalic headscales.

Internasals.- The scales located between therostral, nasals, and frontonasals (Peters, 1964:170). In the genus Echinosaura the internas almar be paired or single. Uzzell (1965) used the

350 T. H. FRITTS ET AL.

terms internasal and frontonasal interchange-ably in describing the scale immediately poste-rior to the rostral. This inconsistency is a poten-tial source of confusion for students of thisgroup of lizards.

Frontonasals.- The scale or scales located be-tween the internasal, frontal, and loreal. In liz-ards with paired prefrontals, the frontonasalmar be bounded posteriorly by the prefrontalsor by the prefrontals and frontal instead of thefrontal only (Peters, 1964:132). Normally thefrontonasal is confined to the dorsal surface ofthe snout and largely anterior to the interorbitalafea.

Prefrontals.-Paired or single scales betweenthe frontonasal and frontal, in Borne lizards,which occupy the anterior margin of the inter-orbital afea. Prefrontals are usually absent inthe genus Echínosaura but are variably presentin Teuchocercus and Borne Echínosaura. In distin-

guishing prefrontals from frontonasals, prefron-tals are nearly as wide as long because they oc-cupy the are a where the head expands to ac-commodate the orbits, whereas frontonasalsmar be longer than wide and primarily con-fined to the snout anterior to the interorbitalafea.

Frontals.-Normally an unpaired scale orscales occupying the space between middle andanterior halves of the semicircles defined by theorbits (Peters, 1964:131). In Borne individuals ofEchínosaura, the frontal can be divided longitu-dinally (paired) or transversely into anteriorand posterior segments. Distinguishing a frag-mented posterior portion of the frontal from thescales normally termed frontoparietals wouldbe difficult unless geographic markers related tothe orbit were used, and OUTdescriptions havebeen expanded to reflect this need.

Frontoparíetals.-Paired or multiple scales ly-ing posterior to the frontal and between theposterior halves of the semicircles and anteriorto scales in the parietal and interparietal re-gions. The frontoparietals of Echínosauraare var-iable in size and number and consequently aredifficult to distinguish from fragments of thefrontal and the similarly variable parietal scaleslocated posterior to them. As noted above, thedefinition of these scales is problematic, and wehave limited our description of frontoparietalsin taxa discussed in this study to "the scalesbordering the posterior margin of the frontal"even though frontoparietals and parietals marbe clearly identified in other groups of gymno-phthalmid lizards.

Interparíetal and Paríetals.-Unpaired medialand paired scales, respectively, in the parietalregion of the head are small and irregular inEchínosaura and Teuchocercus. They are present

and easily identifiable in species of Neustícurusexcept in Neustícurus apodemus.

RESULTS

Status ofEchinosaura spp.-Uzzell consideredtwo specimens from Paramba, Prov. Imbabura,Ecuador, intermediate between horrída and pal-merí because they had the paravertebral crestseparated in two distinct rows, two internasals,snout short, frontal short, Brilles on the dorso-lateral region of the neck not greatly elongate,and Brilles of lateral neck short. The presence oftwo internasals in conjunction with a shortsnout and short frontal prompted us to hypoth-esize that these specimens represented Teucho-cercus keyí, and this identification was confirmedby examination of the specimens by one of us(SS). Thus, no intergrades are known from theParamba locality.

Uzzell considered two specimens from IslaGorgona, Colombia as intergrades between E.horrída and Echínosaura palmerí because they hadparavertebral rows in contact and a single in-ternasal (like horrída), but they differed in thedegree of development of Brilles on the neckand in the form of the frontoparietal, character-istics which are difficult to assess and which wefind to vary within single localities. We exam-ined an additional specimen from Isla Gorgona(MCZ 160153) and find it to agree with E. hor-rída in characteristics of the paravertebrals andinternasal. Thus, we identify the lizards fromIsla Gorgona as a previously unreported popu-lation of horrída in Colombia and note the lackof evidence of intergradation between E. horrídaand E. palmerí.

