HERP QL 666 -L25 T67 ‘velentific Papers Natural History Museum The University of Kansas 16 June 2000 Number 15:1-38 Ecuadorian Lizards of the Genus Stenocercus (Squamata: Tropiduridae) By OMAR ToRRES-CARVAJAL Natural History Museum and Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-2454, USA CONTENTS JNIBISIPRUAN CP eaceecesacenooicecuuegtdoueeke003e EESELCh83 oc EBC HERES cose EEE ACO AEE HOSES EE EEE EEE HOSES Ee SaR Ese EG SEE RenapERBSENC 1 IRI ERSIUIM BIN sccosdsocaccccsstodbecadacsdeecoodetacdeeGscecasee-05oe83 CelGeeoC ac HADOOLe Gc SEaSE SEEESS GaECEHOE aoesL UD oa neSeasncecceesbeneueceooronen D, HSPN RO) DUKA COUNT, ccsacccneccascedcontcbocktcbececec eB e eco SO eee Eecnreeo EES OUEEC Ee JBeee: GRE CEES SE san eec Sor eS CBee sone ceeeeREoHaceEcanec” y) | HUGO ICAI, SUIMIM INS” ccosccocceededoacdeeenctdecseestsde eae esuscse-3= BEE HOSE BCOTE SESE nOOeeE Ho ceeconb ee Jena seaeedsaneanSaccbeaascos 2 PN ETRNIORTILISIDI GIMTSINTTIS 3, canaccatabaoobona socks eee cB On GSEES Ee TOPE ABE OO DEBDE-E SOE DE GEE cr SBE REECE EOE CECE CEE ace ECE EEEEECOSE 3 NEGATE RU ATEVAIN TD VIE AE @ 1D) Siteccessvcescccsecessesoresccsesassncecencesecccecssouccrechossvassieuansavecceeessseeete aaeeeeee sree 3 SUMMAR YEO Eal-Ax ONO MIG CHARA GIE RS eeeeeesrrsccrscetercteseeceerrere ere 4 SIPEXCHIBS, ANCOTUINAINS, Sone csacacoodaiacaBendsescectoatece: Backer c ERCEREEE CROCE dec SBE BER SURO RaE Enc eee ROMCE ER ert cei nec teeire 5 BSI KOXGH EKO RUANI PAE Co 3 sakes ras SSE anc COSA EDR aar SERED ERASERS PRE SEES ee SCE eee Bee cL See ee Ree SSE 34 GEOGRAPHICAT DISTRIB UM ONiceccssssceccccocsorcccancasscocscter Cae cetaces sae Naeceee sncaeccaenss wateceacsdsseccecersadcnsean ste 34 | EXGGOIL(OXGNC/A | DISH RAT ERO (ON) cecaoceceacsdeooteecssocsecot eco see Coe ee SSE Ee Reese EB eRO SER ace. Boa Bea D Oconee sec oRcRaa ec 35 KEY TO THE SPECIES OF STENOCERCUS OF ECUADOR ..............:.cc:cccssccessecesseesesestseenseesees SD JETS TY BA RANG PLU] 28 Be CC a am SE cee ner cee ea eee or ener ere Ree ese eee 36 JRIPIPIBINI DID scacsaad eS aopeodtnac ce naene nod dcosus COB HEGEE ROBO BRC nc Baan heSoCC RBC SPST SCe ce Scene Cer rene Re sen eae eRc eee recientes 37 ABSTRACT A taxonomic review of lizards of the genus Stenocercus in Ecuador revealed that col- oration and certain external morphological characters, such as scales around midbody, the relation between tail length and total length, and number of subdigital lamellae on Finger IV are important taxonomic characters. Fourteen species, including two species new to science are recognized: Stenocercus aculeatus, S. angel sp. nov., S. carrioni, S. chota sp. nov., S. festae, S. guentheri, S. haenschi, S. humeralis, S. iridescens, S. limitaris, S. ornatus, S. rhodomelas, S. simonsii, and S. varius. Of the new species, Stenocercus angel occurs at elevations of 3015-3560 m in Provincia Carchi and Provincia Sucumbios, and Stenocercus chota occurs at elevations of 1575-1940 m in the Chota Valley, Provincia Imbabura. All species are redescribed, except for Stenocercus carrioni, S. limitaris, and S. simonsti, which have recent and appro- © Natural History Museum, The University of Kansas ISSN No. 1094-0782 ScreNtTIz=IC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS priate complete descriptions. A neotype for S. festae is proposed. Sexual variation in morphological characters is described for all species, along with geographical variation in S. guentheri, S. iridescens, and S. varius. The distribution of each species is mapped, and new color descriptions and ecological and reproductive data are provided. A dichotomous key is also included to assist in the identification of specimens. Key words: Tropiduridae, Stenocercus, Ecuador, Taxonomy, Ecology, Distribution. RESUMEN Una revision taxonomica de las lagartijas del género Stenocercus del Ecuador revel6 que la coloracion y ciertos caracteres de morfologia externa, tales como el numero de escamas alrededor del medio cuerpo, la relacion entre la longitud de la cola y la longitud total y el numero de lamelas del cuarto dedo de la mano son caracteres taxondmicos importantes. Se reconocen 14 especies, incluyendo dos nuevas para la ciencia: Stenocercus aculeatus, S. angel sp. nov., S. carrioni, S. chota sp. nov., S. festae, S. guenthert, S. haenschi, S. humeralis, S. tridescens, S. limitaris, S. ornatus, S. rhodomelas, S. simonsti y S. varius. De las especies nuevas, Stenocercus angel habita entre los 3015 y 3560 m en las provincias de Carchi y Sucumbios, y Stenocercus chota habita en el valle del Chota, provincia de Imbabura, entre los 1575 y 1940 m. Todas las especies fueron redescritas, excepto S. carrioni, S. limitaris y S. simonsii, las cuales cuentan con descripciones recientes y apropiadas. Se propone un neotipo para S. festae. También se indica la variacion sexual de los caracteres morfol6gicos para todas las especies y la variacion geografica para S. guentheri, S. iridescens y S. varius. Se presentan mapas de distribucién para todas las especies y se proveen nuevos datos ecoldgicos y reproductivos, asi como nuevas descripciones de color. También se incluye una clave dicotomica para ayudar en la identificacién de los especimenes. Palabras claves: Tropiduridae, Stenocercus, Ecuador, Taxonomia, Ecologia, Distribucion. INTRODUCTION Tropiduridae is a family of medium-sized Neotropi- cal iguanian lizards composed of nine genera (Frost, 1992). Stenocercus Duméril and Bibron (1837) occurs in western South America from northern Colombia to northern coastal Argentina. These lizards occur in a variety of habitats: dry and humid tropical forests, montane forests, and paramo, at elevations of 0-4000 m. The phylogeny of Stenocercus has not been studied because several alpha-taxonomic problems have been pending. Fritts (1974) briefly charac- terized 29 species and reported nine species for Ecuador. Cadle (1991) provided amended descriptions of two Ec- uadorian species (S. simonsti and S. carrioni). He also de- scribed S. limitaris (Cadle, 1998). Because little has been done to resolve the taxonomic problems of the Ecuador- ian members of this group, specimens in several collec- tions have remained unidentified or have been misidentified. The objective of this study is to provide a better understanding of the diversity of this group of liz- ards in Ecuador. HIsTORICAL SUMMARY Thirty-one percent of the species of Stenocercus were described in the last three decades (Ayala and Castro, 1982; Cadle, 1991, 1998; Corredor, 1983; Fritts, 1972). Gray (1845) provided the first description of Stenocercus from Ecuador (Leiocephalus ornatus). Most Ecuadorian species of Stenocercus were described in the second half of the Nine- teenth Century. Liocephalus iridescens and Microphractus humeralis were described by Gtinther (1859a, b), and Leiocephalus aculeatus by O’Shaughnessy (1879); these de- scriptions were based on collections made by Roff depos- ited in the British Museum (Natural History). Specimens from subsequent collections deposited in the British Mu- seum by explorers and naturalists (Brenchley, Fraser, Buckley, and Simons) were described by Boulenger (1885, 1899) as L. guentheri, L. rhodomelas, S. varius, and S. simonsii. Liocephalus festae was described by Peracca (1897), and L. haenschi by Werner (1901); the latter description was based on material collected by Richard Haensch and deposited in the Zoologisches Museum of Berlin. The description of S. carrioni (Parker, 1934) is based on specimens deposited in the British Museum from Provincia Loja and Provincia Zamora by Carrion. After a hiatus of 40 years, during which there were no noteworthy contributions to our knowledge of Stenocercus in Ecuador, Fritts (1974) published a brief characterization of nine species of Ecuadorian Stenocercus. Of the 12 Ecuadorian species of Stenocercus, only four (S. carrioni, limitaris, simonsii, and varius) were described originally as members of this genus. Stenocercus humeralis was described as Microphractus humeralis, and the remain- ing species (S. aculeatus, festae, guentheri, haenschi, iridescens, ornatus, and rhodomelas) were placed originally in Leiocephalus. The latter genus was erected by Gray (1827) and often has been misspelled as Liocephalus. Etheridge (1966) distinguished South American from West Indian species of Leiocephalus and allocated the South American species to the genus Ophryoessoides. He acknowledged the ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS ) likely association of some species of Ophryoessoides with Stenocercus. Fritts (1974) redefined the genera Ophryoessoides and Stenocercus. Frost and Etheridge (1989) placed Stenocercus in the family Tropiduridae. In his phy- logenetic analysis of the Tropidurinae, Frost (1992) pro- posed that Ophryoessoides and Proctotretus should be placed in synonymy with Stenocercus, which he placed in a new tribe, Stenocercini. Four species, Stenocercus boettgeri, S. erythrogaster, S. formosus, and S. ornatissimus were erroneously considered to occur in Ecuador (Almendariz, 1992; Miyata, 1982; Pe- ters, 1967; Peters and Donoso-Barros, 1970). Stenocercus boettgeri, S. formosus, and S. ornatissimus occur only in Peru (Fritts, 1974). Stenocercus erythrogaster is known only from northern Colombia (Peters and Donoso-Barros, 1970). The objectives of my work with Ecuadorian Stenocercus are to (1) describe two new taxa, (2) redescribe poorly known species, and (3) add new data on the morphology, biology, ecology, and distribution of all species. ACKNOWLEDGMENTS I thank the following persons for the loan of specimens: Ana Almendariz (EPN), Franco Andreone (MZUT), Collin McCarthy (BM), Luis A. Coloma (QCAZ), William E. Duellman (KU), Rainer Gtinther (ZMB), Ronald Heyer (USNM), Gunther Kohler (GMF), Manuel Morales (UDAR), and Jean-Marc Touzet (FHGO). I extend my deepest thanks to many field companions—Mayra Araque, Veronica Barragan, Carlos Boada, Santiago Burneo, Carlos Carpio, Amaranta Carvajal, Luis Coloma, Lucia de la Torre, Marcelo Diaz, Juan Guayasamin, Aura Paucar, Alexandra Quiguango, and Santiago Ron. I am grateful to Analia Pugener for providing one of the illustrations and Chris- topher J. Raxworthy for his advice. Color photographs re- produced in figures 4 and 5 were provided by Luis A. Coloma (LAC) and William E. Duellman (WED); Coloma took the photographs of preserved specimens.. This study is based ona research carried out in the Museo de Zoologia (QCAZ) that was completed as a requirement for the de- gree of “Licenciatura” at the Department of Biological Sci- ences of the Pontificia Universidad Catdlica del Ecuador. It was approved by a committee composed of Luis Coloma (major advisor), Ana B. Mafla, and Tjitte de Vries. I thank all of them, especially Luis Coloma, for their encourage- ment and criticism of the present work. The manuscript also benefited from critical review by Richard E. Etheridge, Darrel R. Frost, David A. Kizirian and careful editing by William E. Duellman and Linda Trueb. Finally, I am in- debted to Luis Coloma, Giovanni Onore, and my family for moral support. MATERIALS AND METHODS All of the specimens examined in the course of this study are listed in the Appendix. Throughout the text and appendix, the specimens are listed with their catalog num- bers, preceded by the following codes: BM = British Mu- seum (Natural History); EPN = Escuela Politécnica Nacional, Quito, Ecuador; FHGO = Fundacion Herpetologica Gustavo Orcés, Quito, Ecuador; KU = Natu- ral History Museum, The University of Kansas; MRHN = Institut Royal des Sciences Naturelles de Belgique, Brus- sels; MZUT = Museo Regionale di Scienze Naturali, Torino; QCAZ = Museo de Zoologia, Pontificia Universidad Catolica del Ecuador, Quito; SMF = Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Ger- many; UDAR = Museo de la Universidad del Azuay, Cuenca, Ecuador; USNM = Smithsonian Institution, Na- tional Museum of Natural History; ZMB = Universitat Humboldt, Zoologisches Museum, Berlin. Measurements were made with calipers and recorded to the nearest 0.1 mm with the exception of the snout-vent length (SVL) and tail length (TL); the latter two measure- ments were taken with a ruler and recorded to the nearest 1 mm. The sex of each specimen examined was determined either by dissection, or by noting the presence of hemipenes or another sexually dimorphic character (e.g., black spots in the gular region of females of Stenocercus chota). Clutch size was determined by counting oviductal eggs. Egg vol- ume was calculated by the formula for a prolate spheroid: V =4/3 nm (length/2) - (width/2)?. Drawings of specimens immersed in alcohol were made with a Wild-M8 stereo microscope equipped with a camera lucida. Ecological distributions of each species are given in the Ecuadorian life zones defined by Caftadas-Cruz (1983); climatological data for the life zones also are from Canadas- Cruz. Names for the physiographic regions (e.g., Cordil- lera Occidental, Saraguro Basin) were taken from Duellman (1979). Locality records shown on the distribution maps are based on specimens examined and literature records. Diagnoses include several characters not used by other authors (e.g., Cadle, 1991, 1998; Fritts, 1974). Data included in the diagnoses were taken from the specimens examined, except for data taken from Cadle (1991, 1998) for S. carriont, limitaris, and simonsii. Descriptions of syntypes, holotypes, and a neotype are provided. I did not examine the whole type series of any of the species for which I describe a syntype. The purpose of describing a syntype is to add data to the original description of each taxon based on one of the type specimens. Data on sexual variation are pre- sented for most species and data on intraspecific geo- graphic variation are presented for Stenocercus iridescens, S. guentheri, and S. varius. A dichotomous key is included to aid the identification of specimens. 4 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS SUMMARY OF TAXONOMIC CHARACTERS The taxonomic conclusions of this study are based on the observation of external morphological features and color patterns. Character definitions follow those of Cadle (1991), Fritts (1974), Frost (1992), and Vitt and de la Torre (1996). Measurements and characters of scutellation are summarized in Table 1. MEASUREMENTS Head length-head width ratio (HL/HW).—Head length is the linear distance from the tip of the snout to the anterior edge of the tympanum. Head width was measured at the widest point of the head. Head height-head width ratio (HH/HW).—Head height was measured at the level of the highest point of the head. Body height-body width ratio (BH/BW).—Both mea- surements were recorded at midbody (i.e., halfway be- tween the insertion of the fore and hind limbs). Tail length—total length ratio (TL/ToL).—Total length is the sum of SVL and tail length. Specimens with broken or regenerated tails were not included. Regenerated tail length—tail length ratio (RT/TL).— In this case, the tail length is the sum of the non-lost-tail length and the regenerated tail length. SCUTELLATION Scales around midbody (SM).—Number of longitu- dinal rows of scales counted in a transverse line around midbody (i.e., halfway between the insertion of the fore and hind limbs). Vertebral scales (VS).—Number of middorsal scales from the occipital scales to the level of the posterior edge of the thigh when the latter was extended perpendicular to the body. Paravertebral scales (PS).—Number of scales in the scale row laterally adjacent to the vertebral row. These were counted from the occipital scales to the level of the poste- rior edge of the thigh when the latter was extended per- pendicular to the body. Gular scales (GS).—Number of scales of the gular re- gion counted between the ventral edges of the tympani. Tiny scales within the ear margin were not included. I fol- lowed Cadle’s (1991) procedure—i.e., started and ended the counts with the first enlarged scale ventral to the tympani. Supraocular scales (SS).—Number of scales on the dorsal surface of the orbit counted in a transverse line across its greatest width. Internasal scales (IS)—_Number of scales between the nasal scales. Internasals are located immediately posterior to postrostrals. They can be distinguished from the latter by their anteroposterior elongate shape, in contrast to the transverse elongate shape of postrostrals. Posterior thigh scales.—The scales of the posterior sur- face of the thighs are smooth, keeled, keeled and mucr- onate, or granular. Occipital scales.—The scales of the occipital region on the dorsal surface of the head are juxtaposed or imbricate. They are smooth, wrinkled, or keeled. Ventral scales.—The scales of the ventral body sur- face, from the neck to the vent, are smooth, keeled, or keeled and mucronate. Angulate temporal scales.— There are one or more en- larged, keeled scales that are approximately aligned with the superciliaries and that are located along the border between the lateral temporal scales and the posterior dor- sal head scales. In many species of Stenocercus, these scales are not differentiated. One Ecuadorian species (S. aculeatus) has angulate temporals that bear a bladelike vane; I refer to this condition as projecting angulate temporals. Apical pits.—Gular or ventral scales may bear an api- cal pit. Subdigital lamellae on Finger IV (SF).—Number of subdigitals counted from the point of attachment of Fin- gers III and IV to the terminus of Finger IV. Subdigital lamellae on Toe IV (ST).—Number of subdigitals counted from the point of attachment of Toes III and IV to the terminus of Toe IV. Neck FoLps Stenocercus exhibits various kinds of neck folds (Fig. 1), which are described below. Antehumeral fold.—A vertical or oblique fold that ex- tends across the scapular region. Posthumeral fold.—A fold extending obliquely down- ward the dorsolateral fold behind the forelimb. Gular fold.—A fold immediately in front of the fore- limb; it extends ventrally onto the neck. Dorsolateral fold.—This fold is variable in length. It extends along the body and over the forelimb insertion of the antehumeral fold, with which it usually is confluent. Ventrolateral fold.—A fold that extends along the ven- trolateral region of the body posterior to the forelimb. Supra-auricular fold.—This fold lies between the antehumeral fold and the ear and represents a continua- tion of the dorsolateral fold. Oblique neck fold.—The oblique neck fold usually is located about midway between the posterior border of the ear and the forelimb. It is approximately parallel to the antehumeral fold. ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 5 supra-auricular postauricular antegular longitudinal neck Fig. 1. Antegular fold.—This fold traverses the ventral sur- face of the neck anterior to the gular fold. Longitudinal neck fold.—This longitudinal fold ex- tends along the ventrolateral part of the body from the posteroventral edge of the ear to the forelimb. Postauricular fold.—This fold lies immediately behind the ear usually is confluent with the longitudinal neck fold and the supra-auricular fold. Rictal fold.—A fold ventral and anteroventral to the ear. POSTHUMERAL AND POSTFEMORAL POCKETS Type 1.—Pocket absent, no apparent skin modifica- tion. Type 2.—A noticeable skin modification, such as bare oblique neck antehumeral dorsolateral Ki: pepe eg ventrolateral patch of skin, a series of wrinkles in the skin, or a shallow depression lined with scales different from surrounding body scales. gular Neck folds in Stenocercus. Illustration by L. Analia Pugener. Type 3.—A shallow pocket, similar to Type 2, but with the addition of an overhanging fold of skin or thickened border. Its depth is generally less than half the diameter of the opening. Type 4._A moderate to deep pocket usually having a broad circular opening (narrow in Stenocercus aculeatus); its depth is greater than half the diameter of the opening. Type 5.—A moderate to deep pocket with a narrow, slitlike opening; its depth is generally greater than half the greatest diameter of the opening. This type was observed only in postfemoral pockets in the species studied. SPECIES ACCOUNTS Stenocercus aculeatus (O’Shaughnessy) Leiocephalus aculeatus O’Shaughnessy, 1879:303. Syntypes: BM 1946.8.12.33-36, from “Moyobamba, [Departamento San Martin], Peru.” Leiocephalus angulifer Werner, 1901:595. Holotype: ZMB 16594, from “Ec- uador.” Synonymy fide Burt and Burt, 1931. Leiocephalus iridescens aculeatus (part)—Burt and Burt, 1931:269. Ophryoessoides aculeatus—Etheridge, 1966:88. Stenocercus aculeatus—Frost, 1988:78. Diagnosis.—(1) Maximum total length in males 393 mm (n = 4); (2) maximum total length in females 322 mm (n = 8); (3) vertebral scales 38-43; (4) scales around mid- body 36-45; (5) internasal scales 4-8; (6) gular scales 17- 19; (7) lamellae on Finger IV 15-19; (8) lamellae on Toe IV 19-26; (9) posthumeral pocket Type 4; (10) postfemoral pocket Type 3; (11) projecting angulate temporals present; (12) one row of enlarged supraoculars that are more than twice the size of scales in adjacent rows; (13) occipital scales large, wrinkled, juxtaposed; (14) ventral scales keeled, imbricate, mucronate; (15) scales on posterior surfaces of thighs keeled, imbricate, mucronate; (16) antehumeral fold 6 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 1. Summary of scutellation and measurements of the Ecuadorian species of Stenocercus. Range (first line), + SD (second line) and n (third line) are given. Character S. aculeatus S. angel S. carrioni S. chota S. festae S. guenthert S. haenschi Scales around midbody 36-45 52-60 66-96 45-59 47-64 59-83 57-64 40.50 + 2.58 55.00 + 2.71 82.43 + 8.13 50.93 + 2.96 54.91 +4.43 68.00 + 4.97 60.50 + 4.95 12 10 21 43 43 75 2 Vertebral scales 38-43 39-52 55-72 39-50 39-55 44-64 50 40.25 + 2.05 45.20 + 3.43 64.43 + 5.18 45.19 + 2.79 46.70 + 3.50 55.36 + 3.74 — 12 10 21 42 43 75 1 Paravertebral scales 45-54 50-67 76-96 55-74 46-72 59-89 64 48.83 + 2.79 58.40 + 5.08 86.67 + 5.36 61.60 + 4.65 62.02 + 5.37 74.07 + 6.07 — 12 10 21 42 43 75 1 Gular scales 17-19 20-25 37-52 18-25 16-29 21-31 56 18.25 + 0.62 21.70 + 1.64 46.48 +4.14 20.52 + 1.82 22.74 + 2.30 25.53 + 2.12 — 12 10 21 42 43 75 1 Supraocular scales 45 4-5 4 4-6 4-6 5-7 5 4.08 + 0.29 4.80 + 0.42 — 5.28 + 0.50 5.02 + 0.51 5.71 + 0.56 — 12 10 21 43 43 75 1 Internasal scales 4-8 14 5-7 2-4 2-4 2-4 3 5.67 + 1.37 2.40 + 0.84 5.81 + 0.56 3.26 + 0.73 3.44 + 0.70 3.28 + 0.83 — 10 10 21 43 43 75 1 Subdigital lamellae: Finger IV 15-19 13-22 23-28 14-20 14-22 15-23 26-28 17.50 + 1.38 17.20 + 2.25 25.19 + 1.25 17.64 + 1.48 17.40 + 1.75 18.17 + 1.62 27.00 + 1.41 12 10 21 42 42 75 2 Subdigital lamellae: Toe IV 19-26 20-28 27-32 23-31 21-33 22-36 30 23.25 + 1.82 25.20 + 2.15 29.80 + 1.44 26.29 + 1.58 26.57 + 2.17 27.39 +2.41 — 12 10 20 42 42 74 1 Head length/head width 1.12-1.37 1.16-1.30 — 1.17-1.41 1.14-1.43 1.10-1.35 1.34 1.23 + 0.09 1.22 + 0.05 — 1.28 + 0.06 1.29 + 0.06 1.26 + 0.05 — 12 10 — 4] 42 73 1 Head height/head width 0.79-1.07 0.78-0.93 — 0.67—0.97 0.75-1.16 0.75-1.05 0.86 0.87 + 0.07 0.88 + 0.05 a 0.84 + 0.06 0.86 + 0.07 0.87 + 0.07 —_— 12 10 “= 41 42 72 1 Body height/body width 0.67-1.15 0.71-0.97 — 0.42-1.04 0.63-1.18 0.50-1.15 0.8 0.95 + 0.17 0.83 + 0.09 — 0.69 + 0.16 0.83 + 0.12 0.81 + 0.14 —_— 10 9 — 39 31 68 il Regen. tail length/tail length — 0.46-1.63 — 0.03-3.07 0.27-4.14 0.11-3.40 — _ 0.85 + 0.46 — ill se 11{0)5) 1.45 + 1.31 1.64 + 0.99 — _ 5 — 9 10 13 — Tail length/total length 0.65-0.74 0.62-0.65 0.56-0.60 0.55-0.66 0.52-0.71 0.57-0.72 — 0.69 + 0.02 0.64 + 0.01 0.57 + 0.01 0.63 + 0.02 0.64 + 0.04 0.64 + 0.03 — 12 4 10 29 22 53 _— Maximum size males (mm) 393 214 — 207 241 255 — Maximum size females (mm) 322 155 — 172 206 188 = nearly inconspicuous; (17) tail strongly compressed later- ally; (18) gular region of males black; (19) dorsum bronzed green in males and pale brown in females. Stenocercus aculeatus differs from other species of Stenocercus by the combination of the following charac- ters: keeled ventrals, large occipital scales, one row of en- larged supraoculars that are more than twice the size of the scales in adjacent rows, one canthal scale, two project- ing angulate temporals, distinct dorsolateral crest from neck to base of tail, posthumeral pocket Type 4, and postfemoral pocket Type 3. Description of syntype BM 1946.8.12.34.—Head wider than high (HH/HW = 0.91); occipitals , parietals, and interparietal large, slightly wrinkled, and juxtaposed; 5 internasals; 1 canthal on each side, separated from the nasal by tiny scale; supraoculars slightly wrinkled, 1 row of scales being more than twice size of scales in adjacent rows; lateral temporals imbricate, slightly keeled; 2 pro- jecting angulate temporals on each side of head; parietal eye not visible; gulars keeled, imbricate; mental in contact with first pair of infralabials and first pair of postmentals. Dorsal and lateral scales of body and neck keeled, imbricate; vertebrals large, forming prominent, serrate vertebral crest (Fig. 2); distinct dorsolateral crest from neck to base of tail; ventrals keeled, imbricate, mucronate; preau- ricular fringe absent; antehumeral fold nearly inconspicu- ous; limb scales keeled, imbricate, mucronate; 19 lamellae on Finger IV; 26 lamellae on Toe IV; tail strongly com- ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 7 Table 1 Continued Character S. humeralis S. iridescens S. limitaris S. ornatus S. rhodomelas S. simonsit S. varius Scales around midbody 98-119 35-52 39-54 46-58 43-55 79-102 74-88 108.38 + 6.06 41.57 +3.31 47.30 + 3.16 51.95 + 2.73 48.81 + 2.72 94.20 + 6.63 82.28 + 4.38 34 42 43 40 32 20 25 Vertebral scales 81-102 40-52 40-52 36-50 43-54 59-98 60-74 90.73 + 6.01 45.31 + 3.22 45.70 + 2.54 44.03 + 3.48 47.69 + 3.24 73.75 + 9.76 67.64 + 3.57 33 42 43 39 32 20 25 Paravertebral scales 106-145 43-58 — 53-66 49-61 94-118 76-97 127.33 + 7.70 49.50 + 3.76 — 60.00 + 3.55 55.09 + 2.97 107.20 + 9.02 87.24+5.31 33 42 = 39 eZ 20 25 Gular scales 43-68 16-20 17-23 15-23 17-20 36-57 42-60 49.44 + 4.43 18.19 + 1.09 20.10 + 1.28 18.45 + 1.47 18.31 + 0.93 49.26 + 5.33 50.64 + 4.46 34 42 43 40 32 19 25 Supraocular scales 6-8 2-5 3-5 S/ 3-6 4 5-7 6.97 + 0.46 3.33 + 0.69 — 5.43 + 0.59 5.22 + 0.66 — 5.52 + 0.59 34 42 44 40 32 19 25 Internasal scales 2-4 2-4 45 2-4 2-4 6-9 4-5 3.88 + 0.41 2.45 + 0.59 — 3.63 + 0.67 2.47 + 0.62 7.11 £0.76 4.04 + 0.20 34 42 44 40 32 19 25 Subdigital lamellae: Finger IV 24-33 15-18 W238 17-25 14-21 24-28 24-28 29.09 + 2.01 15.90 + 0.89 19.70 + 1.26 21.23 + 1.98 16.81 + 1.53 26.15 + 1.23 25.61 +1.16 34 4] 44 40 32 20 23 Subdigital lamellae: Toe IV 28-40 22-28 24-32 26-37 22-29 28-37 27-34 36.79 + 2.52 24.73 + 1.57 27.50 + 1.93 30.03 + 2.28 25.69 + 1.62 31.15 + 2.46 29.78 1.51 33 41 44 40 32 20 23 Head length/head width 1.15-1.43 1.09-1.31 — 1.06-1.37 0.98-1.50 — 1.21-1.50 1.28 + 0.06 1.23 + 0.05 _ 1.28 + 0.06 1.25 + 0.09 — 1.32 + 0.07 34 42 -- 40 30 — 25 Head height/head width 0.67-0.93 0.72-1.04 — 0.71-0.98 0.69-0.99 — 0.73-1.04 0.78 + 0.06 0.88 + 0.07 — 0.85 + 0.05 0.83 + 0.07 — 0.83 + 0.07 34 42 — 40 29 — 25 Body height/body width 0.56-0.9 0.66-1.19 -— 0.62-1.36 0.64-1.03 _ 0.54-0.96 0.70 + 0.08 0.86 + 0.14 — 0.82 0.16 0.81 + 0.10 = 0.73 + 0.09 28 36 — 34 16 a 23 Regen. tail length/taillength —1.58-3.84 0.55-3.38 = 0.01-2.76 0.32-3.82 = 0.01-2.87 2.51 +0.81 1.97 + 2.00 — 1.41 + 0.95 1.16 + 1.33 — aS E 129 6 2 — 10 6 — 6 Tail length/total length 0.62-0.68 0.63-0.70 0.66-0.70 0.60-0.72 0.57-0.66 0.59-0.65 0.60-0.66 0.66 + 0.01 0.67 + 0.02 — 0.68 + 0.02 0.63 + 0.02 0.63 + 0.01 0.63 + 0.02 25 30 — 26 23 5 15 Maximum size males (mm) 301 292 — 294 235 — 234 Maximum size females (mm) 272 294 — 207 236 — 208 pressed; all caudals keeled, imbricate, and mucronate; ver- tebral crest extending more than half length of tail; tail length 74% of total length; posthumeral pocket Type 4 with narrow opening; postfemoral pocket Type 3. Data on syntype BM 1946.8.12.34: Male (Fig. 2); SVL = 103 mm; TL = 290 mm; HW = 17.5 mm; HL = 23.7 mm; HH = 15.4 mm; SM = 36; VS = 38; PS = 48; SS = 4; IS = 4; GS = 17; SF = 19; ST = 26. Variation.—Measurements and scutellation of Stenocercus aculeatus in Table 1. Head (Fig. 3A and C) nor- mally wider than high (HH/HW = 0.79-1.07, x = 0.87 + 0.07, n = 12); 4-8 internasals, usually 6 (50% of the speci- mens); body usually wider than high (BH/BW =0.67-1.15, X= 0.95 + 0.17, n = 10); 15-19 lamellae on Finger IV; 19-26 lamellae on Toe IV; tail length 65-74% of total length. Sexual variation for mensural characters in Stenocercus aculeatus is presented in Table 2. Color in life: Dorsum bronzed-green in males; flanks brown; white stripe extending from ear to forelimb; white stripe extending from above ear to shoulder; another white stripe extending from dorsolateral crest to forelimb; black gular region (O'Shaughnessy, 1879, 1881). An adult male (KU 121093) differs from the above de- scription by the following features: dorsum brown; flanks lighter, almost yellow-brown; black supraxillary patch; face black; throat black; venter pink with bluish-gray sides; sides of tail with blue spots; ventral surface of tail blue (John D. Lynch, field notes, 24 July 1968). Female KU 8 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 2. Stenocercus aculeatus, syntype, BM 1946.8.12.34, male. 121092: dorsum of body pale brown; flanks, dorsum of head, and base of tail dark brown; cream-brown line in front of shoulder; reddish-brown line below eye with black edges; greenish-brown venter with some pink coloration; tongue reddish brown anteriorly and cream posteriorly; iris pale copper-brown (Fig. 4G; John D. Lynch, field notes, 4 July 1968). Color in preservative: Males with dark brown or dark olive dorsum; dark triangular marks pointing backwards on vertebral line; sides of neck with scattered pale blue and green scales; white stripe descending from dorsolat- eral crest to insertion of forelimb; head dark brown or ol- ive-green; large white mark behind each eye in 10% of specimens; gular region black; venter iridescent pink or grayish green with scattered blue and pink scales; tail ol- ive-green, with large dark transverse bands on 90% of specimens. Females dark brown; sides and dorsum of head pale brown in contrast to body color in some specimens. Dark brown stripe extending posteroventrally from eye to supralabials in both sexes. Natural history.—One female (EPN 5902) collected in May 1954 contained two oviductal eggs; the sizes of these eggs are 22.8 mm x 8.9 mm and 23.2 mm x 9.8 mm, and their respective volumes are 945.6 mm? and 1166.6 mm‘*. The smallest individual (QCAZ 1635) was collected on 01 May 1993 and has a total length of 112 mm (SVL=39 mm, TL =73 mm). Distribution and ecology.—In Ecuador, Stenocercus aculeatus is known from elevations of 537-1200 m on the eastern slopes of the Cordillera Oriental. The species in- habits the upper valleys of the Rio Napo and Rio Pastaza (Atlantic Drainage) in Provincia Pastaza (Fig. 6). The dis- tribution lies mainly within Very Humid Premontane For- est and Pluvial Premontane Forest life zones. The mean Table 2. Sexual variation in scutellation and measurements of Stenocercus aculeatus. Range (first line), X + SD (second line) and n (third line) are given. Character Females Males Scales around midbody 37-45 36-43 40.38 + 2.39 40.75 + 3.30 8 4 Vertebral scales 38-43 38-43 40.25 + 1.91 40.25 + 2.63 8 4 Paravertebral scales 45-54 48-53 48.00 + 2.83 50.50 + 2.08 8 4 Gular scales 18-19 17-19 18.38 + 0.52 18.00 + 0.82 8 4 Supraocular scales 4-5 4 4.13+0.35 — 8 4 Internasal scales 4-8 47 5.63 + 1.51 5.75 + 1.26 8 4 Subdigital lamellae on Finger IV 15-19 18-19 17.00 + 1.41 18.50 + 0.58 8 4 Subdigital lamellae on Toe IV 19-25 23-26 22.88 + 1.96 24.00 + 1.41 8 4 Head length/head width A213 7, 1.20-1.35 1.2 £0.09 1.28 + 0.06 8 4 Head height/head width 0.79-1.07 0.80—-0.88 0.88 + 0.08 0.85 + 0.04 8 4 Body height/body width 0.67-1.04 IEA) 0.88 + 0.14 1.13 + 0.03 7 3 Tail length/total length 0.65—-0.72 0.69-0.74. 0.68 + 0.02 0.71 + 0.02 8 4 Maximum size (mm) 322 393 annual temperature is 18-24°C in both zones; the mean annual precipitation is 2000-4000 mm and 4000-8000 mm, respectively. O'Shaughnessy (1879, 1881) reported specimens from Canelos (1°36' S, 77°45' W, Provincia Pastaza) and Pallatanga (Provincia Chimborazo), Ecuador, and from Moyobamba (Departamento San Martin, Peru). Pallatanga (01°59' S, 78°58' W) is on the western slopes of the Cordil- lera Occidental of the Andes. I consider this locality to be erroneous for two reasons. First, this record would be the only one from the western slopes of the Andes. Second, for the same reason, Pallatanga has been considered to be erroneous for other taxa (e.g., Osteocephalus buckleyi, Prionodactylus manicatus) of the same collection (Trueb and Duellman, 1971; Uzzell, 1973). Etheridge (1966) and Pe- ters and Donoso-Barros (1970) reported Stenocercus aculeatus from Peru, but they did not mention specific ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 9 BES CSS. ane es A See [SN ee: SS SS — reales SS eeeeet ies 1B L Ds eI Sy LET Se Fig. 3. Dorsal and lateral views of the heads of two species of Stenocercus. A and C S. aculeatus, EPN 4051, male. B and D S. angel, holo- type, QCAZ 3733, male. Damaged tissue is stippled. Scale bars = 5 mm. locality. Avila-Pires (1995) and Cadle (1991) reported this species from several localities in Provincia Pastaza, Ecuador. Stenocercus angel new species Holotype —OQCAZ 3733, an adult male, from 8 km NE El Angel on road to Tulcan (00°40' N, 77°52' W), Provincia Carchi, Ecuador, collected on 17 May 1997 by Jaime A. Chaves and Juan M. Guayasamin. The holotype is situ- ated in the Museo de Zoologia, Pontificia Universidad Catolica del Ecuador. Paratypes—OQCAZ 1354, 3732, 4117-4119 with same locality data as holotype; QCAZ 1358, El Angel (00°38' N, 77°56' W, 3015 m), Provincia Carchi; QCAZ 3777, Estacion Biologica Guanderas, Provincia Carchi; QCAZ 1322, El Playon de San Francisco (00°38' N, 77°38' W, 3300 m), Provincia Sucumbios; OCAZ 3792, 13.6 km W Tulcan on road to Tufino (00°49' N, 77°49' W, 3040 m), Provincia Carchi. Diagnosis ——(1) Maximum total length in males 214 mm (n = 8); (2) maximum total length in females 155 mm (n = 2); (3) vertebral scales 39-52; (4) scales around mid- body 52-60; (5) internasal scales 1-4; (6) gular scales 20- 25; (7) lamellae on Finger IV 13-22; (8) lamellae on Toe IV 20-28; (9) posthumeral pocket Type 2; (10) postfemoral pocket Type 2; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, keeled or wrinkled, juxtaposed; (14) ventral scales smooth or slightly keeled, imbricate; (15) scales on posterior surfaces of thighs keeled, imbricate; (16) antehumeral fold absent; (17) tail not laterally compressed; (18) gular region of males black in 25% of the specimens; (19) dorsum green or dark brown in males and dark brown in females. Stenocercus angel is distinguished from other species of Stenocercus by the combination of the following charac- ters: smooth ventrals, imbricate scales on posterior sur- face of thigh, posthumeral pocket Type 2, postfemoral pocket Type 2, 52-60 scales around midbody, and dark green or dark brown background color. Description of holotype —Head (Fig. 3B and D) wider than high (HH/HW = 0.93); occipitals, parietals, interpa- rietal, and postparietals small, keeled or wrinkled, juxta- posed; 2 postrostrals, each wider than long; 4 internasals; 2 canthals on each side, anterior one in contact with nasal; supraoculars keeled or wrinkled, juxtaposed; enlarged supraocular scales absent; lateral temporals keeled, slightly imbricate; parietal eye visible; gulars large, smooth, im- bricate, bearing apical pit; mental in contact with first infralabials and first postmentals. Body wider than high (BH/BW = 0.78); dorsal and lat- eral scales of body and neck keeled, imbricate, slightly mu- cronate; vertebrals large, forming prominent, serrate, ver- tebral crest; ventrals smooth or slightly keeled, imbricate; preauricular fringe moderately developed; neck folds ab- sent; dorsal scales and proximal ventral scales on forelimbs keeled, imbricate; dorsal scales of hands and distal ven- tral scales of forelimbs smooth, imbricate; palmar scales keeled, imbricate, mucronate; dorsal scales on hind limbs, plantar scales, and scales on posterior surface of thigh keeled, imbricate; ventral scales on hind limbs smooth, imbricate; 17 lamellae on Finger IV; 25 lamellae on Toe IV; tail not compressed; dorsal and lateral caudal scales keeled, imbricate, mucronate; anterior ventral caudal scales smooth, imbricate, bearing apical pit; posterior ventral caudal scales keeled, imbricate; posthumeral and postfemoral pockets Type 2. Part of tail regenerated; there- fore, TL/ToL not calculated. In life, the holotype had the following color features: dorsum dark green with 8-10 transverse brown stripes over vertebral line, from neck to anterior part of tail; dor- sum of head and tail dark brown; gular region black; first two pairs of genials and first infralabials gray; ventral sur- face of body, limbs, and tail with irregular yellow or green spots; iris copper (Fig. 4A, B). Data on holotype: Male; SVL = 82 mm; TL = 70 mm; RT = 32 mm; HW = 15.3 mm; HL = 18.4 mm; HH = 14.3 mm; BW = 22.1 mm; BH = 17.3 mm; SM = 58; VS = 48; PS = 56; SS = 5; IS = 4; GS = 23; SF = 17; ST = 25. Variation.—Measurements and scutellation of Stenocercus angel in Table 1. Head wider than high (HH/ HW = 0.78-0.93, x = 0.88 + 0.05, n = 10); 1-4 internasals, usually 2 (50% of the specimens); parietal eye visible in 90% of the specimens; body wider than high (BH/BW = ScrentiFic Papers, NATURAL History Museum, THE UNIVERSITY OF KANSAS Stenocercus aculeatus, KU 121092, female, 82 mm SVL (WED). Stenocercus festae, KU 134608, male, 83 mm SVL (WED). Fig. 4. Four species of Stenocercus. ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS fal ne i Stenocercus guentheri, QCAZ 4153, male, 72 mm SVL (LAC). Stenocercus guentheri, QCAZ 4108, male, 70 m i m SVL (LAC). Stenocercus humeralis, KU 121138, male, 74 mm SVL (WED). (Seon IN ae Pus le d * e git a os aw i i os a AVA SSR NRE ss Stenocercus iridescens, KU 164168, male, 78 mm SVL (WED). Stenocercus ornatus, KU 121126, male, 73 mm SVL (WED). Stenocercus simonsii, KU 134165, female, 72 mm SVL (WED). Stenocercus varius, QCAZ 2015, male, 69 m m SVL (LAC). Fig. 5. Six species of Stenocercus. 