No intergrades between E. palmerí and Echí-nosaura panamensís were noted by Uzzell (1965),but he considered "the differences between

them to be slight." We noted a striking differ-ence in the structure of the tail in these taxa.The tail of palmerí is similar to that of horrída inhaving longitudinal rows of enlarged scales inthe paravertebral, dorsolateral, and lateral re-gions of the tail. The rows are made of series ofenlarged scales increasing in size from the an-terior border of each autotomic segment to theposterior border of the segment and then re-peating the pattern on the successive segments.The tail of horrída (illustrateq.by Fritts andSmith [1969:fig. 2]) is typical of the conditionexisting in palmerí. The tail of panamensís lackscontinuous longitudinal rows because the scaleson the anterior margins of each autotomic seg-ment are small in size and similar to adjacentscales. At the posterior margin of the segments,enlarged scales exist in the paravertebral, dor-solateral, and lateral regions, but the successive-ly smaller scales anterior to these enlargedscales grade into the small scales anteriorly, dis-

NEW SPECIES OF ECHINOSAURA

rupting the rows. Most specimens of panamensísalgo differ from palmerí and horrída in having thefrontal divided transversely and irregularlyfragmented. One specimen from Panama(USNM203665)has an entire frontal but retainsthe caudal morphology typical of panamensís.

Thus, we find evidence to suggest that thethree taxa recognized by Uzzell (1965) as sub-species are test recognized as distinct species.The recognition of E. panamensís as distinct fromE. palmerí is tentative based on the materialavailable to liS, but careful revision of these liz-ards with additional material will be needed tofurther clarify their status.

Dístríbutíonal Relatíonshíps.- To date none ofthe three taxa has been reported to occur insympatry, but the possibility exigís near the Ec-uador-Colombia border where E. horrída isknown from lowland localities and E. palmeri isknown from higher elevations.

Echinosaura horrída and T. keyí occur in sym-patry at El Placer, Esmeraldas Province, Ecua-dor; USNM 196093 and USNM 196094 repre-sent the two taxa, respectively. They algo occurin sympatry at Mataje, Esmeraldas Province(NMW 31999 and NMW 32001:1, respectively)and at La Florida, Pichincha Province (KU218389-395 and KU 218397-98, respectively).The specimens of Teuchocercus from Pichincha,Carchi, and Esmeraldas Provinces (listed in theSpecimens Examined) augment our knowledgeof fue distribution of the species, which untilrecently was known only from the holotype andparatype of the species collected at the type 10-cality in Pichincha Province of Ecuador. Teucho-cercus keyi is readily distinguishable from Echi-nosaura in having a markedly shorter snout,wider head, prefrontal scales irregularly pre-sent, and in adults, the conical caudal scales not-ed in the original diagnosis by Fritts and Smith(1969). It algo has granular scales covering thetympanum of adult specimens, but the smallscales are not visible on the central part of thetympanum in small juveniles.

A morphologically distinctive and unde-scribed taxon of Echínosaura is known to occurin clase proximity to (but not necessarily insympatry with) horrída in Esmeraldas Provinceof Ecuador, near the Colombian border in theMataje Region where Teuchocercusis now knownto occur. We name this species in honor of Gus-tavo arces V. in recognition of his contributionto the knowledge of the vertebrates of Ecuadorand his guidance of students and other investi-gators ayer several decades.

Echínosaura orcesí sr. nov.Figure 1

Holotype.-AMNH 109822; adult male collect-ed by C. W. Myers (CWM 11457); Type Locality:

351

Colombia: Valle: approximately 13 km west ofDagua, Rio Anchicaya drainage, 820 m.

Paratypes.-EPN 2574; adult female; Ecuador:Provincia Esmeraldas: Centro Comunal Mataje,Reserva Awa, near Rio Mataje at point where itparallels Rio Mira, 01°12'04"N, 78°33'42"W, 250m; collected by Ana Almendáriz on 6 February1988. NMW 32000:1; adult male; Ecuador: Prov-incia Carchi: San Marco, 670 m; collected by E.Samec in 1988. NMW 32000:2; adult female; Ec-uador: Provincia Carchi: San Marco, 700 m; col-lected by E. Samec in 1988.