12 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS 81° 80° -9 20 40 60 80 100 Yt et fd Kilometers [==] <300m 300-2000 m 2000-4000 m >4000 m Permanent snow VS. aculeatus @S. angel @S. carrioni AS. chota HS. festae CG ot Fig. 6. Distribution of five species of Stenocercus in Ecuador. 0.71-0.97, x= 0.83 0.09, n = 9); ventral scales on proximal parts of forelimbs smooth and imbricate in some speci- mens; 13-22 lamellae on Finger IV; 20-28 lamellae on Toe IV; tail length 62-65% of total length. Color in life: The background color in Stenocercus angel varies between dark green and dark brown. One male paratype (QCAZ 4117) with the same pattern as the holo- type differs from it by having a dark brown dorsum. Another male paratype (QCAZ 3777) differs from the holotype in the following color features: dorsum of head green with small black spots; black stripe extending from inferior edge of orbit to superior border of tympanum; some dorsal scales blue; yellow vertical lines on flanks; gular region bluish green; venter grayish blue; throat yel- low; yellow anal mark extending onto ventral surface of hind limbs and anterior ventral portion of tail. A female (QCAZ 3732) has a dark brown dorsum with 8-10 yellowish-brown transverse stripes and a yellowish- brown venter with scattered dark brown flecks. Two sub- adults (OQCAZ 4118, 4119) differ from adults in having a pair of lateral whitish stripes on each side of body. One stripe extends posteriorly from the angle of the mouth below the tympanum to insertion of forelimb. The other stripe extends posteriorly from the eye over the tympa- num to the insertion of hind limb, where it becomes nearly inconspicuous. Also, in QCAZ 4119, the gular region is bright yellow, and in QCAZ 4118, there is a narrow black vertebral line over the tail. Color in preservative: Dorsum of males dark brown or greenish blue with 8-10 pale brown, greenish-brown, or black, short transverse stripes middorsally, from neck to anterior part of tail; head pale brown, dark brown, or ol- ive-green; gular region cream, pale green, or black; venter cream or pale yellow with pale green; some ventral scales with black borders; pale gray longitudinal midventral stripe in 37% of the specimens. Dorsum of females pale brown; head dark brown; gular region brown with scat- tered yellowish-brown, dark brown, or yellowish-green marks; venter pale green or yellow with scattered dark brown flecks; tail dark brown. Natural history —One female (QCAZ 3732) collected on 17 May 1997 contained two oviductal eggs. The sizes of these eggs are 16.1 x 9.46 mm and 16.23 x 9.39 mm, their volumes are 754.4 mm? and 749.3 mm‘°, respectively. The smallest juveniles (QCAZ 4118, 4119) were collected in January 1999; each has a total length of 138 mm (SVL = 52 mm, TL = 86 mm and SVL = 48, TL = 90, respectively). Distribution and ecology.—Stenocercus angel is known from elevations of 3015-3560 m in the northern Andes (Nudo de Pasto). The species inhabits the upper drainage systems of the Rio Aguarico (Atlantic Drainage) and Rio Mira (Pacific Drainage) in Provincia Sucumbios and Provincia Carchi, respectively (Fig. 6). The distribution lies mainly within Humid Montane Forest, Very Humid Mon- tane Forest, and Low Humid Montane Forest life zones. The mean annual temperature is 7- 12°C in the former two life zones and 12-18 °C in the third. The mean annual pre- cipitation is 500-1000 mm in the Humid Montane Forest and 1000-2000 mm in the Very Humid and Low Humid Montane Forests. Most specimens were found in undisturbed areas of paramo. They were basking on the ground near plants of the thorny bromeliad genus Puya , in which they hide when disturbed. Specimen QCAZ 4118 was sleeping near a leptodactylid frog (Eleutherodactylus curtipes; QCAZ 12820) at the base of a plant genus Cortaderia (Poaceae) late in the afternoon. Etymology.—The specific name is a noun in apposi- tion and refers to the Paramo del Angel, where the species was found initially. This paramo is considered to be one of the most humid Ecuadorian paramos, and it includes one of the few populations of Espeletia (Asteraceae) in Ecua- dor (Coello, 1994); Espeletia is endemic to the northern Andes. ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 13 Stenocercus carrioni Parker Stenocercus carrioni Parker, 1934:268. Holotype: BM 1933.6.24.75, a male from “Zamora, 3250 ft (= 990.6 m), [Provincia Zamora Chinchipe] Ecuador.” Diagnosis.—(1) Maximum SVL in males 74 mm (Cadle, 1991); (2) maximum SVL in females 71 mm (Cadle, 1991); (3) vertebral scales 55-72; (4) scales around mid- body 66-96; (5) internasal scales 4; (6) gular scales 37-52; (7) lamellae on Finger IV 23-28; (8) lamellae on Toe IV 27- 32; (9) posthumeral pocket Type 1 or 2; (10) postfemoral pocket Type 3; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, smooth or slightly wrinkled, juxtaposed; (14) ventral scales smooth, imbricate; (15) scales on posterior surfaces of thighs granular; (16) antehumeral fold well developed; (17) tail not or only slightly compressed later- ally; (18) gular region in males not black; (19) dorsum greenish brown in males. Stenocercus carrioni differs from other species of Stenocercus by the combination of the following charac- ters: smooth ventrals, granular scales on posterior surface of thigh, distinct vertebral row on neck and at least on an- terior part of the body, 2 caudal whorls per autotomic seg- ment, distal portion of tail with caudal whorls alternating in size, posthumeral pocket Type 1 or 2, postfemoral pocket Type 3, 66-69 scales around midbody, 23-28 lamellae on Finger IV, no black collar in front of forelimbs, and no dif- ferences in pattern between sexes. Description and variation —Cadle’s (1991) descrip- tion is sufficient. Measurements and scutellation of S. carrioni summarized in Table 1 were taken from Cadle (1991), who also commented on the holotype. Distribution and ecology.—Stenocercus carrioni occurs at elevations of 1320-1900 m in southern Ecuador. It in- habits the upper valley of the Rio Chira (Pacific Drainage) in Provincia El Oro and Provincia Loja (Fig. 6). The distri- bution lies within the Humid Premontane Forest, where the mean annual precipitation is 1000-2000 mm and the mean annual temperature is 18-24°C. Stenocercus carrioni is sympatric with S. limitaris in Alamor (04 ‘02'S, 80°02' W, 1325 m), Provincia Loja. Fritts (1974) mentioned the possi- bility that S. carrioni occurs in sympatry with S. ornatus. Remarks.—The holotype of Stenocercus carrioni is the only specimen that has been reported from the eastern slopes of the Cordillera Oriental of the Andes (Zamora 0404'S, 78°56' W). All other specimens are from the west- ern slopes of the Cordillera Occidental. Fritts (1974) ques- tioned the authenticity of Zamora as the type locality of S. carrioni because some other taxa recorded by Parker (1934) from Zamora (e.g., Atelopus and Pholidobolus) have been collected subsequently only from higher elevations. The holotype of S. carrioni is part of a collection obtained by Clodoveo Carrion (an Ecuadorian naturalist) deposited in the British Museum (Natural History). The Atelopus in the collection could be either Atelopus boulengeri or an undescribed species (Luis A. Coloma, pers. comm.), and the Pholidobolus in the collection could be either P. annectens or P. macbrydei, taxa known from Provincia Loja and Provincia Zamora-Chinchipe (Montanucci, 1973). Montanucci (1973) did not report any species of Pholidobolus from Zamora, but he did not check specimens in the British Museum. On the other hand, Parker (1934) recorded from Zamora other species, such as Anolis fuscoauratus and A. ortonit, that are known to occur there. Although Cadle (1991) did not question Zamora as the type locality of S. carrioni, the presence of the species at Zamora is highly questionable. Stenocercus chota new species Holotype.—QCAZ 3768, an adult male, from the Panamerican Highway 5 km E Chota (00°28' N, 78°01' W), Valle del Chota, Provincia Imbabura, Ecuador, collected on 8 July 1997 by Luis M. Diaz and Omar Torres-Carvajal. The holotype is situated in the Museo de Zoologia, Pontificia Universidad Cato6lica del Ecuador. Paratypes.—QCAZ 2768, 3755, 3757, 3762-3776, with same locality data as holotype; QCAZ 806, 897-902, 3791, 3794, Ambuqui—Monte Olivo Road, 6.5 km E Panamerican Highway, 00°25' N, 77°55’ W, 1940 m, Provincia Imbabura; QCAZ 799, Ambuqui (00°27' N, 78°01' W, 1780 m), Provincia Imbabura; OCAZ 2654, 2655, 2773-2778, Chota (00°28' N, 78°04' W), Provincia Imbabura; EPN 5848, in the vicinity of Lago Yaguarcocha, Provincia Imbabura; EPN 5858-5860, 5862-5864, Rio Cachabi, Provincia Esmeraldas; MZUT R2154.1-5, R2154.25-29, La Concepcion (00°35' N, 78 °07' W, 1575 m), Provincia Carchi. Diagnosis.—(1) Maximum total length in males 207 mm (71 = 11); (2) maximum total length in females 172 mm (n = 21); (3) vertebral scales 39-50; (4) scales around mid- body 45-59; (5) internasal scales 2-4; (6) gular scales 18- 25; (7) lamellae on Finger IV 14-20; (8) lamellae on Toe IV 23-31; (9) posthumeral pocket Type 2; (10) postfemoral pocket Type 2; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, keeled or wrinkled, juxtaposed; (14) ventral scales smooth or slightly keeled, imbricate; (15) scales on posterior surfaces of thighs keeled, imbricate; (16) antehumeral fold absent; (17) tail not laterally compressed; (18) gular region of males not black; (19) dorsum grayish brown or pale brown in males and females. The following combination of characters distinguish Stenocercus chota from other species of Stenocercus: smooth ventrals, imbricate scales on posterior surface of thigh, posthumeral pocket Type 2, postfemoral pocket Type 2, 45-59 scales around midbody, large black spots in gular 14 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS ne Ws Be rk i Fig. 7. Dorsal and lateral views of the heads of two species of Stenocercus. A and C S. chota, holotype, QCAZ 3768, male. B and D S. festae, QCAZ 1347, male. Scale bars = 5 mm. region of females, and large blue longitudinal mark on each side of dark midventral stripe in males. Description of holotype.—Head (Fig. 7A and C) wider than high (HH/HW = 0.82); occipitals, parietals, interpa- rietal, and postparietals small, keeled, and slightly imbri- cate; 4 postrostrals, 2 wider than long and 2 as wide as long; 4 internasals; 2 canthals on each side, the most ante- rior separated from the nasal by 2 tiny scales; supraoculars keeled or wrinkled and juxtaposed; row of enlarged supraocular scales absent; lateral temporals slightly keeled and imbricate; parietal eye visible; gulars large, smooth, and imbricate each with an apical pit; mental in contact with the first infralabials, first postmentals and with a tiny scale between postmentals. Body wider than high (BH/BW = 0.78); dorsal and lat- eral scales of body and neck keeled, imbricate, and slightly mucronate; vertebrals large, forming a prominent serrate vertebral crest; ventrals smooth and imbricate; preauricu- lar fringe well developed; neck folds absent; limb scales keeled and imbricate except the ventral scales of hind limbs that are smooth and imbricate; dorsal scales of hind limbs and plantar scales mucronate; 20 lamellae on Finger IV; 28 lamellae on Toe IV; tail laterally compressed; caudals keeled and imbricate except those of the anterior ventral region that are smooth and imbricate; tail length 62% of total length; posthumeral and postfemoral pockets Type 2. In life, the coloration of the holotype was: dorsum gray- ish brown with 12 transverse brown stripes longitudinally disposed over vertebral line, from neck to anterior part of tail; white and dark brown spots on dorsum of body and flanks; limbs with more white spots than dorsum and with brown narrow transverse lines (5-6 on forelimbs and 8-10 on hind limbs); dorsum of head dark gray; sides of head white with gray spots; gular region reddish cream; small dark midventral longitudinal stripe extending from throat to point before insertion of hind limbs; large blue longitu- dinal mark on each side of dark midventral stripe; rest of ventral surface faintly orange with several tiny brown spots; yellow anal mark extending to ventral surfaces of thighs and anterior ventral surface of tail; ventral surface of tail cream; iris bronzed green. Data on holotype: Male; SVL = 69 mm; TL = 114 mm; HW = 14 mm; HL = 16.8 mm; HH = 11.5 mm; BW = 19 mm; BH = 14.9 mm; SM = 59; VS = 49: PS = 71;SS =5; IS= 4; GS = 22; SF = 20; ST = 28. Variation.—Measurements and scutellation of - Stenocercus chota in Table 1. Head wider than high (HH/ HW = 0.67-0.97, x = 0.84 + 0.06, n = 41); usually 2 postrostrals, wider than long; 2-4 internasals, usually 3 (42% of the specimens) or 4 (42%); the most anterior can- thal in contact with the nasal in some specimens; parietal eye visible in 87% of the specimens; lateral temporal scales juxtaposed in some specimens; body wider than high (BH/ BW = 0.42-1.04, X = 0.69 + 0.16, n = 39); ventrals slightly keeled in some specimens; anterior ventral scales of tail rarely keeled; tail not compressed in females; lamellae on Finger IV 14-20; lamellae on Toe IV 23-31; tail length 55- 66% of total length; posthumeral and postfemoral pockets Type 2. Sexual variation in measurements and scutellation in S. chota is presented in Table 3. Color in life: Two male paratypes (QCAZ 3775, 3776) differ slightly in coloration from the holotype by having yellow spot in each axilla; dark midventral stripe indis- tinct; yellow anal mark posteriorly delineated by black; iris bronzed red. Also, a dark midventral longitudinal stripe extends onto the ventral surfaces of the thighs in OCAZ 3755. Marked sexual dichromatism exists in Stenocercus chota (Fig. 4C, D, E, F). Females have a cream ventral surface of body without markings (dark midventral stripe in males) and several large black spots in the gular region (absent in males). The sides of the head are gray with cream and black marks (sides of head white with gray marks in males). There is a longitudinal black stripe ex- tending from the eye to a point that corresponds to the posterior border of the occipital region; another black stripe extends from the comisure to the tympanum (those stripes are absent in males). The dorsum usually is pale brown in both sexes. Color in preservative: Ground color of dorsum of head and body dark brown or gray, dark brown transverse stripes on dorsum; rostrum white, cream, or gray; pale gray, venter reddish cream or cream in females; gular region of females with large black spots; dark midventral stripe and blue ventral marks indistinct or absent in males. ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 15 Table 3. Sexual variation in scutellation and measurements of Stenocercus chota. Range (first line), x + SD (second line), and n (third line) are given. Character Females Males Scales around midbody 45-56 46-59 50.62 + 2.69 51.91 + 4.25 21 11 Vertebral scales 39-50 42-49 45.20 + 3.29 45.64 + 2.58 20 11 Paravertebral scales 55-71 58-74 61.30 + 4.56 64.64 + 4.99 20 11 Gular scales 18-23 19-23 20.57 + 1.57 20.64 + 1.36 Dil 11 Supraocular scales 4-6 5-6 5.29 + 0.56 5.36 + 0.50 21 11 Internasal scales 2-4 2-4 3.38 + 0.74 3.27 + 0.65 21 11 Subdigital lamellae of Finger IV 14-19 17-20 W714! se 1345) 18.64 + 1.03 21 11 Subdigital lamellae of Toe IV 24-30 23-28 26.05 + 1.28 26.09 + 1.45 21 11 Headlength/head width 1.29-1.39 1.17-1.25 1.29 + 0.04 1.21 + 0.03 Dil 10 Head height/head width 0.76-0.94 0.77-0.90 0.84 + 0.05 0.84 + 0.05 21 10 Body height/body width 0.50-0.94 0.63-1.04 0.69 + 0.15 0.83 + 0.13 21 9 Regenerated tail length/tail length 0.59-3.07 0.55-0.63 1.57 + 1.00 0.26 + 0.10 6 2 Tail length/total length 0.58-0.65 0.55—0.63 0.63 + 0.02 0.61 + 0.03 Maximum size (mm) W772: 207 Natural history —One female (QCAZ 3767) collected on 8 July 1997 contained two oviductal eggs. The sizes of these eggs are 16.59 x 7.81 mm and 15.95 x 7.67 mm; their volumes are 529.8 mm® and 491.3 mm’, respectively. The smallest individual (QCAZ 902) was collected on 4 June 1989 and has a total length of 83 mm (SVL = 30 mm, TL = 53 mm). Distribution and ecology.—Stenocercus chota is known from elevations of 1575-1940 m in the inter-Andean val- ley El Chota (Ibarra Basin) in northern Ecuador. It inhabits the upper valley of the Rio Mira (Pacific Drainage) in Provincia Carchi and Provincia Imbabura (Fig. 6). Six speci- mens (EPN 5858-5860, 5862-5864), purportedly from Rio Cachabi, Provincia Esmeraldas, were not used in the dis- tribution analysis. Moreover, the upper valley of the Rio Cachabi (Pacific Drainage) is approximately at 800 m about 55 km west of the closest locality confirmed for S. chota (La Concepcion, Provincia Carchi, 00°35' N, 78°07' W, 1575 m). The distribution lies mainly within the Thorny Premontane Thicket, where the mean annual temperature is 18-24°C, and the mean annual precipitation is 250-500 mm. Most individuals observed in the field were basking. These lizards escape by running directly into holes among stones, vegetation thickets, fallen logs, or branches, or they hide in the base of thorny shrubs. Specimens were found in undisturbed areas as well as very disturbed areas such as sugar cane plantations. A colubrid snake Drymarchon corais (QCAZ 3753), with the same collection data as the holotype of Stenocercus chota, contained a piece of tail and two eggs similar in shape and size to the eggs of S. chota. Etymology.—The specific name is a noun in apposi- tion and refers to the Chota Valley (Ibarra Basin), inhab- ited by this species. Stenocercus festae (Peracca) Liocephalus festae Peracca, 1897:6. QCAZ 4059, an adult male from Sevilla de Oro (02°48' S, 78°39' W), 2630 m, Provincia Azuay, Ecuador, col- lected on 17 September 1998 by P. Menéndez and P. Salazar, herein designated as neotype. Leiocephalus ornatus ornatus (Part)—Burt and Burt, 1931:271. Ophryoessoides festae—Etheridge, 1966:88. Stenocercus festae—Fritts, 1974:49. Comments on the neotype.—The description of Stenocercus festae was based on three specimens from Cuenca, Provincia Azuay (Peracca, 1897). Peters (1967) er- roneously referred to specimen TurM (MZUT) 2619 as the holotype of Stenocercus festae. MZUT 2619 is a no longer valid number that corresponds to MZUT R2158.1-62, a se- ries of 62 specimens reported by Peracca (1904) from La Concepcion, Provincia Imbabura (Franco Andreone, pers. comm.). Peracca (1904) misidentified these specimens as Stenocercus festae; | checked them and they correspond to Stenocercus chota. According to Fritts (1974), the type ma- terial of Stenocercus festae corresponds to three syntypes deposited in the Torino Regional Museum of Natural His- tory with unknown MZUT numbers. Nonetheless, the type material of S. festae cannot be located and a name-bearing type is necessary to define this taxon objectively. Diagnosis.—(1) Maximum total length in males 241 mm (71 = 10); (2) maximum total length in females 206 mm (n = 22); (3) vertebral scales 39-55; (4) scales around mid- body 47-64; (5) internasal scales 2-4; (6) gular scales 16- 29; (7) lamellae on Finger IV 14-22; (8) lamellae on Toe IV 21-33; (9) posthumeral pocket Type 2 or 3; (10) postfemoral pocket Type 3; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, keeled, juxtaposed; (14) ventral scales smooth or slightly keeled in adults and keeled in juveniles, imbri- cate; (15) scales on posterior surfaces of thighs keeled, im- bricate; (16) antehumeral fold absent; (17) tail not or slightly compressed laterally; (18) gular region of males may be 16 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS black; (19) dorsum pale green. Stenocercus festae differs from other species of Stenocercus by the combination of the following charac- ters: smooth ventrals (slightly keeled in juveniles), imbri- cate scales on posterior surface of thigh, no enlarged supraoculars, posthumeral pocket Type 2 or 3, and postfemoral pocket Type 3. Description of neotype-——Head higher than wide (HH/HW = 1.07); occipitals, parietals, interparietal, and postparietals small, keeled, and juxtaposed; 2 postrostrals; 4 internasals; 2 canthals, the most anterior in contact with the nasal; supraoculars keeled; enlarged supraoculars ab- sent; lateral temporals keeled and slightly imbricate; pari- etal eye visible; gulars smooth and imbricate, bearing an apical pit; mental in contact with first pair of infralabials and first pair of postmentals. Body wider than high (BH/BW = 0.99); dorsal and lat- eral scales of body and neck strongly keeled, imbricate, and mucronate; vertebrals large, forming a prominent ser- rate vertebral crest (Fig. 8); ventrals smooth and imbricate, bearing an apical pit; preauricular fringe nearly inconspicu- ous; neck folds absent; dorsal scales of hand, proximal ventral scales of forelimbs, and ventral scales of hind limbs smooth and imbricate; rest of limb scales imbricate and keeled; 22 lamellae on Finger IV; 31 lamellae on Toe IV; dorsal caudals keeled, imbricate, and mucronate; ventral caudals smooth and imbricate proximally and keeled and imbricate distally; tail length 70% of total length; posthumeral and postfemoral pockets Type 3. Data on neotype: Male (Fig. 8); SVL = 81 mm; TL = 164 mm; HW = 14.32 mm; HL = 19.83 mm; HH = 15.39 mm; BW = 17.15 mm; BH = 17.11 mm; SM = 62; VS = 49; PS = 65; SS = 63 1S S42 ES = 28s SF = 22 Sl = Bil. Variation.—Measurements and scutellation of Stenocercus festae in Table 1. Head (Fig. 7B and D) usually wider than high (HH/HW = 0.75-1.16, x = 0.86 + 0.07, n = 42); 2-3 postrostrals; 2—4 internasals, usually 4 (56%); pari- etal eye visible in 77% of the specimens; body usually wider than high (BH/BW = 0.63-1.18, x = 0.83 + 0.12, n = 31); ventrals smooth or slightly keeled in adults and keeled in juveniles; 14-22 lamellae on Finger IV; 21-33 lamellae on Toe IV; tail length 52-71% of total length; posthumeral pocket Type 2 or 3, normally Type 3 (80%). Sexual variation in measurements and scutellation are given in Table 4. Color in life: Dorsum of males grayish tan, grayish brown, or pale olive brown with roughly paired irregular green or yellow transverse stripes; flanks black with green or yellow flecks and blotches; antehumeral region black in some specimens; dorsal surfaces of limbs with yellow or green flecks or spots; supralabials and infralabials black; loreal and infraorbital scales yellowish green or yellow; Fig. 8. Stenocercus festae, neotype, QCAZ 4059, male. chin pale yellow or pale yellowish green; throat black; wide black midventral stripe; rest of venter yellowish green or pale yellow with black flecks; in some individuals, entire ventral surface of body black; in some specimens, anal re- gion, ventral surfaces of hind limbs, and anterior ventral surface of tail yellow; ventral surface of thighs heavily suffused with black in some individuals; (Fig. 4H; Tho- mas H. Fritts, field notes, 5 and 8 June 1970). Dorsum of females brown with or without short dark transverse stripes on vertebral line; pale gray dorsolateral stripe present or not on each side; chin and chest yellow; venter pinkish white or pale yellow, rarely with gray flecks or blotches (Thomas H. Fritts, field notes, 5 and 20 June 1970). Juveniles have grayish-beige dorsum with numerous dark brown dorsal blotches; beige dorsolateral stripe on ante- rior half of body; whitish-beige infraorbital stripe; chin and venter pale gray with slight to extensive dark reticulation (Thomas H. Fritts, field notes, 8 and 20 June 1970). Color in preservative: Dorsum in males dark brown or gray with dark brown transverse bars middorsally to tip of tail; black spot present or not on shoulder; limbs dark brown dorsally with or without black and cream reticula- tion; head dark brown, speckled with black in some indi- viduals; supralabials and infralabials dark brown or black; black line extending from posterior border of orbit to tym- panum; black spot behind eye; gular region usually white or cream, or with black spots; black band on throat; pecto- ral region cream speckled with black and brown; wide or narrow black midventral stripe; ventral surface of limbs and tail cream, with black or brown spots in some indi- viduals. Dorsum of females dark brown or gray, with or ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 17 Table 4. Sexual variation in scutellation and measurements of Stenocercus festae. Range (first line), x= SD (second line) and n (third line) are given. Character Females Males Scales around midbody 48-63 47-64. 54.64+4.15 56.6 + 5.70 22 10 Vertebral scales 39-55 43-53 46.59 + 4.06 47.30 +3.4 22 10 Paravertebral scales 46-72 55-67 62.00 + 5.68 61.30 + 4.74 22 10 Gular scales 16-29 21-27 22 PY] xe DI 23.60 + 1.96 22 10 Supraocular scales 4-6 46 5.05 + 0.49 5.10 + 0.57 22 10 Internasal scales 2-4 2-4 3.55 + 0.67 3.40 + 0.7 22 10 Subdigital lamellae on Finger IV 14-22 15-21 17.45 + 1.82 17.44+2.19 22 9 Subdigital lamellae on Toe IV 21-33 24-30 26.45 + 2.63 26.56 + 2.07 22 9 Head length/head width 1.19-1.43 1.14-1.29 1.30 + 0.05 1.24 + 0.05 21 10 Head height/head width 0.75-0.95 0.77-1.16 0.84 + 0.04 0.89 + 0.11 21 10 Body height/body width 0.63-1.10 0.71-1.18 0.80 + 0.11 0.89 + 0.16 17 9 Regenerated tail length/tail length 0.27-2.24 0.66-4.14 1.12 + 0.96 2.08 + 1.71 5 4 Tail length/total length 0.52-0.71 0.63—0.67 0.63 + 0.06 0.65 + 0.02 11 4 Maximum size (mm) 206 241 without black spots; dorsum with several transverse bars that are wider and shorter than those of males; dorsal sur- face of limbs with dark brown or black spots but never reticulate as in males; venter cream with several short lon- gitudinal gray flecks which can extend to gular region. Distribution and ecology.—Stenocercus festae occurs at elevations of 2300-3200 m in the Cuenca and Saraguro Basins in southern Ecuador. It inhabits the upper valleys of the Rio Paute (Atlantic Drainage) and Rio Jubones (Pa- cific Drainage) in Provincia Azuay, Provincia Canar and Provincia Loja (Fig. 6). The distribution lies within the Dry Low Montane Forest, Humid Montane Forest, and Subandean Humid Forest (= Subalpean Humid Forest of Canadas-Cruz [1983]) life zones. The mean annual tem- perature in these zones is 12-18°C, 7-12°C, and 6-13°C, respectively. The mean annual precipitation is 500-1000 mm in the first two life zones and 250-500 mm in the third. Stenocercus festae and S. simonsii occur in sympatry in the upper valley of the Rio Jubones (Fritts, 1974). In this area, S. festae utilizes the leaves of agave plants and sur- rounding ground surface for basking and feeding, whereas S. simonsit is restricted to large rocks or rock piles (Fritts, 1974). Also, S. festae possibly occurs in sympatry with S. rhodomelas in the Saraguro Basin. Stenocercus guentheri (Boulenger) Liocephalus guenther1 Boulenger 1885:169. Syntypes: BM 58.7.25.16-18; 59.9.20.6; 60.6.16.18-21; 71.2.7.7-10; 71.4.16.53; 80.12.8.53, from “Guayaquil [Provincia Guayas, Ecuador], Sarayacu [Provincia Pastaza] Ecuador, Western Ecuador, and Colombia” (restricted to San Antonio de Pichincha, 2500 m, Provincia Pichincha, by Fritts [1974]). Leiocephalus ornatus ornatus (Part)—Burt and Burt, 1931:271. Ophryoessoides guentheri—Etheridge, 1966:88. Stenocercus guentheri—Fritts, 1974:54. Comments on the type locality —Fritts (1974) men- tioned that Stenocercus guentheri occurs at none of the origi- nal type localities. However, Castro and Granados (1993) reported S. guentheri from southern Colombia. Fritts (1974) restricted the type locality to the Pichincha Basin (San Antonio de Pichincha) because some type specimens, one of which is the syntype herein described, have a black band on the throat, a character that has been found only in males from that basin. Diagnosis.—(1) Maximum total length in males 255 mim (71 = 38); (2) maximum total length in females 188 mm (n = 33); (3) vertebral scales 44-64; (4) scales around mid- body 59-83; (5) internasal scales 2-4; (6) gular scales 21- 31; (7) lamellae on Finger IV 15-23; (8) lamellae on Toe IV 22-36; (9) posthumeral pocket Type 1 or 2; (10) postfemoral pocket Type 1 or 2; (11) projecting angulate temporals ab- sent; (12) row of enlarged supraoculars absent; (13) occipi- tal scales small, wrinkled or keeled, juxtaposed; (14) ven- tral scales smooth, imbricate; (15) scales on posterior sur- faces of thighs keeled, imbricate; (16) antehumeral fold absent; (17) tail not laterally compressed; (18) gular region of males not black; (19) dorsum bronzed-green, olive-green or brownish green. Stenocercus guentheri is distinguished from other spe- cies of Stenocercus by the combination of smooth ventrals, imbricate scales on posterior surface of thigh, no enlarged supraoculars, posthumeral pocket Type 1 or 2, postfemoral pocket Type 1 or 2, and 59-83 scales around midbody. Description of syntype BM 60.6.16.20.—Head wider than high (HH/HW = 0.88); occipitals , parietals, interpa- rietal, and postparietals small, keeled, and juxtaposed; 4 postrostrals, each of which is longer than wide; 3 18 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 4. Sexual variation in scutellation and measurements of Stenocer- cus festae. Range (first line), x + SD (second line) and n (third line) are given. Character Females Males Scales around midbody 48-63 47-64 54.64+4.15 56.6 + 5.70 22 10 Vertebral scales 39-55 43-53 46.59 + 4.06 47.3 +3.40 22 10 Paravertebral scales 46-72 55-67 62.00 + 5.68 61.3 44.74 22 10 Gular scales 16-29 21-27 22.27 £2.71 23.6 + 1.96 22 10 Supraocular scales 4-6 46 5.05 + 0.49 5.10 + 0.57 22 10 Internasal scales 2-4 2-4 3.55 + 0.67 3.40 + 0.7 22 10 Subdigital lamellae on Finger IV 14-22 15-21 17.45 + 1.82 17.44 + 2.19 22 9 Subdigital lamellae on Toe IV ZN 33) 24-30 26.45 + 2.63 26.56 + 2.07 22 9 Head length/head width 1.19-1.43 1.14-1.29 1.30 + 0.05 1.24 + 0.05 21 10 Head height/head width 0.75-0.95 0.77-1.16 0.84 + 0.04 0.89 + 0.11 21 10 Body height/body width 0.63-1.10 0.71-1.18 0.80 + 0.11 0.89 + 0.16 17 9 Regenerated tail length/tail length 0.27-2.24 0.664.114 1.12 + 0.96 2.08 + 1.71 5 4 Tail length/total length 0.52-0.71 0.63-0.67 0.63 + 0.06 0.65 + 0.02 11 4 Maximum size (mm) 206 241 internasals; 2 canthals on each side, the most anterior in contact with the nasal; supraoculars keeled; row of en- larged supraocular scales absent; lateral temporals keeled and imbricate; parietal eye visible; gulars smooth and im- bricate; mental in contact with the first pair of infralabials and first pair of postmentals. Dorsal and lateral scales of body and neck strongly keeled and imbricate; vertebrals large, forming a promi- nent serrate vertebral crest; ventrals smooth and imbricate; preauricular fringe poorly developed; neck folds absent; ventral scales of limbs smooth or slightly keeled, imbri- cate; dorsal scales of limbs, plantars, palmars, and scales on posterior surface of thighs keeled and imbricate; dor- sal hand scales smooth and imbricate; 20 lamellae on Fin- Fig. 9. Stenocercus guentheri, syntype, BM 60.6.16.20, male. ger IV; 31 lamellae on Toe IV; tail slightly compressed; dorsal and lateral scales of tail keeled, imbricate, and slightly mucronate; anterior ventral scales of tail smooth and imbricate; posterior ventral scales of tail keeled and imbricate; vertebral crest extending more than half the length of the tail; tail length 62% of total length; posthumeral and postfemoral pockets Type 2. Data on syntype BM 60.6.16.20: Male (Fig. 9); SVL = 96 mm; TL = 159 mm; HW = 17 mm; HL = 20.3 mm; HH = 15 mm; SM = 69; VS = 59; PS = 89: SS = 6; IS = 3; GS = 26; SF = 20; ST = 31. Variation.—Measurements and scutellation of Stenocercus guenthert in Table 1. Head (Fig. 10A and C) usu- ally wider than high (HH/HW = 0.75-1.05, x= 0.87 £0.07, n = 72); usually 2 postrostrals (58%, 3 in 28%, 4 in 14%), each wider than long; 2-4 internasals (usually 4, 50%); anteriormost canthals seldom (12%) separated from the nasals by one or two tiny scales; parietal eye usually vis- ible (80%); gulars bearing apical pit in (98%); mental in contact with a tiny scale (20%) that medially separates first pair of postmentals; body usually wider than high (BH/ BW = 0.50-1.15, x = 0.81 + 0.14, n = 68); ventrals weakly keeled in 45% of the specimens; vertebral crest most con- spicuous in adult males; 15-23 lamellae on Finger IV; 22— 36 lamellae on Toe IV; tail not or slightly compressed; ver- tebral crest extending more than half the length of the tail only in adult males; tail length 57-72% of total length; posthumeral and postfemoral pockets Type 1 or 2. Sexual and geographic variation in scutellation and measurements among samples of S. guentheri in Tables 5 and 6, respec- tively. Color in life: An adult male (QCAZ 3761) from 7.5 km N Otavalo, Provincia Imbabura: dorsum of head, body, limbs, and tail pale olive-green; several irregular small black transverse bars on vertebral line; numerous pale green scales on flanks and dorsal surfaces of limbs; irregu- ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 19 3 rd > oe Es (] \) <] a» Dy we se Ee LEX Oi Fig. 10. Dorsal and lateral views of the heads of two species of Stenocercus. A and C S. guentheri, QCAZ 775, male. B and D S. haenschi, holotype, ZMB 16595, male. Scale bars = 5 mm. lar black transverse lines on dorsal surface of hind limbs; iridescent pale green gular region; ventral surfaces of body and limbs also pale green, but less iridescent; ventral sur- faces of thighs and anal region pale yellow; ventral sur- face of tail pale yellow proximally and dark yellow dis- tally. Another adult male (QCAZ 4108) from Lloa, Provincia Pichincha, differs noticeably from the former in the fol- lowing characters: dorsum of head dark brown; dorsum of body, limbs and tail dark greenish brown; several small yellow transverse marks on vertebral line and small yel- low dots on flanks and dorsal surface of limbs; one dorso- lateral black stripe on each side extending from the supe- rior edge of tympanum to the base of tail; gular region dark brown, with several yellowish-brown flecks; throat black with small yellow flecks; throat pale yellow with dark brown flecks; dark brown longitudinal midventral stripe extending from throat to point before the insertion of hind limbs; ventral surface of body on each side of midventral stripe bluish gray with several white irregular marks; ven- tral surface of forelimbs and tail dark brown with yellow marks, distal portion of tail with pink marks (Fig. 5C). An adult female (QCAZ 3659) from 14.5 km N Tixan, Provincia Chimborazo: dorsum dark olive-green with scattered small black spots; short transverse black bars on vertebral line; region between flanks and venter intense yellow with sev- eral dark olive-green spots; pale yellow venter with small gray flecks in gular region. A female (QCAZ 3660) from the same locality differs from the former by having whit- ish-cream venter and small gray flecks in mental, gular, and pectoral regions. Color in preservative: Dorsum of head and body in males dark brown, reddish-brown, greenish-brown, or gray; Table 5. Sexual variation in scutellation and measurements of Stenocer- cus guentheri. Range (first line), X+ SD (second line) and n (third line) are given. Character Females Males Scales around midbody 59-82 60-83 67.27 + 4.49 68.79 + 5.25 33 38 Vertebral scales 46-64 46-63 55.67 + 3.80 55.34 + 3.45 33 38 Paravertebral scales 63-89 59-89 74.00 + 5.40 74.74 + 6.42 33 38 Gular scales 21-31 22-30 DRY 3 ANG) 25.89 + 2.09 33 38 Supraocular scales 5-7 5-7 5.61 +0.61 5.84 + 0.49 33 38 Internasal scales 2-4 2-4 3.27 + 0.88 3.24 + 0.82 33 38 Subdigital lamellae on Finger IV 16-22 15-23 17.76 + 1.28 18.66 + 1.74 33 38 Subdigital lamellae on Toe IV 22-34 23-36 26.88 + 2.55 27.92 + 2.27 32 38 Head length/head width 1.13-1.35 1.10-1.33 1.27 + 0.04 1.26 + 0.06 32 38 Head height/head width 0.75-1.01 0.79-1.05 0.85 + 0.06 0.89 + 0.07 32 37 Body height/body width 0.50-1.03 0.59-1.15 0.78 0.13 0.84 + 0.14 31 34 Regenerated tail length/tail length 0.58-3.40 0.11-3.00 2.09 + 1.16 1.44 + 0.90 4 9 Tail length/total length 0.57-0.72 0.58—0.68 0.63 + 0.03 0.65 + 0.02 25 25 Maximum size (mm) 188 255 small black spots on head in 34% of specimens; black band (incomplete medially in 20%) on throat of 56% of the males from Machachi, San Antonio, Quito, Uyumbicho, and Tabacundo (Fritts [1974] referred to this black band as “black patches on the gular region” and he mentioned that these patches can be found only in populations of the Pichincha Basin); gular region gray, reddish cream, or cream; dorsum and flanks with several small cream, blu- ish-green or pale blue spots (92%); short dark transverse bars on vertebral line of body and tail (73%); dark trans- verse lines on dorsal surfaces of limbs (86%); ventral sur- faces of body, limbs, and tail, reddish cream, cream, or gray. Dorsum of head and body in females greenish-brown, brown, or pale gray; short transverse dark bars on verte- 20 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 6. Geographic variation in scutellation and measurements of Stenocercus guentheri. Range (first line), x+ SD (second line) and n (third line) are given. Palmira, Guayllabamba, Machachi, Quito, San Antonio, Picaihua, Character Chimborazo Pichincha Pichincha Pichincha Pichincha Tungurahua Scales around midbody 68-72 64-70 59-70 59-74 60-74 67-83 69.67 + 2.08 67.00 + 2.58 66.00 + 3.16 67.50 + 5.32 67.63 43.80 75.33 + 8.02 3 4 9 6 24 3 Vertebral scales 46-56 53-56 54-57 53-61 53-64 56-63 52.00 + 5.29 54.00 + 1.41 54.67 + 1.00 56.67 + 3.20 56.63 42.83 60.33 £3.79 3 4 9 6 24 3 Paravertebral scales 63-68 71-78 71-81 69-86 70-85 73-82 65.33 + 2.52 73.75 £3.10 Ys}-319)35 3}.3)9) 75.83 + 5.91 76.38 £2.98 76.33 +4.93 3 4 9 6 24 3 Gular scales 25-27 21-29 23-28 24-31 22-30 26-30 26.00 + 1 24.25 + 3.40 Ps Mil ae WP 26.33 £ 2.73 2518324 27.3823 3 4 9 6 24 3 Supraocular scales 5-6 S=7/ 5 5-6 6-7 6-7 5.33 + 0.58 5.75 + 0.96 — 5.50 + 0.55 6.04 + 0.20 6.67 + 0.58 3 4 9 6 24 3 Internasal scales 4 3-4 3-4 2-4 2-4 4 = 3.50 + 0.58 3.78 + 0.44 3.67 + 0.82 2.58 + 0.72 = 4 9 6 24 3 Subdigital lamellae on Finger IV 16-18 17-20 16-19 16-18 16-21 19-22 17.00 + 1.00 18.50 + 1.29 17.11 £0.93 17.50 + 0.84 18:71 2130) 2033 Eeio8 3 4 9 6 24 3 Subdigital lamellae on Toe IV 22-24 27-28 24-29 24-29 24-31 29 PB So ecalello 27.25 + 0.50 25.67 + 1.80 27.00 + 1.79 28.17 + 1.74 = 8 4 9 6 24 3 Head length/head width ESS IeZal 1.27-1.31 1.19=1.33 122-125 22> 1635) 1.23-1.29 1.18 + 0.04 1.29 + 0.02 1.25 + 0.04 1.24 +0.01 1.29 + 0.03 1.26 + 0.03 3 4 9 5 23 3 Head height/head width 0.75-0.81 0.78-1.01 0.78-0.93 0.76-1.05 0.79-1.05 0.81—0.88 0.77 + 0.03 0.90 + 0.10 0.88 + 0.05 0.89 + 0.11 0.88 + 0.07 0.84 + 0.03 3 4 9 5 23 3 Body height/body width 0.59-0.67 0.69—-0.86 0.69-1.15 0.62-0.99 0.50-1.01 0.75—0.79 0.62 + 0.04 0.76 + 0.09 0.89 + 0.14 0.85 + 0.17 0.80 + 0.14 0.78 + 0.03 3 4 9 5 23 3 Regenerated tail length/tail length 2.05 3 = 1.35 0.16-2.33 2.09 — — = = 1.15 + 0.98 = 1 1 — 1 4 1 Tail length/total length 0.63 0.60—-0.65 0.59-0.67 0.63-0.72 0.57—-0.67 0.62-0.63 = 0.63 + 0.02 0.63 + 0.02 0.65 + 0.05 0.64 + 0.03 0.63 + 0.01 1 3 9 4 17 2 Maximum size (mm) 144 166 234 215 215 169 bral line of body and tail (75%); dark transverse lines on dorsal surfaces of limbs (65%); small black spots on head (90%); three parallel black lines on each side of the head (75%), one extending from angle of mouth to inferior edge of tympanum, other extending from inferior border of the orbit to superior edge of tympanum, one from posterior edge of orbit to neck; loreal region cream (85%) or pale brown; venter greenish cream, blue-cream, or cream; gu- lar region normally with reticulate gray or dark brown flecks; small dark flecks on venter (80%). Natural history—One female (QCAZ 713) collected on 17 June 1989 contained two oviductal eggs. The sizes of these eggs are 19.7 mm x 10.2 mm and 19.3 mm x 10.3 mm; their volumes are 1071.1 mm? and 1078.2 mm’, respectively. The smallest individual (QCAZ 737) was collected on 5 June 1988 and has a total length of 72 mm (SVL = 20 mm, TL=52 mm). Fritts (1974) reported one female covering a small hole containing two eggs on 11 June 1970 near Guamote (Provincia Chimborazo) and 18 females that de- posited two eggs each in the laboratory between 16 May and 18 June 1972. Distribution and ecology.