Díagnosís.-A species of the genus Echinosaurawith an extremely long snout, unpaired inter-nasal, reduced numbers of supralabials and in-fralabials, enlarged caudal scales arranged incontinuous transverse rings spanning the dorsaland lateral surfaces of the tail at the posteriormargin of each autotomic segment; and elevatednumbers of subdigital lamellae on the fourthtoe. In contrast to other members of the genus,Echínosaura orcesí lacks continuous rows of slightto moderately enlarged scales in the paraverte-bral afea of the body.

Descriptíon (basedon holotype).-Scales of headfUgase with numerous longitudinal ridges;snout elongate, narrow; head markedly wider atand posterior to orbital region. A single inter-nasal scale, longer than wide, with prominentlongitudinal rugosities, medial groove, and aconvex posterior margin, bordered posteriorlyby a pair of frontonasals and laterally by pos-terior extension of rostral and anterior half ofnasal scales. Frontonasals only slightly longerthan the sum of their widths, marked with lon-gitudinal rugosities, bordered posteriorly byconvex anterior margin of frontal scale at levelof presupraocular scales. Frontal narrowingposteriorly at middle of orbits, with a convexposterior margin at suture between first andsecond supraoculars contacting paired scales infrontoparietal region. "Frontoparietals" bor-dered laterally by second supraoculars and bymost medial of a series of enlarged scales bor-dering posterior margin of orbits. Scales poste-rior to "frontoparietals" and on regí of posteriorhead small, irregular, flattened in medial occip-ital region, somewhat convex and tubercular intemporal region.

. Nasal scale about twice as long as high withnóstril in middle; groove extending from nostrilto border of internas al. A single scale slightlylonger than high separates nasal from the an-terior angle of orbit. Three supralabials, with su-ture between first and second just anterior toposterior edge of nasal, suture between secondand third just anterior to middle of eJe. Threeinfralabials. Mental extending further posteriorthan rostral; a single postmental bounded pos-teriorly by a pair of enlarged scales that laterally

352 T. H. FRITTS ET AL.

FIC. 1. Dorsal and ventral views of AMNH 109822, the holotype of Echinosaura orcesi.

contact second infralabials. All other scales onruin and guIar regían irregularly arranged,small. Tympanum small, lacking scales on sur-face but having tiny white flecks.

Scales of neck and body small, granular, notin transverse or longitudinal rows, intermixedwith enlarged tubercular scales with longitudi-nal keels. Enlarged scales often arrangedobliquely in short series of 3-8 scales formingirregular chevrons on dorsal field of body andbroken undulating rows of scales along dorso-lateral body. No continuous vertebral or para-vertebral rows of enlarged scales. Chevrons ex-tend anta base of tail only; tail incomplete, re-generated posterior to basal tour autotomic seg-ments. Scales on original tail small, irregularlyarranged except at posterior margin of each au-totomic segment where enlarged keeled scalesare arranged in transverse series extendingfrom paravertebral afea laterally to longitudinalcrease on lateral surface of tail. Scales anteriorto the enlarged scales only slightly enlarged rel-ative to granular caudal scales covering most ofeach caudal segment. Scales on regenerated seg-ment of tail granular, uniform in size, irregular-ly arranged. Ventral scales loosely arranged in

transverse rows between pectoral and pelvic re-gions and in eight longitudinal rows at mid-body. Ventrals smooth, except for traces of keelson some scales at lateral margin of ventero Digitsof hands and feet five, with prominent claws.Dorsal surface of upper forearm with enlarged,keeled scales; dorsal surface of lower forearmwith smaller, keeled scales. Subdigitallamellaeon fourth finger 20; lamellae on fourth toe 30.Femoral pares 15-15 extending anta pelvic are aswith 4-5 scales separating right and left seriesof pares. About 44 scales on midline of venterfrom collar fold to vent.