—Stenocercus guentheri oc- curs at elevations of 2135-3890 m in the Ecuadorian Andes, especially in the Cordillera Occidental and the Alausi, Latacunga, Otavalo, Quito, and Riobamba inter-Andean basins (Fig. 11). It inhabits the upper valleys of several riv- ers of the Pacific (Rio Mira, Rio Esmeraldas, and Rio Guayas) and Atlantic (Rio Pastaza and Rio Paute) drain- ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS Ail 81° 80° 7° 20 40 60 80 100 ee Kilometers © Permanent snow Fig. 11. Distribution of three species of Stenocercus in Ecuador. ages in the following Provincias: Chimborazo, Cotopaxi, Imbabura, Pichincha, and Tungurahua. The distribution lies within the Low Humid Montane Forest, Low Dry Montane Forest, Low Thorny Montane Steppe, and Mon- tane Steppe life zones. The mean annual temperature is 12-18°C in the former three life zones and 7-12°C in the fourth. The mean annual precipitation is 1000-2000 mm in Low Humid Montane Forest, 500-1000 mm in Low Dry Montane Forest, and 250-500 mm in the latter two zones. Fritts (1974) gave Sevilla de Oro (02°48' S, 78°39' W, 2630 m; Provincia Azuay) as the southernmost locality for Stenocercus guentheri. However, I examined two of the specimens from Sevilla de Oro that Fritts (1974) checked (JAP 6522, 6525, 6527, 6528, 6530, USNM 201222, 201223) and they correspond to S. festae. Furthermore, the neotype of S. festae was collected in Sevilla de Oro. Stenocercus guentheri also has been reported from the Andes of southern Colombia (Ayala and Castro, 1982; Castro and Granados, 1993; Corredor, 1983), where it oc- curs at elevations of 2000-3900 m in departamentos of Narino and Cauca (Castro and Granados, 1993). Males tend to occupy rocks and other elevated posi- tions for basking and feeding, whereas females prefer the ground (Fritts, 1974). At localities lacking exposed rocks or rock walls, both sexes seek refuge in holes in the ground at the bases of plants such as Agave and Stipa (Fritts, 1974). De Vries et al. (1983) reported S. guentheri as one of the main prey items of the Andean falcon Phalcoboenus carunculatus (Carunculated Caracara). Remarks.—Fritts (1974) described some of the consid- erable geographic variation in Stenocercus guentheri. He noted that the number of scales around midbody is higher in some populations in the Cotopaxi and Chimborazo Ba- sins than in those in the Imbabura and Pichincha Basins. Fritts (1974) also mentioned that males from the Imbabura and Pichincha Basins have a narrow yellow midventral stripe with lateral areas of venter bluish to greenish gray, whereas males from the Cotopaxi and Chimborazo Basins have yellow venters with numerous dark spots, and the population from Alausi (southern Chimborazo Basin) in- cludes males with reddish-orange, pale yellow, or bluish- green venters. However, a specimen (QCAZ 4153) from Pusuqui (Pichincha Basin) has a reddish-orange venter with a yellow midventral stripe (Fig. 5A, B), and a speci- men (QCAZ 4108) from Lloa (Pichincha Basin) has a dark brown midventral stripe (Fig. 5C). Males from Colombia (departamentos Narino and Cauca) have a cream venter with scattered brown marks (Castro and Granados, 1993). The Cotopaxi, Chimborazo, Imbabura, and Pichincha Ba- sins respectively correspond to the Latacunga, Riobamba, Otavalo, and Quito Basins mentioned by Duellman (1979). Stenocercus guenthert is probably a species complex because of the considerable geographic variation (especially color patterns of males). However, clarification of the taxonomic status of the populations of this species requires further collection and study. Stenocercus haenschi (Werner) Liocephalus formosus Boulenger, 1880:43. Holotype: MRHN 2007 from “Andes of Ecuador” (restricted to Balsapamba, 750 m, Provincia Bolivar, Ecuador, by Fritts [1974]). Replacement of the name Liocephalus formosus with the junior synonym Liocephalus haenschi was necessary because L. formosus became a junior secondary hom- onym of Scelotrema formosum when both taxa were combined into Stenocercus. Synonymy fide Fritts, 1974:55. Liocephalus haenschi Werner, 1901:595. Holotype: ZMB 16595 from “Balsapamba [Provincia Bolivar], 750 m, Ecuador’. Ophryoessoides haenschi—Etheridge, 1966:88. Stenocercus haenschi—Fritts, 1974:55. Diagnosis.—(1) Maximum SVL in males 76 mm; (2) unknown; (3) vertebral scales 50; (4) scales around mid- body 57-66; (5) internasal scales 3; (6) gular scales 56; (7) lamellae on Finger IV 26-28; (8) lamellae on Toe IV 30; (9) posthumeral pocket Type 2; (10) postfemoral pocket Type 2; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, DD SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS wrinkled, juxtaposed; (14) ventral scales smooth, imbri- cate; (15) scales on posterior surfaces of thighs granular; (16) antehumeral fold present; (17) tail not laterally com- pressed; (18) gular region in males not black; (19) dorsum bluish green. Stenocercus haenschi differs from other species of Stenocercus by the combination of smooth ventrals, granu- lar scales on posterior surface of thigh, no enlarged supraoculars, no vertebral crest, 3 caudal whorls per autotomic segment, caudals not mucronate, posthumeral pocket Type 2, postfemoral pocket Type 2, and 26-28 lamel- lae on Finger IV. Description of holotype.—Head (Fig. 10B and D) wider than high (HH/HW = 0.86); occipitals, parietals, interparietal, and postparietals small, wrinkled, and jux- taposed; 3 internasals; 2 canthals, the most anterior in con- tact with the nasal; supraoculars smooth; 1 row of supraocular scales more than twice size of scales in adja- cent rows; lateral temporals small, keeled, juxtaposed; parietal eye not visible; gulars smooth and imbricate; men- tal in contact with first pair of infralabials and first pair of postmentals. Body wider than high (BH/BW = 0.80); dorsal scales of neck and body keeled and imbricate; lateral scales of neck granular; lateral scales of body keeled, imbricate, decreasing to less than one half size of dorsals laterally; vertebrals inconspicuous, not forming crest; ventrals smooth, imbricate, same size as largest dorsals; preauricu- lar fringe well developed; antegular, gular, longitudinal neck, oblique neck, postauricular, rictal, and ventrolateral folds present; palmars, plantars, and dorsal scales of fore- limbs and hind limbs keeled, imbricate; dorsal scales of hands and ventral scales of hind limbs smooth, imbricate; scales on posterior surface of thighs granular; 28 lamellae on Finger IV; 30 lamellae on Toe IV; tail not compressed; all caudals keeled and imbricate except scales of anterior ventral surface smooth and imbricate; posthumeral and postfemoral pockets Type 2. Tail of holotype broken; there- fore, TL/ToL not estimated. In life, the holotype had the following color features: dorsum bluish green; flanks grayish green with small pale green spots; large black mark on each shoulder; irregular dark transverse stripes on limbs; dorsum of head pale ol- ive-brown; gular region grayish green with pale spots; venter bluish green; dorsal surface of tail brown with dark transverse bands; ventral surface of tail gray with narrow pale transverse bands (Werner, 1901). The preserved holotype has the following color fea- tures: dorsum and flanks bluish gray; flanks, lateral neck surfaces and gular region speckled with several small white spots; large rhomboidal mark in front of each shoulder; Fig. 12. Stenocercus haenschi, holotype, ZMB 16595, male. ventral surface gray; dorsal surface of tail greenish gray; ventral surface of tail cream with irregular greenish-gray marks. Data on holotype: Male (Fig. 12); SVL = 76 mm; HW = 14.7 mm; HL = 19.7 mm; HH = 12.6 mm; BW = 19.8 mm; BH = 15.9 mm; SM = 66; VS = 50; PS = 64; SS =5; IS =3; GS = 56; SF = 28; ST = 30. Distribution and ecology.—Stenocercus haenschi is known only from the type locality (Balsapamba, Provincia Bolivar) at 750 m on the western slopes of the Cordillera Occidental, (Fig. 11). This locality is in the upper valley of the Rio Babahoyo (Pacific Drainage), within the Humid Premontane Forest, where the mean annual temperature is 18-24 ’C and the mean annual precipitation is 1000-2000 mm. Remarks.—Except for the type material (MRHN 2007 and ZMB 16595), no specimens of Stenocercus haenschi have been collected. I visited the type locality (Balsapamba, Provincia Bolivar) in June 1997 but found no specimens. Natural vegetation of Balsapamba has been replaced mostly by orange, banana, and coffee plantations. Possi- bly, the population of this species has been severely af- fected by human activities. More exhaustive collections at the type locality are necessary to address the population status of Stenocercus haenschi. Stenocercus humeralis (Gunther) Microphractus humeralis Gunther, 1859a:90. Syntypes: BM 1946.8.11.76- 77, from “Andes of Ecuador” (restricted to Loja, 2150 m, Provincia Loja, Ecuador, by Fritts [1974)). Stenocercus humeralis—Peters, 1967:34. Diagnosis.—(1) Maximum total length in males 301 mm (7 = 17); (2) maximum total length in females 272 mm (n = 16); (3) vertebral scales 81-102; (4) scales around mid- body 98-119; (5) internasal scales 2-4; (6) gular scales 43— 68; (7) lamellae on Finger [V 24-33; (8) lamellae on Toe IV 28-40; (9) posthumeral pocket Type 1 or 2; (10) postfemoral pocket Type 3; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 23 Fig. 13. Stenocercus humeralis, syntype, BM 1946.8.11.76, female. scales small, smooth, juxtaposed; (14) ventral scales smooth, imbricate; (15) scales on posterior surfaces of thighs granular; (16) antehumeral fold present; (17) tail not laterally compressed; (18) gular region of males not black; (19) dorsum bluish green or brownish green. Stenocercus humeralis is distinguished from other spe- cies of Stenocercus by the combination of smooth ventrals, granular scales on posterior surface of thigh, 3 caudal whorls per autotomic segment, caudals not mucronate, posthumeral pocket Type 1 or 2, postfemoral pocket Type 3, 98-119 scales around midbody, 24-33 lamellae on Fin- ger IV, and 81-102 vertebral scales. Description of syntype BM 1946.8.11.76.—Head wider than high (HH/HW = 0.76); occipitals, parietals, interparietal, and postparietals small, smooth, and juxta- posed; 4 postrostrals, the 2 innermost longer than wide; 4 internasals; 2 canthals on each side, most anterior sepa- rated from nasal by tiny scales; supraoculars smooth; en- larged supraoculars absent; lateral temporals smooth and small, approximately half size of dorsal head scales; pari- etal eye not visible; gulars smooth, imbricate; mental in contact with first pair of infralabials and first pair of postmentals. Dorsal and lateral scales of neck granular; dorsal scales of body imbricate, slightly keeled, becoming gradually granular toward flanks; vertebral crest nearly inconspicu- ous; ventrals smooth, imbricate; preauricular fringe poorly developed; antehumeral, gular, antegular, longitudinal neck, oblique neck, rictal, postauricular, supra-auricular, dorsolateral, and ventrolateral folds present; dorsal scales of forelimbs and hind limbs keeled, imbricate; ventral scales of forelimbs granular proximally, smooth and im- bricate distally; ventral scales of hind limbs smooth, im- bricate; palmar and plantar scales slightly keeled, imbri- cate; scales on posterior surface of thighs granular; 24 Table 7. Sexual variation in scutellation and measurements of Stenocercus humeralis. Range (first line), x + SD (second line) and n (third line) are given. Character Females Males Scales around midbody 98-119 100-119 106.69 + 6.22 109.65 + 5.74 16 17 Vertebral scales 81-101 83-102 91.13 + 6.37 90.76 + 5.82 15 17 Paravertebral scales 106-145 114-140 126.53 + 9.21 128.29 + 6.44 15 17 Gular scales 43-68 45-56 49.25 + 5.63 49.82 + 3.15 16 17 Supraocular scales 6-7 6-8 6.88 + 0.34 7.06 + 0.56 16 17 Internasal scales 3-4 24 3.94 + 0.25 3.82 + 0.53 16 17 Subdigital lamellae on Finger IV 24-33 26-31 29.06 + 2.35 29:29 =F 1-57, 16 17 Subdigital lamellae on Toe IV 28-40 34-40 36.56 + 3.14 37.19 + 1.72 16 16 Head length/head width 1.25-1.43 1.15—1.35 1.31 +0.05 1.26 + 0.06 16 17 Head height/head width 0.67-0.93 0.70-0.88 0.78 + 0.07 0.78 + 0.05 16 iW Body height/body width 0.56-0.90 0.61-0.89 0.70 + 0.1 0.71 0.07 13 15 Regenerated tail length/tail length 1.74 1.58-3.84 = 2.67 0.8 1 5 Tail length/total length 0.63-0.67 0.62-0.68 0.65 + 0.01 0.66 + 0.02 13 11 Maximum size (mm) 272 301 lamellae on Finger IV; 28 lamellae on Toe IV; tail not com- pressed; caudals keeled, imbricate; tail length 57% of total length; posthumeral pocket Type 1; postfemoral pocket Type 3. Data on syntype BM 1946.8.11.76: Female (Fig. 13); SVL = 76; TL = 101 mm; HW = 13.3 mm; HL = 17.1 mm; HH = 10.1 mm; SM = 119; VS = 82; PS = 131; SS =7 (right side), 8 (left side); IS = 4; GS = 68; SF = 24; ST = 28. Variation.—Measurements and scutellation of Stenocercus humeralis in Table 1. Head (Fig. 14A and C) wider than high (HH/HW = 0.67-0.93, x = 0.78 £0.06, n = 34); 2-4 internasals, usually 4 (91% of specimens); canthals separated from nasals by 1 or 2 tiny scales (73%); mental in contact with small scale between the first pair of 24 ScrEnTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 14. Dorsal and lateral views of the heads of two species of Stenocercus. Aand C S. humeralis, EPN 5828, female. B and D S. iridescens, QCAZ 3322, female. Scale bars = 5 mm. postmentals in 37 % of specimens; body wider than high (BH/BW = 0.56-0.90, x = 0.70 + 0.08, 1 = 28); 24-33 lamel- lae on Finger IV; 28-40 lamellae on Toe IV; tail length 57— 68% of total length; posthumeral pocket Type 1 or 2. Sexual variation in scutellation and measurements of S. humeralis is presented in Table 7. Color in life: Dorsum in males (Fig. 5D) yellow-green with a few transverse rows of yellow spots extending onto flanks; black dorsal and lateral collar extending to inser- tion of forearm on each side; venter pale yellow (Thomas H. Fritts, field notes, 28 June 1970). Dorsum and flanks in females yellowish green; scattered black flecks on dorsum; chin and gular region pale yellowish green; ventral sur- face of body and limbs pale yellow; subcaudal region beige (Thomas H. Fritts, field notes, 26 and 27 June 1970). Dor- sum in juveniles yellowish beige or pale brown; transverse black bars on vertebral line; black flecks and blotches might be present on dorsum; chin and venter yellow (Thomas H. Fritts, field notes, 28 June 1970). Color in preservative: Dorsum with several dark short transverse bars on vertebral line; black transverse stripe across dorsum between shoulders; ground color of head, body and tail gray; ventral surfaces cream, reddish cream, or pale gray. Natural history.—One female (EPN 5829) collected in December 1968 contained four oviductal eggs, two in each oviduct. Mean size of these eggs is 17.83 mm x 10.74 mm. Mean volume of the eggs is 1076.9 mm’®. The smallest indi- vidual (FHGO 1495), collected on 14 October 1996, has a total length of 118 mm (SVL = 40 mm, TL = 78 mm). Distribution and ecology.—Stenocercus humeralis oc- curs at elevations of 2000-3000 m in the Loja Basin in south- ern Ecuador. It inhabits the upper valleys of the Rio Catamayo (Pacific Drainage) and Rio Zamora (Atlantic Drainage) in Provincia Loja (Fig. 11). The distribution lies within the Dry Low Montane Forest and Humid Premontane Forest. The mean annual temperature in these zones is 12-18 °C and 18-24°C, respectively. The mean an- nual precipitation is 500-1000 mm in the former and 1000- 2000 mm in the latter. Stenocercus humeralis and S. ornatus occur in sympatry in the upper valley of the Rio Zamora; S. humeralis is abun- dant on small shrubs and trunks of eucalyptus trees in mesic valleys, as well as on agave plants and top of fences, whereas S. ornatus is confined to more open habitats at bases of shrubs and in low rows of agave (Fritts, 1974). Stenocercus iridescens (Gunther) Liocephalus iridescens Gunther, 1859b:409. Syntypes: BM 60.6.16.2-7, from “Andes of western Ecuador.” Leiocephalus iridescens iridescens (Part)—Burt and Burt, 1931:269. Ophryoessoides iridescens—Etheridge, 1966:88. Stenocercus iridescens—Frost, 1992:43, Diagnosis.—(1) Maximum total length in males 294 mm (71 = 17); (2) maximum total length in females 213 mm (n = 16); (3) vertebral scales 40-52; (4) scales around mid- body 35-52; (5) internasal scales 2-4; (6) gular scales 16- 20; (7) lamellae on Finger IV 15-18; (8) lamellae on Toe IV 22-28; (9) posthumeral pocket Type 1 or 2; (10) postfemoral pocket Type 1; (11) projecting angulate temporals absent; (12) one row of enlarged supraoculars, more than twice size of scales in adjacent rows; (13) occipital scales large, smooth, slightly imbricate; (14) ventral scales keeled, im- bricate; (15) scales on posterior surfaces of thighs keeled, imbricate; (16) antehumeral fold absent; (17) tail strongly compressed laterally; (18) gular region of males black; (19) dorsum brownish green or brownish blue in males and dark brown in females. Stenocercus iridescens differs from other species of Stenocercus by the combination of the following charac- ters: keeled ventrals, large and smooth occipital scales, one row of enlarged supraoculars more than twice the size of scales in adjacent rows, two canthal scales, posthumeral pocket Type 1 or 2, postfemoral pocket Type 1, absence of dorsolateral crest, and absence of projecting angulate temporals. Description of syntype BM 60.6.16.2—Head wider than high (HH/HW = 0.90); occipitals, parietals, interpa- rietal, and postparietals large, smooth, slightly imbricate; 2 postrostrals, wider than long; 2 internasals; 2 canthals on each side, most anterior one separated from nasal by tiny scales; supraoculars smooth, slightly imbricate, 1 row being more than twice size of scales in adjacent rows; lat- eral temporals imbricate, slightly keeled; parietal eye vis- ible; gulars imbricate, slightly keeled, each bearing apical ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 25 Fig. 15. Stenocercus iridescens, syntype, BM 60.6.16.2, male. pit; mental in contact with first pair of infralabials and first pair of postmentals. Dorsal and lateral scales of body and neck keeled, imbricate; vertebrals large, forming prominent serrate ver- tebral crest; ventrals keeled, imbricate; preauricular fringe absent; neck folds absent; limb scales keeled, imbricate; ventral scales of hind limbs smooth or slightly keeled; 15 lamellae on Finger IV; 27 lamellae on Toe IV; tail strongly compressed; all caudals keeled, imbricate; vertebral crest extending more than half length of tail; tail length 65% of total length; posthumeral pocket Type 2; postfemoral pocket Type 1. Data on syntype BM 60.6.16.2: Male (Fig. 15); SVL = 82 mm; TL = 155 mm; HW = 14 mm; HL = 17.8 mm; HH = 12.6 mm; SM = 49; VS = 51; PS = 57; SS = 4; IS = 2; GS = 20; SF = 1S: Sila 27, Variation.—Measurements and scutellation of Stenocercus iridescens in Table 1. Head (Fig. 14B and D) nor- mally wider than high (HH/HW = 0.72-1.04, x = 0.88 + 0.07, n = 42); 1 or 2 postrostrals, wider than long; 2-4 internasals, usually 2 (60% of the specimens); parietal eye visible in 79% of the specimens; gulars bearing an apical pit in 91% of the specimens; postmentals in contact medi- ally or separated by a tiny scale (11% of the specimens); body generally wider than high (BH/BW = 0.66-1.19, x= 0.86 + 0.14, n = 36); vertebral crest more developed in adult males; 15-18 lamellae on Finger IV; 22-28 lamellae on Toe IV; tail length 63-70% of total length; posthumeral pocket Type 1 or 2. Sexual and geographic variation in measure- ments and scutellation of S. iridescens in Tables 8 and 9 respectively. Table 8. Sexual variation in scutellation and measurements of Stenocer- cus iridescens. Range (first line), x + SD (second line) and n (third line) are given. Character Females Males Scales around midbody 38-52 35-49 42.13 + 3.36 41.71 +3.48 16 17 Vertebral scales 40-52 41-51 44.50 + 3.22 45.88 + 2.96 16 7 Paravertebral scales 44-52 44-57 48.56 + 2.19 50.18 + 3.86 16 17 Gular scales 16-20 17-20 17.94 + 1.24 18.41 + 0.87 16 Th Supraocular scales 2-4 3-5 3.31 £0.70 3.41 + 0.62 16 7 Internasal scales 2-4 2-3 2.69 + 0.70 2.24 + 0.44 16 17 Subdigital lamellae on Finger IV 15-17 15-17 15.63 + 0.62 15.88 + 0.89 16 16 Subdigital lamellae on Toe IV 23-27 22-27 24.69 + 1.54 24.50 + 1.55 16 16 Head length/head width 1.19-1.3 1.09-1.30 1.23 + 0.04 1.23 + 0.07 16 17 Head height/head width 0.79-0.97 0.72-1.04 0.88 + 0.05 0.89 + 0.08 16 7 Body height/body width 0.66-1.12 0.70-1.19 0.84 + 0.12 0.90 0.13 16 14 Regenerated tail length/tail length 3.38 — 1 = Tail length/total length 0.21-0.70 0.65—-0.70 0.63 + 0.14 0.68 + 0.01 11 15 Maximum size (mm) 213 294 Color in life: Dorsum (Fig. 5E) of adult male (KU 142684) brown with dark brown chevrons, especially on the neck where they are black; side of neck pale greenish gray and black; pale greenish-yellow spots on thighs; chin yellow anteriorly, pink or gray posteriorly; gular region black; throat and ventral surfaces of forelimbs bright yellow; belly and ventral surface of proximal half of tail lavender-pink; ventral surfaces of hind limbs bronze-gray; iris copper; tongue white (William E. Duellman, field notes, 4 July 1971). Dorsum of adult female (QCAZ 3322) brown; white line extending longitudinally from tympanum to midbody; another white line extending vertically from the former to shoulder, this line bordered anteriorly with black; sides of head (including supralabials and infralabials) white with 26 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 9. Geographic variation in scutellation and measurements of Stenocercus iridescens. Range (first line), X= SD (second line), and n (third line) are given. Balzar, Cerro Blanco, Tonsupa, Cabo Pasado, Rio Palenque, Character Guayas Guayas Esmeraldas Manabi Los Rios Scales around midbody 35-41 40-43 43-45 39-44 39-45 38.86 + 1.86 41.25 + 1.50 44.00 + 0.82 42.00 + 2.16 42.50 + 2.65 7 4 4 4 4 Vertebral scales 40-51 42-50 43-47 43-49 40-45 45.14 + 3.53 46.50 + 3.32 45.25 + 1.71 45.75 + 2.75 42.25 + 2.22 7 4 4 4 4 Paravertebral scales 47-55 48-56 47-53 46-51 44-48 50.14 + 2.79 52.25 + 3.86 49.75 + 2.50 48.75 + 2.06 45.00 + 2.00 7 4 4 4 4 Gular scales 18 18-20 19-20 16-18 17-18 _ 19.00 + 0.82 19.25 + 0.50 17.00 + 0.82 17.75 + 0.50 7 4 4 4 4 Supraocular scales 3-4 3-4 3 2-4 3-4 3.71 + 0.49 3.75 + 0.50 = 3.00 + 0.82 3.25 + 0.50 7 4 4 4 4 Internasal scales 23 2-3 28} 2-4 2 2.29 + 0.49 2.25 + 0.50 2.50 + 0.58 3.00 + 0.82 = 7 4 4 4 4 Subdigital lamellae on Finger IV 15-16 ISS17, 15-16 15-16 I=Al7/ 15.43 + 0.53 16.25 + 0.96 15.25 + 0.50 15.75 + 0.50 16.00 + 1.15 7 4 4 4 4 Subdigital lamellae on Toe IV 22-26 24-27 24-26 23-27 22-25 23.86 + 1.35 25.50 + 1.29 25.25 + 0.96 25.00 + 1.63 23.5 + 1.29 7 4 4 4 4 Head length/head width 1.09-1.28 1.22-1.26 1.24-1.31 1.20-1.30 1.13-1.29 1.20 + 0.06 1.24 + 0.02 1.28 + 0.04 1.23 + 0.05 1.22 + 0.07 7 4 4 4 4 Head height/head width 0.79-0.93 0.84—0.96 0.88-1.04 0.87-0.97 0.82-0.98 0.86 + 0.05 0.88 + 0.06 0.96 + 0.08 0.91 + 0.05 0.89 + 0.08 7 4 4 4 4 Body height/body width 0.75—-0.96 0.69-0.87 0.72-1.12 0.66-0.77 0.80-0.95 0.85 + 0.08 0.79 + 0.08 0.98 + 0.22 0.73 + 0.05 0.90 + 0.07 6 4 3 4 4 Tail length/total length 0.67—-0.70 0.67—-0.69 0.67—-0.68 0.21-0.70 0.68—-0.69 0.68 + 0.01 0.68 + 0.01 0.67 + 0.01 0.56 + 0.23 0.68 + 0.01 7 3 2 4 3 Maximum size (mm) 246 246 292 203 294 small brown dots; dark brown oblique stripe extending posteroventrally from lower border of eye to infralabials; dark brown interorbital bar present; one short horizontal white stripe on the posterior surface of each thigh; gular region and ventral surfaces of limbs and body whitish cream; belly iridescent rose. Color in preservative: Dorsum of males pale green or greenish-brown with a series of V-shaped dark brown marks on vertebral line with apex pointing posteriorly; V- shaped marks extending on proximal portion of tail in some individuals; flanks bluish gray; dorsum of head dark green with several small black dots; sides of head bluish gray, yellowish brown, or pale brown; pale brown stripe extending downward from eye to angle of mouth; gular region pale blue or pale gold; black band on throat; pink ventral surface of body, sometimes pale blue laterally; ven- tral surface of limbs yellow; dorsal surface of tail brown or yellowish brown; ventral surface of tail yellow proximally and brown distally. Dorsum and flanks of females dark brown or greenish brown; V-shaped marks over vertebral line; white line extending from shoulder to level of eye in some specimens; sides of head white or pale brown with small dark brown dots; pale brown stripe extending down- ward from eye to angle of mouth; gular region white or cream with several short longitudinal dark brown lines; ventral surface of body cream, greenish cream or dark yel- lowish brown; ventral surface of limbs pale yellow; tail yellowish brown. Distribution and ecology.—In western Ecuador, Stenocercus iridescens occurs at elevations of 0-2000 m in the provinces of Azuay, Cotopaxi, El Oro, Esmeraldas, Guayas, Los Rios, and Manabi (Fig. 16). The distribution lies mainly within the Tropical Desertic Thicket, Very Dry Tropical Forest, Dry Tropical Forest, Premontane Humid ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 27 81° 80° 29 20 40 60 80 100 ——— 1 Kilometers ( <300m 9 300-2000 m GB 2000-4000 m Gy > 4000m © Permanent snow ®@ S. iridescens | YS. limitaris BS. ornatus A S. rhodomelas Fig. 16. Distribution of four species of Stenocercus in Ecuador. Forest, and Very Humid Premontane Forest life zones. The mean annual temperature is 24—26°C in the former two life zones, 24—25°C in the third and 18-24°C in the latter two. The mean annual precipitation is 125-250 mm in the first one, 500-1000 mm in the second one, 1000-2000 mm in the third and fourth ones, and 2000-4000 mm in the lat- ter one. This species also occurs in northwestern Peru (Cadle, 1991). Stenocercus limitaris Cadle Stenocercus limitaris Cadle, 1998:261. Holotype: AMNH 22183 a male from Alamor, 04°02' S, 80°02' W, 1325 m, Provincia Loja, Ecuador. Diagnosis.—(1) Maximum SVL in males 97 mm (Cadle, 1998); (2) maximum SVL in females 82 mm (Cadle, 1998); (3) vertebral scales 40-52; (4) scales around mid- body 39-54; (5) internasal scales 4-5; (6) gular scales 17- 23; (7) lamellae on Finger IV 17-23; (8) lamellae on Toe IV 24-32; (9) posthumeral pocket Type 3 or 4; (10) postfemoral pocket Type 4 or 5; (11) projecting angulate temporals ab- sent; (12) one row of enlarged supraoculars more than twice size of scales in adjacent rows; (13) occipital scales large, keeled or wrinkled, juxtaposed; (14) ventral scales mucr- onate, keeled, imbricate; (15) scales on posterior surfaces of thighs mucronate, keeled, imbricate; (16) antehumeral fold nearly inconspicuous; (17) tail strongly compressed laterally; (18) gular region in males not black; (19) dorsum brown in males and females. Stenocercus limitaris is distinguished from other spe- cies of Stenocercus by the combination of keeled ventrals, keeled, wrinkled, or multicarinate large occipital scales, one row of enlarged supraoculars more than twice the size of scales in adjacent rows, two canthal scales, posthumeral pocket Type 3 or 4, postfemoral pocket Type 4 or 5, usu- ally more than 40 scales around midbody, absence of dor- solateral crest, and absence of projecting angulate temporals. Description and variation.—Cadle’s (1998) descrip- tion is sufficient. Measurements and scutellation of S. limitaris (from Cadle,1998) are summarized in Table 1. Distribution and ecology. —Stenocercus limitaris occurs at elevations of 1000-1300 m in southwestern Ecuador. It inhabits the upper valley of the Rio Chira (Pacific Drain- age) in Provincia El Oro and Provincia Loja (Fig. 16). The distribution lies within the Humid Premontane Forest, where the mean annual precipitation is 1000-2000 mm and the mean annual temperature is 18-24°C. Stenocercus limitaris is sympatric with S. carrioni and S. iridescens. This species occurs also in northwestern Peru (Cadle, 1998). Stenocercus ornatus (Gray) Leiocephalus ornatus Gray, 1845:219. Holotype: BM 1946.8.29.72 from “Guayaquil [Provincia Guayas, Ecuador]” (restricted to Loja, 2150 m, Provincia Loja, Ecuador, by Fritts [1974]). Leiocephalus ornatus ornatus (Part)—Burt and Burt, 1931:271. Ophryoessoides ornatus—Etheridge, 1966:88. Stenocercus ornatus—Fritts, 1974:62. Diagnosis ——(1) Maximum total length in males 294 mm (7 = 15); (2) maximum total length in females 207 mm (n = 11); (3) vertebral scales 36-50; (4) scales around mid- body 46-58; (5) internasal scales 2-4; (6) gular scales 15- 23; (7) lamellae on Finger IV 17-25; (8) lamellae on Toe IV 26-37; (9) posthumeral pocket Type 4; (10) postfemoral pocket Type 5; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, keeled or wrinkled, juxtaposed; (14) ventral scales smooth or slightly keeled, imbricate; (15) scales on posterior surfaces of thighs keeled, imbricate; (16) antehumeral fold nearly inconspicuous; (17) tail slightly or moderately compressed laterally; (18) gular region of males not black; (19) dorsum brown. Stenocercus ornatus is distinguished from other species of Stenocercus by the combination of smooth ventrals (slightly keeled in 43% of the specimens), imbricate scales on posterior surface of thigh, no enlarged supraoculars, slightly imbricate occipitals, posthumeral pocket Type 4, 28 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS EDe i Sa G nals essere Ne eee Be Sci Gu) DX zs 300) See A B — S ie WE OSD Fig. 17. Dorsal and lateral views of the heads of two species of Stenocercus. A and C S. ornatus, FHGO 679, male. B and D S. rhodomelas, QCAZ 3663, male. Scale bars = 5 mm. postfemoral pocket Type 5, 46-58 scales around midbody, 17-25 lamellae on Finger IV, 26-37 lamellae on Toe IV, no black gular patch in males, vertical black bar on antehumeral fold in males, and antehumeral, supra-au- ricular, and dorsolateral folds. Description and variation.—Measurements and scutellation of Stenocercus ornatus in Table 1. Head (Fig. 17A and C) wider than high (HH/HW = 0.71-0.98, x = 0.85 + 0.05, n = 40); occipitals, parietals, interparietal, and postparietals small, keeled, slightly imbricate; 2-4 postrostrals (wider than long when there are 2 or 3 but as wide as long when there are 4); 2-4 internasals, usually 4 (64% of specimens); 2 canthals, most anterior one in con- tact with nasal or separated from it by 2 tiny scales; supraoculars keeled; enlarged supraoculars absent; lateral temporals slightly keeled, imbricate; parietal eye visible in 82% of specimens; gulars imbricate and smooth, each bearing apical pit; mental in contact with first pair of infralabials and first pair of postmentals; postmentals in contact medially or sometimes separated by tiny scale (14% of specimens). Body normally wider than high (BH/BW = 0.62-1.36, X= 0.82 + 0.16, n = 34); dorsal scales of neck and body and lateral scales of body keeled, imbricate, mucronate; lateral scales of neck keeled, imbricate; vertebrals large, forming prominent, serrate vertebral crest; slight dorsolateral crest on each side of neck (extending behind shoulder in some specimens); ventrals imbricate, smooth, or slightly keeled (43% of the specimens); preauricular fringe nearly incon- spicuous; antehumeral, supra-auricular, and dorsolateral folds weakly developed, most evident in adult males; ven- tral scales of forelimbs, dorsal scales of forelimbs and hind Table 10. Sexual variation in scutellation and measurements of Stenocercus ornatus. Range (first line), x+SD (second line) and n (third line) are given. Character Females Males Scales around midbody 47-56 46-58 51.95 + 2.26 51.95 + 3.19 20 20 Vertebral scales 39-50 36-50 45.35 + 3.01 42.63 + 3.47 20 19 Paravertebral scales 53-65 54-66 60.30 + 3.37 59.68 + 3.8 20 19 Gular scales 15-20 16-23 18.1 + 1.33 18.8 + 1.54 20 20 Supraocular scales 4-6 S=i/ 5.25 + 0.55 5.6 + 0.60 20 20 Internasal scales 24 2-4 3.55 + 0.76 3.77 + 0.57 20 20 Subdigital lamellae on Finger IV 17-24 17-25 20.65 + 1.95 21.8 + 1.88 20 20 Subdigital lamellae on Toe IV 26-34 27-37 29.2 + 1.85 30.85 + 2.41 20 20 Head length/head width 1.16-1.37 1.06-1.34 1.29 + 0.06 1.26 + 0.06 20 20 Head height/head width 0.71-0.93 0.78-0.98 0.83 + 0.05 0.87 + 0.05 20 20 Body height/body width 0.62-1.05 0.69-1.36 0.74 + 0.10 0.89 + 0.17 16 18 Regenerated tail length/tail length 0.01-2.76 0.95-2.13 1.44 + 1.20 1.36 + 0.55 6 4 Tail length/total length 0.66-0.69 0.6-0.72 0.68 + 0.01 0.69 + 0.03 1 15 Maximum size (mm) 207 294 limbs, palmars and plantars imbricate, keeled (mucronate in some specimens); ventral scales of hind limbs smooth, imbricate; scales on posterior surface of thighs keeled, imbricate; dorsal scales of hands and feet imbricate, keeled, or smooth (some with apical pit); 17-25 lamellae on Fin- ger IV; 26-37 lamellae on Toe IV; tail slightly compressed in females, moderately compressed in males; caudals keeled, imbricate, mucronate (anterior ventrals not mucr- onate); vertebral crest extending more than half length of tail; tail length 60-72% of total length; posthumeral pocket Type 4; postfemoral pocket Type 5. Sexual variation in scutellation and measurements of Stenocercus ornatus in Table 10. Color in life: Dorsum of males (Fig. 5F) pale brown or gray brown; narrow or wide black transverse blotches on ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 29 dorsum between indistinct beige dorsolateral stripes; flanks suffured with pink or reddish cream; conspicuous broad black bar on antehumeral fold; chin and gular re- gion pale red or pinkish red (with black suffusion in some specimens); yellow patch on pectoral region; broad black stripe on each side of midventral line between pectoral and pelvic regions; rest of ventral surface of body pink or reddish cream; anal region, ventral surface of thighs, and base of tail yellow (heavily suffused with black in some specimens); lateral and ventral surface of tail pink (Tho- mas H. Fritts, field notes, 21 June 1970). Dorsum of an adult female (QCAZ 3790) brown with black marks; pale brown irregular bands in flanks; white scale below each eye; chin and labials yellow; venter pink with gray dotted pattern medially; distal half of tail greenish brown; iris pale brown (John J. Wiens, field notes, 27 April 1990). Color in preservative: Dorsum of males dark brown, ol- ive-green, or bluish gray; 6-8 dark brown, short transverse bars on vertebral line from neck to base of tail; 4—5 black transverse lines on dorsal surface of limbs; large black mark anterior to each shoulder; gular region bluish gray, dark brown, pale blue, or pale red; pectoral region black (with several pale blue, cream, or gray scales in some specimens); black midventral stripe; rest of belly pink, gray, or dark brown; ventral surface of limbs pale blue, brown, or gray; dark transverse rings intercalated with pale rings on tail; black mark on the anterior ventral region of the tail in 18% of specimens. Dorsal color of females same as in males, but transverse dark bars narrower; pale dorsolateral line extending from behind eye to hind limbs in 60% of speci- mens; black mark in front of shoulder absent; loreal re- gion cream; dark brown or black stripe from angle of mouth, through tympanum, to, or behind shoulder; supralabials and infralabials dark brown or black; ventral surfaces bluish gray, cream, reddish cream, or yellowish green, with small irregular dark marks in some specimens; scattered dark brown flecks in gular region; tail with ringed pattern as in males. Natural history —One female (EPN 5879) contained two oviductal eggs. The sizes of these eggs are 17.01 mm x 9.47 mm and 18.07 mm x 9.32 mm; their volumes are 798.7 mm~? and 821.8 mm’, respectively. Distribution and ecology.—Stenocercus ornatus occurs at elevations of 1500-3000 m in the Loja Basin in southern Ecuador. It inhabits the upper valleys of the Rio Catamayo (Pacific Drainage) and Rio Zamora (Atlantic Drainage) in Provincia Loja (Fig. 16). The distribution lies within the Dry Low Montane Forest and Humid Premontane Forest. The mean annual temperature in these zones is 12-18°C and 18-24°C, respectively. The mean annual precipitation is 500-1000 mm in the former and 1000-2000 mm in the latter. Stenocercus ornatus and S. humeralis occur in sympatry in the upper valley of the Rio Zamora. According to Fritts (1974), S. humeralis is abundant on small shrubs and trunks of eucalyptus trees in mesic valleys, as well as on agave plants and on top of fences, whereas S. ornatus is confined to more open habitats at bases of shrubs and in low rows of agave. Stenocercus rhodomelas (Boulenger) Liocephalus rhodomelas Boulenger, 1899:455. Syntypes: BM 1946.8.29.77- 80, from “Ona [Provincia Azuay], Ecuador.” Ophryoessoides rhodomelas—Etheridge, 1966:88. Stenocercus rhodomelas—Fritts, 1974:63. Diagnosis.