Color in Life.-Dull brown with vague blackdorsal bands (on neck, body, and base of tail).Venter dull yellow with black transverse bandsacross ruin; a weak suffusion of white (individ-ual granules) under head posterior to ruinmarkings. Iris brown with heavy gray suffusion.Anterior part of tongue black, posterior part un-pigmented. Description from field notes of C.W. Myers 28 January 1973.

Color in Preservative.-Black transverse mark-ings on ruin outline off-white markings coincid-ing with sutures between infralabials. Whitemarks at suture between second and third in-

NEW SPECIES OF ECHINOSAURA

fralabials extend dorsally to suture between sec-ond and third supralabials and to ventral mar-gin of orbit. Tip of mental off-white. Lower eye-lid unpigmented, translucent. Six irregularblack blotches on each side of ventrolateral bodywith trace of white spot between consecutivepairs of dark blotches. A pair of smaller darkblotches and corresponding white spot is evi-dent anterior to larger blotches, posterior to pec-toTal regían. A broad white band at base of tail.Diffuse white markings on sutures of supraocu-lar scales.

Notes on Variation.-The tour specimens athand are adults, two females and two males.The number of femoral pares differs betweenthe sexes: males with 14-15 pares on each side,females with 5-6 on each side. The width of thehead is greater in the holotype and less mark-edly so in the other male (MNW 32000/2); thehead of females is narrow. The three specimenswith complete tails have long tails (2 X snout-vent length, SVL). Ratios of SVL/Taillength forthe two females were 0.50-0.51 and 0.45 for onemaleo Males were 69-70 mm SVL, females 76-81 mm.

Presence of prefrontal scales between thepaired frontonasals and the unpaired frontal isvariable with two specimens lacking prefron-tals, one having a pair of prefrontals, and onehaving a single prefrontal on only one side withthe frontal and frontonasal in contact on theother side. Supralabials are 3-4 and infralabials2-3. Subdigital lamellae on the fourth toes are30-31.

Arrangement of enlarged tubercles on neck,body, and head varies slightly. All individualslack discrete rows of enlarged scales in the ver-tebral or paravertebral afeas. Enlarged tuberclesin the dorsal field are arranged in irregularlypaired diagonal series that commonly producetubercular chevrons. Three individuals have tu-bercular scales along the dorsolateral line ar-ranged in a broadly undulating nearly completetubercular crest. The type differs in having thescales in the dorso lateral afea arranged in dis-continuous and obliquely oriented series.

Color pattern of the paratypes is more diffusethan in the holotype. All animals have a prom-inent off-white band across the base of the tail,conspicuous white marking across the orbit andinterorbital afea, and the black-and-white mark-ings on the infralabials and chino Only the typehas conspicuous dark blotches in the ventrolat-eral regían of the body.

DrscussrON

Comparisonswith Other Echinosaura.-Echino-sauraorcesi differs markedly from other Echino-saurain having a more elongate snout, reducednumbers of infralabials and supralabials, in-

353

creased number of subdigital lamellae on thefourth toes, enlarged scales of dorsal and lateralsurfaces of the tail arranged in transversewhorls at the posterior margin of each autotom-ic segment, and in lacking a distinct vertebralcrest or paravertebral crests. Echinosaura orcesi isunique in its tail scutellation (consisting of con-tinuous transverse rows of enlarged scales onthe dorsal surface of the tail), lack of a contin-uous vertebral or paravertebral crest, and largenumber of femoral pares for females. Overall ithas a less spinose appearance than other speciesof the genus Echinosaura.

The elongate snout of E. orcesi relative to thatof other species is evidenced by ratios of thefrontonasal length/ internas al length. This ratiowas 0.63:0.80 for E. orcesi versus 1.2:3.0 for in-dividuals of other species. The ratio of frontallength/ internasallength is also diagnostic withratios of 0.85:1.0for orcesiand 1.4:4.0for otherspecies. Potentially, the tail of orcesi is longerwith SVL/TL ratios of 0.45:0.51 versus 0.56:0.95for other species.