—(1) Maximum total length in males 235 mm (7 = 10); (2) maximum total length in females 236 mm (n = 13); (3) vertebral scales 43-54; (4) scales around mid- body 43-55; (5) internasal scales 2-4; (6) gular scales 17- 20; (7) lamellae on Finger IV 14-21; (8) lamellae on Toe IV 22-29; (9) posthumeral pocket Type 4; (10) postfemoral pocket Type 5; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, keeled or wrinkled, juxtaposed; (14) ventral scales smooth, imbricate; (15) scales on posterior surfaces of thighs keeled, imbricate, mucronate; (16) antehumeral fold absent; (17) tail not laterally compressed; (18) gular region of males black; (19) dorsal ground color of males brown. Stenocercus rhodomelas differs from other species of Stenocercus by the combination of smooth ventrals, imbri- cate scales on posterior surface of thigh, no enlarged supraoculars, posthumeral pocket Type 4, postfemoral pocket Type 5, 43-55 scales around midbody, 17-20 gular scales, black gular patch in males, black ventral surface of hindlimbs in males, and no neck folds. Description of syntype BM 1946.8.29.77.—Head wider than high (HH/HW = 0.86); occipitals, parietals, interparietal and postparietals small, keeled or wrinkled, juxtaposed; 2 postrostrals, wider than long; 2 internasals; 2 canthals, most anterior one in contact with nasal; supraoculars keeled or wrinkled, juxtaposed; enlarged supraoculars absent; lateral temporals keeled, imbricate; parietal eye visible; gulars large, smooth, imbricate, each with an apical pit; mental in contact with first pair of infralabials and first pair of postmentals. Dorsal and lateral scales of neck and body keeled, imbricate, mucronate; vertebrals large, forming prominent vertebral crest; ventrals smooth, imbricate, each with an apical pit; preauricular fringe well developed; neck folds absent; forelimb scales keeled, imbricate, mucronate (some dorsal scales of hand smooth, imbricate); dorsal scales of hind limb keeled, imbricate, mucronate; ventral scales of hind limbs smooth or weakly keeled and imbricate; pal- mar and plantar scales keeled, imbricate; 18 lamellae on Finger IV; 29 lamellae on Toe IV; tail slightly compressed; 30 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 18. Stenocercus rhodomelas, syntype, BM 1946.8.29.77, male. caudals keeled, imbricate, mucronate (anterior ventrals smooth, imbricate); vertebral crest extending more than half the length of the tail; tail length 64% of total length; posthumeral pocket Type 4; postfemoral pocket Type 5. Data on syntype BM 1946.8.29.77: Male (Fig. 18); SVL = 82 mm; TL = 146 mm; HW = 13.2 mm; HL = 17.8 mm; HH = 11.3 mm; SM = 50; VS =51; PS = 56;5S =5; IS = 2; GS = 1192 Sis = ie Sir = 28), Variation.—Measurements and scutellation of Stenocercus rhodomelas in Table 1. Head (Fig. 17B and D) wider than high (HH/HW = 0.69-0.99, x = 0.83 + 0.07, n = 29); 2-3 postrostrals, wider than long; if 3 postrostrals, middle one as wide as long; 24 internasals, usually 2 (67% of specimens); most anterior canthals separated from na- sals by 2 tiny scales (29% of specimens); parietal eye vis- ible in 67% of specimens; apical pits of gulars and ventrals most evident in adult males; body normally wider than high (BH/BW = 0.64-1.03, x = 0.81 + 0.10, n = 16); verte- bral crest most conspicuous in adult males; 14-21 lamel- lae on Finger IV; 22-29 lamellae on Toe IV; vertebral crest extending more than half length of tail in males, less than half length in females; tail length 57-66% of total length. Sexual variation in scutellation and measurements of S. rhodomelas is presented in Table 11. Color in life: Dorsum and flanks of an adult male (QCAZ 3663) dark brown with several pink, cream, or black scales distributed irregularly; vertebral crest dark brown with a few yellow scales, some with black marks; dorsal black V-shaped mark between shoulders with vertex pos- teriorly; three faint V-shaped marks behind the latter; small black mark anterior to each shoulder; dorsal surface of head darker than body; infralabials, supralabials, and rostral black; sublabials, lorilabials, suboculars, and first pair of postmentals yellow; gular region black except for pale yel- low dot on chin; throat pink with thin black midventral stripe continuous with black gular patch; anterior part of pectoral region greenish blue; posterior part of pectoral region, midventral line, pelvic region, and ventral surfaces Table 11. Sexual variation in scutellation and measurements of Stenocer- cus rhodomelas. Range (first line), x+ SD (second line) and n (third line) are given. Character Females Males Scales around midbody 44-55 43-51 49.21 + 2.84 48.23 + 2.52 19 13 Vertebral scales 43-54 43-52 47.63 + 3.44 47.77 + 3.06 19 13 Paravertebral scales 49-60 52-61 54.79 + 3.21 55.54 + 2.63 19 13 Gular scales 17-20 17-20 18.26 + 0.93 18.38 + 0.96 19 13 Supraocular scales 3-6 5-6 5.00 + 0.67 5.54 + 0.52 9 13 Internasal scales 2-3 2-4 2.26 + 0.45 2.77 + 0.73 19 13 Subdigital lamellae on Finger IV 14-18 15-21 16.21 + 1.27 17.69 + 1.49 19 13 Subdigital lamellae on Toe IV 22-27 23-29 AR Ail se 13383 26.54 + 1.66 19 13 Head length/head width 1.19-1.36 0.98-1.50 1.27 + 0.05 1.23 + 0.13 18 12 Head height/head width 0.72-0.90 0.69-0.99 0.81 + 0.06 0.85 + 0.08 17 12 Body height/body width 0.64-0.89 0.65-1.03 0.77 + 0.08 0.83 + 0.12 13 9 Regenerated tail length/tail length 0.32-0.77 1.10-3.82 0.52 + 0.21 2.46 + 1.93 4 2 Tail length/total length 0.61—-0.66 0.57—-0.64 0.64 + 0.01 0.62 + 0.02 13 10 Maximum size (mm) 236 235 of thighs and shanks black, rest of ventral surface of body cream; small black triangular mark behind vent pointing backwards; ventral surface of tail pink proximally, cream distally (Omar Torres-Carvajal, field notes, 11 January 1997). Color in preservative: Dorsum of males dark greenish brown or pale brown with irregular black dots; 6-7 dark transverse stripes on dorsal surface of body, dark brown stripe at the level of shoulders extends to point anterior to the insertion of forelimbs; short dark stripe parallel to lat- ter at insertion of forelimbs; gular region black; throat slightly pink; anterior part of pectoral region pale blue or whitish cream; posterior part of pectoral region, midventral line, pelvic region, and ventral surfaces of thighs and shanks black; small black irregular or triangular mark ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS Bil pointing posteriorly behind vent in 60% of specimens; ven- tral surface of tail pink or cream proximally and cream distally. Dorsum of females dark brown; gular region cream with dark brown flecks; ventral surface cream. Distribution and ecology.—Stenocercus rhodomelas is known from the western slopes of the Cordillera Occiden- tal and Saraguro Basin in southern Ecuador (Provincia Azuay and Provincia Loja). It occupies the upper valley of the Rio Jubones (Pacific Drainage) at elevations of 730- 2100 m (Fig. 16). The distribution lies mainly within the Low Humid Montane Forest and Thorny Premontane Thicket life zones. The mean annual temperature is 12- 18°C and 18-24°C, respectively. The mean annual precipi- tation is 1000-2000 mm and 250-500 mm. The area occupied by this species is xeric with sparse vegetation of cactus and agave plants; individuals can be found on large rocks and on the ground at the bases of cactus (Fritts, 1974). Although Fritts (1974) mentioned that Stenocercus simonsii and S. rhodomelas are allopatric at el- evations above 2200 m in the upper valley of the Rio Jubones, both species occur in sympatry (Ona, 1981 m, Provincia Azuay). Also, S. rhodomelas possibly is sympat- ric with S. festae in the Saraguro Basin. Stenocercus simonsii Boulenger Stenocercus simonsti Boulenger, 1899:454. Syntypes: BM 1946.8.11.73-74, from “Ona, 6500 ft (= 1981.2 m), [Provincia Azuay] Ecuador.” Diagnosis.—(1) Maximum SVL in males 88 mm (Cadle, 1991); (2) maximum SVL in females 79 mm (Cadle, 1991); (3) vertebral scales 59-98; (4) scales around mid- body 79-102; (5) internasal scales 4; (6) gular scales 36-57; (7) lamellae on Finger IV 24-28; (8) lamellae on Toe IV 28— 37; (9) posthumeral pocket Type 1; (10) postfemoral pocket Type 3; (11) projecting angulate temporals absent; (12) row of enlarged supraoculars absent; (13) occipital scales small, smooth, juxtaposed; (14) ventral scales smooth, imbricate; (15) scales on posterior surfaces of thighs granular; (16) antehumeral fold well developed; (17) tail laterally com- pressed distally; (18) gular region of males not black; (19) dorsum greenish gray or pale gray. Stenocercus simonsii is distinguished from other spe- cies of Stenocercus by the combination of smooth ventrals, granular scales on posterior surface of thigh, dorsal sur- face of neck, and anterolateral surface of trunk, complete vertebral row, 2 caudal whorls per autotomic segment, caudal whorls subequal in size, posthumeral pocket Type 1, postfemoral pocket Type 3, 79-102 scales around mid- body, 36-57gular scales, 94-118 paravertebral scales, 24— 28 lamellae on Finger IV, and 6 or more scale rows within antegular fold. Description and variation.—Cadle’s (1991) descrip- tion is sufficient. Measurements and scutellation of S. simonsii (from Cadle, 1991) are summarized in Table 1. Color in life: Males with broad black collar posterior to antehumeral fold, incomplete at vertebral line; small black and yellow spots on flanks; white stripe from infraorbital scales to shoulder in some specimens; hind limbs dull black with pinkish-beige transverse stripes; limbs and tail with alternate black and white bars in some specimens; chin and gular region pale yellowish green; gular fold black interiorly; ventral surface of forelimbs, body, and base of tail orange-yellow (Thomas H. Fritts, field notes, 6 June 1970). Dorsum of females (Fig. 5G) greenish brown; chin and gular region usually with scattered brown or black spots; venter pale yellowish beige (Thomas H. Fritts, field notes, 6 June 1970). Fritts (1974) mentioned that individu- als from Giron (Provincia Azuay) differ in dorsal colora- tion from those from Saraguro (Provincia Loja). Lizards from the former locality have a pale gray-green dorsum with distinct black transverse blotches, whereas lizards from Saraguro have a pale gray dorsum with smaller, less distinct blotches. Distribution and ecology.—Stenocercus simonsti occurs at elevations of 1980-2500 m in southern Ecuador (Provincia Azuay and Provincia Loja). It inhabits the up- per valley of the Rio Jubones (Pacific Drainage) on the western slopes of the Cordillera Occidental and Saraguro Basin (Fig. 19). The distribution lies within the Low Dry Montane Forest with a mean annual temperature of 12- 18°C and a mean annual precipitation of 500-1000 mm. Stenocercus simonsii and S. festae occur in sympatry in the upper valley of the Rio Jubones; in this area S. simonsit usually is found on large rocks or rock piles, whereas S. festae utilizes the leaves of agave plants and the surround- ing ground (Fritts, 1974). Although Fritts (1974) mentioned that Stenocercus simonsii and S. rhodomelas are allopatric at elevations above 2200 m in the upper valley of the Rio Jubones, both species occur in sympatry (Ona, 1981 m, Provincia Azuay). Stenocercus varius Boulenger Stenocercus varius Boulenger, 1885:134. Holotype: BM 71.4.16.53, “un- known locality” (restricted to Tandapi, 1460 m, Provincia Pichincha, Ecuador, by Fritts [1974]). Diagnosis.—(1) Maximum total length in males 234 mm (n = 9); (2) maximum total length in females 208 mm (n = 14); (3) vertebral scales 60-74; (4) scales around mid- body 74-88; (5) internasal scales 4—5; (6) gular scales 42- 60; (7) lamellae on Finger IV 24-28; (8) lamellae on Toe IV 27-34; (9) posthumeral pocket Type 1; (10) postfemoral pocket Type 1 or 2; (11) projecting angulate temporals ab- sent; (12) row of enlarged supraoculars absent; (13) occipi- tal scales small, smooth, juxtaposed; (14) ventral scales smooth, imbricate; (15) scales on posterior surfaces of thighs granular; (16) antehumeral fold present; (17) tail 32 SCIENTIFIC PAPERS, NATURAL History MuseuM, THE UNIVERSITY OF KANSAS 81° 80° +? 20 40 60 80 100 | Kilometers {| <300m ==) 300-2000 m (8 2000-4000 m BB >4000 m © Permanent snow Fig. 19. Distribution of two species of Stenocercus in Ecuador. slightly compressed laterally; (18) gular region of males not black; (19) dorsum bluish gray or green. Stenocercus varius differs from other species of Stenocercus by the combination of smooth ventrals, granu- lar scales on posterior surface of thigh, distinct vertebral crest, 3 caudal whorls per autotomic segment, caudal scales without projecting spines, posthumeral pocket Type 1, postfemoral pocket Type 1 or 2, and 74-88 scales around midbody. Description and variation.—Measurements and scutellation of Stenocercus varius in Table 1. Head (Fig. 20A and B) usually wider than high (HH/HW = 0.73-1.04, x= 0.83 + 0.07, n = 25); occipitals, parietals, interparietal, and postparietals small, smooth, juxtaposed; 3 postrostrals, wider than long; 4-5 internasals, usually 4 (96%); 2 canthals, most anterior one in contact with nasal; supraoculars smooth, juxtaposed; enlarged supraoculars absent; lateral temporals smooth, juxtaposed; parietal eye not visible in 92% of specimens; gulars small, imbricate, smooth (most anterior gulars granular); mental in contact with first pair of infralabials and first pair of postmentals. Fig. 20. Dorsal (A) and lateral (B) views of the head of Stenocer- cus varius, QCAZ 3046, female. Scale bar = 5 mm. Table 12. Sexual variation in scutellation and measurements of Steno- cercus varius. Range (first line), x+ SD (second line) and n (third line) are given. Character Females Males Scales around midbody 74-88 79-88 81.36 + 4.16 84.56 + 3.94 14 9 Vertebral scales 60-73 64-74 66.93 + 3.83 69.33 + 2.87 14 9 Paravertebral scales 76-97 83-97 86.79 + 5.67 88.78 + 5.07 14 9 Gular scales 42-58 47-60 49.36 + 4.18 53.00 + 4.18 14 9 Supraocular scales 5-6 5-7 Soy/ ae OSI 5.56 + 0.73 14 9 Internasal scales 4-5 4 4.07 + 0.27 — 14 9 Subdigital lamellae on Finger IV 24-28 24-28 25.54 + 1.13 25.5+1.31 13 8 Subdigital lamellae on Toe IV 27-32 29-34 29.31 + 1.32 30.38 + 1.60 13 8 Head length/head width 1.22-1.46 1.21-1.33 1.32 + 0.06 1.29 + 0.04 14 9 Head height/head width 0.73-0.89 0.76-1.04 0.81 + 0.05 0.87 + 0.09 14 9 Body height/body width 0.58-0.82 0.66-0.96 0.72 + 0.08 0.77 + 0.09 13 9 Regenerated tail length/tail length 0.01—2.87 0.48 1.38 + 1.58 i 4 1 Tail length/total length 0.60-0.64 0.61-0.65 0.62 + 0.01 0.63 + 0.01 9 6 Maximum size (mm) 208 233 ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 3) Table 13. Geographic variation in scutellation and measurements of Stenocercus varius. Range (first line), x SD (second line) and n (third line) are given. Las Pampas, La Favorita, Rio Guajalito, Tandapi, Character Cotopaxi Pichincha Pichincha Pichincha Scales around midbody 81-88 80-88 74-81 79-88 86.14 + 2.54 83.25 + 3.40 77.00 + 3.24 82.00 + 3.52 7 4 5 6 Vertebral scales 64-74 63-70 60-73 66-72 68.29 + 3.20 66.75 + 3.77 66.80 + 4.66 69.50 + 2.07 7 4 5 6 Paravertebral scales 87-97 76-86 77-91 83-97 90.86 + 3.63 82.00 + 4.24 86.20 + 5.89 87.50 + 5.58 7 4 5 6 Gular scales 42-60 46-53 44-56 47-49 53.29 + 5.71 50.25 + 3.10 48.40 + 4.83 48.50 + 0.84 7 4 5 6 Supraocular scales 5-6 5-6 5-6 5-7 5.43 + 0.53 5.50 + 0.58 5.60 + 0.55 5.83 + 0.75 Y 4 5 6 Internasal scales 4 4 4 4-5 _ — — 4.17+0.41 7 4 5 6 Subdigital lamellae on Finger IV 24-28 24-27 25-26 24-28 25.33 + 1.51 25.5 + 1.29 25.75 + 0.50 25.50 + 1.38 6 4 4 6 Subdigital lamellae on Toe IV 27-30 30-32 28-34 29-31 28.83 + 1.17 30.67 +1.15 30.80 + 2.39 29.50 + 0.84 6 3 5 6 Head length/head width 1.21-1.36 1.30-1.50 1.27-1.46 1.22-1.33 1.30 + 0.05 1.38 + 0.09 1.37 + 0.08 1.28 + 0.05 Y 4 5 6 Head height/head width 0.81-1.04 0.79-0.90 0.75-0.85 0.73-0.94 0.87 + 0.08 0.83 + 0.05 0.81 + 0.04 0.83 + 0.08 7 4 5 6 Body height/body width 0.68-0.96 0.54-0.80 0.70-0.78 0.63-0.83 0.77 + 0.09 0.66 + 0.12 0.73 + 0.04 0.74 + 0.08 7 4 3 6 Regenerated tail length/tail length — 0.03-2.87 0.01 2.63 — 1.04 + 1.26 — — — 4 1 1 Tail length/total length 0.61—0.63 = 0.60-0.66 0.64 0.62 + 0.01 _— 0.64 + 0.03 _ 6 — 4 3 Maximum size (mm) 208 180 234 201 Body wider than high (BH/BW = 0.54-0.96, x= 0.73 + 0.09, n = 23); lateral scales of body and dorsal scales of neck and body keeled, imbricate; lateral scales of neck granular; lateral scales of body near venter approximately half size of dorsals; vertebrals small, not forming conspicu- ous vertebral crest; ventrals smooth, imbricate, each with an apical pit; preauricular fringe nearly inconspicuous; antehumeral, gular, antegular, longitudinal neck, oblique neck, rictal, postauricular, supra-auricular, dorsolateral, and ventrolateral folds present; dorsal scales of forelimbs and hind limbs keeled, imbricate; ventral scales of fore- limbs and hind limbs smooth, imbricate; palmars and plantars imbricate, smooth, keeled; 24—28 lamellae on Fin- ger IV; 27-34 lamellae on Toe IV; tail slightly compressed laterally; dorsal and lateral caudals keeled, imbricate; caudals keeled and imbricate ventrally; tail length 60-66% of total length; posthumeral pocket Type 1; postfemoral pocket Type 1 or 2. Sexual and geographic variation in scutellation and measurements of Stenocercus varius is pre- sented in Tables 12 and 13 respectively. Color in life: Dorsum and flanks of an adult male (QCAZ 3845) olive-green with several pale yellowish-green spots; large rhomboidal black mark anterior to each shoulder; dor- sal surface of hind limbs dark brown; dorsal surface of head with black and dark brown marks; eyelids, gular region, and throat yellow; ventral surfaces of body, limbs, and proximal portion of tail yellowish cream; ventral surface of distal por- 34 ScIENTIFIC PAPERS, NATURAL History MusEeuM, THE UNIVERSITY OF KANSAS tion of tail dark brown; iris bronze. Females without rhom- boidal mark anterior to each shoulder (Fig. 5H). Color in preservative: Dorsum gray; 6-8 short black transverse bands on dorsal surface of body from neck to base of tail; pale reticulations on dorsal surface of body and limbs; small white, cream, or bluish-cream dots on flanks in 55% of specimens; black rhomboidal mark ante- rior to each shoulder in males; black marks on dorsal sur- face of head in 95% of specimens; gular region gray, usu- ally with several small, light oval marks; ventral surfaces of body and limbs gray; usually a thin cream midventral line that is interrupted by dark brown rings on ventral surface of tail in some individuals. Natural history.—One female (QCAZ 3046) collected on 15 October 1995 contained two oviductal eggs. The sizes of these eggs are 22.63 mm x 9.14 mm and 22.44 mm x 8.96 mm, their volumes are 989.9 mm? and 943.3 mm’, respec- tively. The same female had a body temperature of 34.2°C when collected (A. Quiguango and Juan M. Guayasamin, field notes, 24 February 1998). The smallest juvenile (QCAZ 719), collected on 17 May 1988, has a total length of 134 mm (SVL = 45 mm, TL = 89 mm). Distribution and ecology.