Echinosaura horrida from Ecuador differs inhaving a conspicuous vertebral crest formed bytwo adjacent and continuous rows of enlargedand somewhat elevated scales, a short single,unpaired, internasal (as opposed to an elongateone) and a short frontal, fewer subdigital la-mellae on the fourth toe, a shorter tail relativeto the body length, a shorter snout, moderate-sized scales in the frontoparietal regían that donot extend past the posterior margin of the su-praoculars, tail ornamented with longitudinalrows of enlarged scales that alternate in size oneach autotomic segment with scales on the pos-terior margin of each segment larger than thegradient of scales anteriorly; enlarged scales notin contact transversely at the posterior marginof the autotomic segments. Echinosaura horridaappears to lack the conspicuous white band onthe base of the tail.

Variation in Echinosaura horrida.- The num-ber of supraoculars was noted by Uzzell to varyfrom two to three. In specimens with three su-praoculars, the first and second are large andthe third is considerably smaller. In specimenswith two supraoculars, the third is absent orsufficiently small to be confused with super-numerary scales at the posterior margin of theomito We found two supraoculars in all speci-mens from Ecuadorian localities in the Provinc-es of Imbabura and Esmeraldas and three su-praoculars in most specimens from the Provinceof Pichincha. Ratios of SVL/Tail Lengths were0.56:0.95. Supralabials 4:5; infralabials 3:4.

The new species resembles E. panamensis fromPanama in lacking enlarged scales on the ante-rior portions of the autotomic segments of thetail but differs in having enlarged scales ar-

354 T. H. FRITTS ET AL.

ranged in nearly continuous rings at the poste-rior margin of the autotomic segments and inhaving an undivided internas al. Panamanianspecimens are also similar in lacking a vertebralrow and in having a sinuous pattern of enlargedscales in the paravertebral region; they differ inhaving the enlarged scales on the central dor-sum in relatively continuous rows instead of be-ing broken into irregular diagonally oriented se-ries. Echinosaura panamensis also differs in hav-ing numerous enlarged scales on the lateralbody arranged in obliquely angled rows insteadof the sparse enlarged scales of E. orcesi,whichdo not contact each other and are arranged inextremely diffuse horizontal rows. Echinosaurapanamensis has SVL/TL ratios of 0.66:0.91; fivesupralabials; and 5 infralabials.

Echinosaura palmeri differs from E. orcesi inhaving paired internasals, in tail scutellation, inhaving continuous paravertebral rows of en-larged scales, and in tending to have the frontaldivided transversely. Echinosaurapalmeri hasSVL/TL ratios of 0.56:0.60; 5:6 supralabials; tourinfralabials.

The new species differs from Teuchocercus inhaving a much more elongate snout, in lackingcontinuous paravertebral rows, and in lackingconical scales on the tail that are separated bygranular scales. Most Teuchocercus have pairedinternasals.

Variation in Teuchocercus keyi.-Althoughsome variation exists, the cephalic scutellationtypically consists of paired internasals, fronton-asals, and prefrontals with a relatively short sin-gle frontal ending anterior to the level of thesecond supraocular. A pair of scales betweenthe second supraoculars bounds the frontal pos-teriorly and is bounded posteriorly by small ir-regular scales. Three supraoculars are presentwith the third much smaller than the others. Su-

perciliary scales separate the second and thirdsupraoculars from the ciliary row along the bor-der of the eJe.

InfrequentIy the internas al is single instead ofpaired; it is wider than long. The presence ofprefrontals in the presupraocular region is var-iable. The frontal is variable, depending on pre-frontal condition, short, usually ending anteriorto the second supraocular. One specimen(EPN2715) differs significantIy in having a sin-gle internas al, a medial scale posterior to thefrontonasals, and a fragmented frontal (appar-entIy irregularly divided longitudinally andtransversely). Teuchocercus have granular scalesthat cover some or all of the tympanum periph-erally, but juveniles and subadults have a por-tion of the tympanum exposed medially. Allshow a reduced amount of the tympanum ex-posed relative to Echinosaura. Four juvenileshave SVL of 24-27 mm and tails 28-43 mm. The

internas al is paired in all specimens examinedexcept EPN2715. Prefrontal scales arrear onone or both sides in nearly all specimens. Thescales of the subcaudal region are relativelylarge and arranged in series of three scales perautotomic unit, even near the base of the tail.The scales on the ventrolateral tail are small butnot granular. The enlarged scales on the dorsaland lateral tail are arranged in six longitudinalrows primarily at posterior margin of each au-totomic segment. The prominence of the para-vertebral rows increases with age and size.Paravertebral rows are separated by 2-4 scalesand weakly undulating. Subadult and juvenilespecimens exhibit a prominent white baTId atthe base of the tail and diffuse white bands onall sutures between supraoculars, but thesecharacteristics were not evident in the adultscomprising the type material (Fritts and Smith,1969). The ventral coloration is variable withbold dark brown and white checkered patternor only diffusely marked.

Acknawledgments.-We are grateful to the cu-rators and staff of the various institutions allow-ing access to the specimens and data in theircare and especially to C. W Myers for infor-mation and photographs for the type specimen.J. Carr and R. Reynolds provided helpful com-ments on drafts of the manuscript. T. Uzzell andN. Scott also provided useful reviews.

LITERATURE CITED

FRITTS,1. H, ANO H. M. SMlTH. 1969. A new teiidlizard genus from western Ecuador. Transactionsof the Kansas Academy of Science 72:54-59.

HARRIS,D. M. 1994. Review of the teiid lizard genusPtychoglossus. Herpetological Monograph 8:226-275.

KIZIRIAN,D. A. 1996. A review of Ecuadorian Proc-toporus (Squamata: Gymnophthalmidae) with de-scriptions of nine new species. HerpetologicalMonograph 10:85-155.

OFTEOAL,o. 1. 1974. A revision of the genus Anadia(Sauria, Teiidae) Arquivos de Zoología. (Silo Paulo)25:203-265.

PETERS,J. A. 1964. Dictionary of Herpetology. HafnerPublishing Co., New York.

UZZELLJR., 1. M. 1965. Teiid lizards of the GenusEchinosaura. Copeia 1965:82-89.

Accepted: 20 October 2001.

ApPENDIX1

Additional Specimens ExaminedEchinosaura horrida.-Ecuador: Pichincha Prov.:

USNM 196100, near Pacto; MCZ 149670-71, Santo Do-mingo de los Colorados; MCZ 149670-149671, SantoDomingo de los Colorados; EPN 2711-13, Santo Do-mingo de los Colorados; USNM 196095-97, 196099,257160-175, Dyott's Farm, km 121, 6 km east of SantoDomingo de los Colorados; KU 154666-67, 6 km eastof Santo Domingo de los Colorados; USNM 283530,

NEWSPECIES OF ECHINOSAURA

Rancho Santa Teresita, km 25 on road to Chane, SantoDomingo de los Colorados; USNM 285471-72, CentroCientifico Rio Palenque, 47 km south of Santo Domin-go de los Colorados; MCZ 145600, Centro CientíficoRio Palenque; MCZ 156145, 156147, 156149, LogoCreek, Centro Cientifico Rio Palenque; KU 152598-152599, Estacion Biologica Rio Palenque, 56 km northof Quevedo, 220 m; MCZ 145600, Centro CientificoRio Palenque, 47 km south of Los Rios; MCZ 156844,Centinella, 14.1 km southeast of Patricia Pilar; USNM285680-81, El Centinela, on the crest of Montañas deHa, 14.1 km east-southeast of Patricia Pilar, 550-600m; MCZ 171866, 4 km east-southeast of El Esfuerzo;MCZ 171869, 6 km east-southeast of El Esfuerzo; KU218389-395, 5 km west of La Florida, 860 m; EPN2710, Rio Salara; Esmeraldas Prov.: USNM 196093, 1.5km west of El Placer, 405 m; EPN2605, Caserio El Pan,Estero El Pan; NMW 31999:1, Rio Mataje; EPN 2609,2649,2614, Centro Comunal Mataje, Reserva Awa, 250m; EPN 2716, no further locality; BMNH 1901.3.29.25,Sto Javier; Imbabura Prov.: KU 133503-133504, Lita;BMNH 1901.3.29.85, Rio Lita; NMW 16950:1-16950:3,Lita; Manabí: KU 218396, 38 km northwest of El Car-men, ca El Carmen-Pedernales Rd., 330 m; "Ecuador,Prov. unknown": NMW 19982:2-19982:8, no furtherlocality; Colombia: Depto. Cauca: MCZ 160153, IslaGorgona.

Echinosauraorcesi.-Colombia: Depto. Valle: AMNH109822, approximately 13 km west of Dagua, Río An-chicaya drainage, 820 m; Ecuador: Esmeraldas Prov.:EPN 2574, Centro Comunal Mataje, near Rio Mataje,OloI2'04"N, 78°33'42"vy' 250 m; Carchi Prov.: NMW32000:1, San Marco, 670 m, NMW 32000:2, San Marco,700 m.

Echinosaurapalmeri.-Panama: Darien Prov.: USNM

355

141789, Rio Tapalisa, 546 m; KU 113612-16, Rio Jaque,1.5 km above Rio Imamado, 50 m; KU 112617, ridgebetween Rio Jaque and Río Imamado, 730 m; KU113618, northeast slope Cerro Sapo, 820 m; KU113620, north end Serranía de Pirre, 320 m; KU113621, Rio Tuira at Rio Mono, 150 m; KU 113622,southeast slope Cerro Pirre, 1520 m; KU 76165, La-guna, 820 m; KU 76167-68, Tacarcuna, 550 m; SanBIas Prov: KU 113619, Camp Sasardi, 12 m; Colombia:Depto. Boyaca: USNM 127097-98, Muzo region; Dep-tú. Valle: USNM 151621, Virology Field Station, RioRaposo near Buenaventura; KU 144149, Quebrada LaGuinea, 2 km east of Cisneros, 400 m. Colombia: Dep-to. unknown: BMNH 1923.10.12.14, no specific local-ity.

Echinosaura panamensis.-Panama: Veraguas Prov.:USNM 266154, Los Girasoles Escuela Agricola AltoPiedra, 5 km northwest of Santa Fe; Canal Zone:USNM 92594, Canal, Rio Grande; Cocle Prov.: KU76169-170, El Valle, 560 m; Darien Prov.: USNM50144, Pirri Range, head Rio Limon; Panama Prov: KU80584, Finca La Sumbadora, 570 m; MCZ 127854-127855, MCZ 127763, Cerro Campana; ANSP 21593,Cerro Campana, 909 m; ANSP 21087, El Valle, 606 m.

Teuchocercus keyi.-Ecuador: Esmeraldas Prov.:USNM 196094, 1 km west of El Placer, 400 m; NMW32001:1, near Mataje, 350 m; NMW 31988, Rio Sabal-era, 620 m; NMW 32001:9-32001:11, Rio Mira, 620 m;EPN 2717, 2719-720, Alto Tambo, 570 m; EPN 2718,Rio Sabalera; Carchi Prov.: EPN 2715, El Pailon,78°14'11"W; 01°00'OTN; NMW 32001:2-32001:4, RioSabalera, near ajala, 630 m; NMW 32001:5, ajala, 400m; NMW 32001:6-32001:8, Rio Sabalera, 620 m; Im-babura Prov.: BMNH 98.4.28.48-98.4.28.49, Paramba;Pichincha Prov.: KU 218397-98, 5 km west of La Flor-ida, 860 m.

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