—Stenocercus varius occurs at elevations of 1460-2200 m on the western slopes of the northern part of the Cordillera Occidental. It inhabits the upper valleys of the Rio Blanco and Rio Toachi (Pacific Drainage) in Provincia Cotopaxi and Provincia Pichincha (Fig. 19). The distribution lies within the Very Humid Premontane Forest and Low Very Humid Montane For- est. The mean annual temperature is 18-24 °C and 12-18 °C, respectively. The mean annual precipitation is 2000-4000 mm in both life zones. Individuals have been found on tree trunks, fallen logs, and rocks in partially cleared areas (Fritts, 1974). Stenocercus varius also occurs in patches of primary forest at the forest edges. BIOGEOGRAPHY GEOGRAPHICAL DISTRIBUTION The distributional range of Stenocercus extends from the western lowlands (S. iridescens) through the Andean cordillera (e.g., S. festae) to the eastern lowlands (S. aculeatus). The species occur mostly on the western slopes of the Cordillera Occidental and in the inter-Andean ba- sins. Ten species (S. carrioni, chota, guentheri, haenschi, humeralis, iridescens, limitaris, rhodomelas, simonsti, and varius) occur at elevations below 2200 m on the western versant of the Andes, whereas only two species (S. carrioni and aculeatus) occur at similar elevations on the eastern versant of the Andes (Fig. 21). Moreover, six species (S. chota, festae, guentheri, humeralis, ornatus and rhodomelas) occupy inter-Andean basins. This pattern contrasts with the trans-Andean distribution of other Ecuadorian herpetofaunal components such as Anolis with 21 western species versus six eastern species (Almendariz, 1992), Proctoporus with nine western species versus seven east- ern species (Kizirian, 1996), Eleutherodactylus with 61 west- ern species (Lynch and Duellman, 1997) versus 74 eastern species (updated from Coloma, 1991) or Colostethus with 15 western species versus 16 eastern species (Coloma, 1995). These lizards occur at elevations between sea level (S. iridescens) and 3890 m (S. guentheri). Stenocercus iridescens has the largest elevational range (0-2000 m), whereas S. haenschi is known from a single elevation (Fig. 22). Exami- nation of altitudinal distributions by 100 m increments re- veals that the greatest concentration of species (71%) is between 1500 m and 2500 m. No species of Stenocercus is known to have a distribu- tional range extending from Colombia to Peru. Three spe- cies occur in both Ecuador and Peru; S. iridescens and S. limitaris occur in western Ecuador and northwestern Peru (Cadle, 1991, 1998) and S. aculeatus inhabits the eastern slopes of the Ecuadorian and Peruvian Andes (Peters and Donoso-Barros, 1970). Only one Ecuadorian species, S. guenthert, also is known from Colombia (Ayala and Castro, 1982; Castro and Granados, 1993). Nevertheless, S. angel probably occurs in southern Colombia because its type locality lies close to the Colombian border. a) 2 [S) ® [om 2) re) ® a £ 5 Zz A B Cc D E F G Geographic Zone Fig. 21. Trans-Andean distribution of the genus Stenocercus in Ecuador. ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS 35 4000 3500 3000 2500 S. rhodomelas S. varius 1) Cc o 1S) 3 8 = (dp) 2000 Elevation (m) # S. carrioni S. festae 1500 # S. aculeatus S. humeralis S. guentheri 8) S. haenschi S. ornatus 1000 500 Species Fig. 22. Altitudinal distribution of species of Stenocercus in Ecuador. Distribution patterns of Stenocercus in southern Ecua- dor and northern Peru are influenced by the Huancabamba Depression, a region widely recognized as a major biogeo- graphic discontinuity for Andean organisms (Duellman, 1979). No species of Stenocercus is presently known from north and south of the depression (Cadle, 1991). Ten species (71%) are endemic to Ecuador. Most of the Ecuadorian species occur south of 1°S Lat. Stenocercus iridescens has the greatest latitudinal distribution range (from 00°53' N to 05°55' S in northwestern Peru), whereas S. haenschi is known only from its type locality. Most of the Ecuadorian species of Stenocercus resemble the Andean liz- ard genera Pholidobolus (Montanucci, 1973) and Proctoporus (Kizirian, 1996) in having limited geographic distributions. Table 14. Distribution of Stenocercus within life zones in Ecuador. Life Zone Number of Species Thorny Premontane Thicket Tropical Dry Forest Tropical Very Dry Forest Premontane Thorny Forest Humid Premontane Forest Very Humid Premontane Forest Pluvial Premontane Forest Low Thorny Montane Steppe Montane Steppe Low Dry Montane Forest Low Humid Montane Forest Low Very Humid Montane Forest Humid Montane Forest Very Humid Montane Forest Sub-Andean Montane Forest PENP WORF PREP WAN Nevertheless, additional collecting is necessary to obtain a clearer understanding of the distributional patterns of the species of Stenocercus. ECOLOGICAL DISTRIBUTION The ecological distribution of the Ecuadorian Stenocercus species is described in the species accounts. For analytical purposes, I used the classification of life zones of Ecuador proposed by Canadas-Cruz (1983), which is based on the Holdridge (1947) system. The distribution of Stenocercus in Ecuador includes 15 life zones (Table 14). Most species inhabit premontane and montane forests. Dry life zones (annual mean precipitation < 1000 mm) are oc- cupied by nine species, six of which (S. angel, guenthert, humeralis, iridescens, ornatus, and rhodomelas) also occur in humid life zones (annual mean precipitation > 1000 mm). Only three species (S. chota, festae, and simonsit) are re- stricted to dry life zones. In contrast, five species (S. aculeatus, carrioni, haenschi, limitaris, and varius) are re- stricted to humid life zones. Within its distribution, Stenocercus iridescens inhabits the driest life zone (Tropical Desertic Thicket), whereas S. aculeatus inhabits the most humid zone (Pluvial Premontane Forest). KEY TO SPECIES OF STENOCERCUS OF ECUADOR 1. Scales on posterior surface of thigh imbricate .......... 2 Scales on posterior surface of thigh granular.......... 10 2. One row of enlarged supraoculars, each more than twice size of scales in adjacent TOWS ...........:ccsceeeeeeees 3 Row of enlarged supraoculars absent ....... seucarstteansebes 5 3. Postfemoral pocket Type 4 or 5 «0... Postfemoral pocket of other types 4. Postfemoral pocket Type 3; projecting angulate HEIEMDOUES, ccnseseccocesnc0s05300000C01202009500795500s0070000000 S. aculeatus Postfemoral pocket Type 1; projecting angulate (EVEN OYONEUS GI SVE coscrocenceacc6s9%650130050003092050505006 S. iridescens 5. Posthumeral pocket Type 4; postfemoral pocket Tf XB cn cncoqasco nace sctecncatosaocnge vas eoco es acu onto oncooasoscos0GaueceIA060: 6 Posthumeral and postfemoral pockets of other types onde de beuo exes tenet Gea sasse atid neuatislaete ir drrreattensarsaravanvee secseatigess 7 6. Antehumeral fold Weak 0.0.0.0... cece S. ornatus Antehumeral fold absent..............::.00000 S. rhodomelas 7. Postfemoral pocket Type 3; gular region of males black; ventral scales of juveniles keeled ..................... S. festae Postfemoral pocket Type 1 or 2; gular region of males 36 SCIENTIFIC PAPERS, NATURAL History MusEeuM, THE UNIVERSITY OF KANSAS usually not black; ventral scales of juveniles and adults moral yas tn @ Othe sersesneceestenseerencses ses conseenscaeereee cee 8 8. Females with large, black spots in gular region; males with dark midventral stripe and large blue ventrolat- ef all IMALKS)«-.sevcseecali sic nen enters ee ties S. chota Females lacking black spots in gular region; males usu- ally lacking blue marks on vente? .............:ccccceseseeeees 9 9. Scales around middle of the body 59 or more............. ssuesvoes isivescstsssesi sbeowpasvontasisvendnestaccvsestiaeaee sets ttr S. guentheri Scales around middle of the body fewer than 59........ sae od pestnagnosibebeisan sapvaseviespezss uo uvve ceeve saan Tanor este S. angel 10. Scales around middle of the body fewer than 62........ 6906036366 0830%090 5653000 CASH GoTIDCERSGNOSonHEBaNOoSoNONoAANE S. haenschi Scales around middle of the body 62 or more......... 11 11. Caudal scales mucronate; two caudal whorls per AUItOLOMMCISCPTMEM tesreccesssseesesesre ene tere 12 Caudal scales not mucronate; three caudal whorls per BUUKON WONT SSA TIES acronnonosocenn-nosococaconsocccosn-cosososocectoscs0" 13 12. Vertebral row discontinuous; distal caudal whorls clearly alternating in Size ............00:ceeeeee S. carrioni Vertebral row continuous; caudal whorls equal in size FE oc CEC EEE EEEEOEC EAC OCO BE DerEE Sco Boor ssebcorcccoacn: S. simonsii 13. Scales around middle of the body 98 or more............. essSeusadessdaze fecazacabracs domes tistet tests aioe sto eee ae S. humeralis Scales around middle of the body fewer than 98........ ste secedesssteecberesetenessssustevesesesteckeets sixth wits aetna S. varius LITERATURE CITED Almendariz, A. 1992. “1991”. Anfibios y reptiles. Pp. 89-162 in R. Barriga, A. Almendariz, and L. Albuja, Lista de vertebrados de Ecuador. Revista Politécnica, Escuela Politécnica Nacional, 16 (3): PAGES ? Avila-Pires, T. C. S. 1995. Lizards of Brazilian Amazonia (Reptilia: Squamata). Nationaal Natuurhistorisch Museum, Zoologische Verhandelingen 299:1—706. Ayala, S., and F. Castro. 1982. Nueva especie de (Sauria: Iguanidae) lagarto collarejo de la zona sur andina de Colombia. Caldasia 13:473-478. Boulenger, G. A. 1880. Reptiles et batraciens recueillis par M. Emile de Ville dans les Andes de L’ Equateur. Bulletin de la Société Zoologique de France 5:43. Boulenger, G. A. 1885. Catalogue of the Lizards in the British Museum (Natu- ral History), Vol. 2, pp.1-497. London: Taylor and Francis. Boulenger, G. A. 1899. Descriptions of new reptiles and batrachians col- lected by Mr. P. O. Simons in the Andes of Ecuador. Annals and Magazine of Natural History 4:454—457. Burt, C. E., and M. D. Burt. 1931. South American lizards in the collection of the American Museum of Natural History. Bulletin of the Ameri- can Museum of Natural History 61(7):227-395. Cadle, J. E. 1991. Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from northern Peru: new species and comments on relationships and distribution patterns. Proceedings of the Acad- emy of Natural Sciences of Philadelphia 143:1—-96. Cadle, J. E. 1998. New species of lizards, genus Stenocercus (Iguania: Tropiduridae), from western Ecuador and Peru. Bulletin of the Mu- seum of Comparative Zoology 155(6):257-297. Canadas-Cruz, L. 1983. El Mapa Bioclimatico y Ecologico del Ecuador. Quito: Editoriales Asociados Cia. Ltda. Castro, F., and H. Granados. 1993. Distribucién de Stenocercus guentheri (Sauria: Iguanidae) en el sur de los Andes de Colombia. Caldasia 17(2):295-300. Coello, F. 1994. Plan de manejo de la reserva ecologica El Angel. Convenio MBS-INEFAN-IICA. Quito, Ecuador. Coloma, L. A. 1991. Anfibios del Ecuador: lista de especies, ubicacion altitudinal y referencias bibliograficas. Reportes Técnicos Ecociencia 2:146. Coloma, L. A. 1995. Ecuadorian frogs of the genus Colostethus (Anura: Dendrobatidae). University of Kansas, Natural History Museum, Miscellaneous Publication 87:1-72. Corredor, V. 1983. Una nueva especie de Stenocercus (Sauria: Iguanidae) de la cordillera oriental de Colombia. Lozania 37:1-10. De Vries, T., J. Black, C. de Solis, and C. Hernandez. 1983. Historia natu- ral del curiquingue (Phalcoboenus carunculatus) en los paramos del Antisana y Cotopaxi del Ecuador. Ediciones de la Universidad Catolica de Quito, Ecuador, pp. 1-83. Duellman, W. E. 1979. The herpetofauna of the Andes: patterns of distri- bution, origin, differentiation and present communities. Pp. 371— 459 in W. E. Duellman (ed.), The South American Herpetofauna: its Origin, Evolution and Dispersal. University of Kansas, Natural His- tory Museum, Monograph 7:1-485. Dumeril, A. M., and G. Bibron. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4, Paris: Librairie Encyclopedique de Roret. Etheridge, R. 1966. The systematic relationships of West Indian and South American lizards referred to the iguanid genus Leiocephalus. Copeia 1966:79-91. Fritts, T. H. 1972. New species of lizards of the genus Stenocercus from Peru (Sauria: Iguanidae). Occasional Papers, Museum of Natural History, University of Kansas 10:1—21. Fritts, T. H. 1974. A multivariate and evolutionary analysis of the andean iguanid lizards of the genus Stenocercus. San Diego Society of Natu- ral History, Memoir 7:1-89. Frost, D. R. 1988. A Phylogenetic Analysis of the Tropidurus Group of Iguanian Lizards, with Comments on the Relationships Within the Iguania (Squamata). Ph. D. Dissertation, University of Kansas. xi + 350 pp. Frost, D. R 1992 Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania: Tropiduridae). American Museum Novitates 3033:1-68. Frost, D. R., and R. Etheridge. 1989. A phylogenetic analysis and tax- onomy of Iguanian lizards (Reptilia: Squamata). Miscellaneous Pub- lication, Museum of Natural History, University of Kansas 81:1-65. Gray, J. E. 1827. A description of new species of saurian reptiles; with a revision of the species of chamaleons. Annals of Philosophy 2:207— 214. Gray, J. E. 1845. Catalogue of the Specimens of Lizards in the Collection of the British Museum. London: British Museum. Gunther, A. 1859a. List of the cold-blooded vertebrata collected by Mr. Fraser in the Andes of western Ecuador. Proceedings of the Zoo- logical Society of London 1859:89-93. Gunther, A. 1859b. Second list of the cold-blooded vertebrata collected by Mr. Fraser in the Andes of western Ecuador. Proceedings of the Zoological Society of London 1859:402-427. Holdridge. L. R. 1947. Determination of world plant formations from simple climatic data. Science 105:367-368. Kizirian, D. A. 1996. A review of Ecuadorian Proctoporus (Squamata: Gymnophthalmidae) with descriptions of nine new species. Her- petological Monographs 10:85—-155. Lynch, J. D.,and W. E. Duellman. 1997. Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador: systematics, ecology and bio- geography. Special Publication, Natural History Museum, Univer- sity of Kansas 23:iv + 236. Montanucci, R. 1973. Systematics and evolution of the andean lizard ge- nus Pholidobolus (Sauria: Teiidae). Miscellaneous Publication, Mu- seum of Natural History, University of Kansas 59:1-52. Miyata, K. 1982. A check list of the amphibians and reptiles of Ecuador ECUADORIAN LIZARDS OF THE GENUS STENOCERCUS Bi with a bibliography of Ecuadorian herpetology. Smithsonian Her- petological Information Service 54:1—70. O'Shaughnessy, A. W. E. 1879. Description of new species of lizards in the collection of the British Museum. Annals and Magazine of Natu- ral History 4:295-303. O'Shaughnessy, A. W. E. 1881. An account of the collection of lizards made by Mr. Buckley in Ecuador, and now in the British Museum, with descriptions of the new species. Proceedings of the Zoological Society of London 1881:227-245. Parker, H. W. 1934. Reptiles and amphibians from southern Ecuador. Annals and Magazine of Natural History 14:268-270. Peracca, M.G. 1897. Viaggio del Dr. Enrico Festa nell ‘Ecuador e regioni vicine. Bollettino del Musei di Zoologia ed Anatomia Comparata della Universita di Torino 12:1—20. Peters, J. A. 1967. The lizards of Ecuador, a check list and key. Proceed- ings of the United States National Museum 119:1—49. Peters, J. A., and R. Donoso-Barros. 1970. Catalogue of the Neotropical Squamata. Part 2, lizards and amphisbaenians. United States Na- tional Museum Bulletin 297:1-293. Trueb, L., and W. E. Duellman. 1971. A synopsis of neotropical hylid frogs, genus Osteocephalus. Occasional Papers, Museum of Natural His- tory, University of Kansas 1:1-47. Uzzell, T. 1973. A revision of lizards of the genus Prionodactylus, with a new genus for P. leucostictus and notes on the genus Euspondylus (Sauria, Teiidae). Postilla 159:1-67. Vitt, L. J., and S. de la Torre. 1996. Guia para la investigacion de las lagartijas de Cuyabeno. Monografias del Museo de Zoologia de la Pontificia Universidad Catolica del Ecuador 1:1-165. Werner, F. 1901. Uber Reptilien und Batrachier aus Ecuador und Neu Guinea. Verhandlungen der zoologisch-botanischen Gesellschaft in Wien 51:593-614. APPENDIX SPECIMENS EXAMINED All specimens are from Ecuador unless noted other- wise. Species are arranged alphabetically. Localities are arranged alphabetically within provinces, which are also arranged alphabetically within each species. Stenocercus aculeatus (12 specimens) Provincia Pastaza: Mera, 1123 m, EPN 1153, 4050, 4051; 3 km S Puyo, 920 m, KU 127094; Rio Solis, EPN 5902-5904; Veracruz, 950 m, KU 121092. Provincia Tungurahua: Rio Negro, 01°24’ S, 7812' W, QCAZ 1635; no spe- cific locality, ZMB 16594. Peru: Departamento San Martin: Moyobamba, BM 1946.8.12.33-34 (syntypes). Stenocercus angel (10 specimens) Provincia Carchi: El Angel, 00°38' N, 77°56' W, 3015 m, QCAZ 1358 (paratype); 8 km NE El Angel on road El Angel-Tulcan, 00°40' N, 77°52' W, 3560 m, QCAZ 1354 (paratype), 3732 (paratype), 3733 (holotype), 4117— 4119 (paratypes); 13.6 km W Tulcan on road Tulcan-Tufino, 00 “49'N, 77°49' W, 3040 m, QCAZ 3792 (paratype); Estacion Biologica Guanderas, QCAZ 3777 (paratype). Provincia Sucumbios: El Playon de San Francisco, 00°38' N, 77°38' W, 3300 m, QCAZ 1322 (paratype). Stenocercus chota (55 specimens) Provincia Carchi: La Concepcion, 00°35' N, 78°07' W, 1575 m, MZUT R2154.1-5, R2154.25-29 (paratypes). Provincia Esmeraldas: Rio Cachabi, EPN 5858-5860 (paratypes), 5862-5864 (paratypes). Provincia Imbabura: Ambuqui, 00°27' N, 78°01' W, 1780 m, QCAZ 799 (paratype); 6.5 km E Panamerican Highway on road Ambuqui-Monte Olivo, 1940 m, QCAZ 806 (paratype), 897-902 (paratypes), 3791 (paratype), 3794 (paratype); Chota, 00°28' N, 78°04' W, QCAZ 2654 (paratype), 2655 (paratype), 2773- 2778 (paratypes); 5 km E Chota on Panamerican Highway, 00 °28' N, 78°01’ W, QCAZ 2768 (paratype), 3755 (paratype), 3757 (paratype), 3762-3767 (paratypes), 3769-3776 (paratypes), 3768 (holotype); Salinas, 00°30' N, 78°08' W, QCAZ 4162; Tumbabiro, 00°28' N, 78°12' W, QCAZ 4161; sur- roundings of Yaguarcocha, EPN 5848 (paratype). Stenocercus festae (55 specimens) Provincia Azuay: Cuenca, 02°53' S, 78°59’ W, 2530 m, UDAR 11; 4 km E Cuenca, 2540 m, KU 134574-134579, 134582, 134583, 134585-134592, 134594; Laguna Zurucuchu, 3200 m, KU 121094; 3.1 km E Sigsig on road Sigsig-Shuso, Rio Santa Barbara, 2450 m, QCAZ 3789; 4 km W San Crist6bal, 2500 m, KU 121095; Sevilla de Oro, 02°48' S, 78°39' W, 2630 m, QCAZ 4059 (neotype); Sigsig-Shiguinda road, 3200 m, QCAZ 1337; Ucubamba, 02°52'S, 78°54' W, 2530 m, UDAR5; no specific locality, USNM 201222, 201223. Provincia Canar: 3 km S Azogues, 2500 m, KU 134602, 134604-134607, 134609; Canar, 02°33' S, 78°56' W, OCAZ 1409; La Carboneria, 02°30' S, 79°01' W, QCAZ 3117; Laguna Culebrillas, 02°25'S, 78°51' W, QCAZ 1346-1348; Cebadas, Pacupala, EPN 2700. Provincia Loja: Chuquiribamba, 03°50' S, 79°20' W, 2700 m, QCAZ 1340; Manu, 03 ae Sy 79°24' W, 2200 m, QCAZ 3599-3602; Saraguro, 03°36' S, 79°13' W, 2500 m, KU 134120, 134122-134126, QCAZ 3113; 14 km NE Urdaneta, 3050 m, KU 179419. Stenocercus guentheri (77 specimens) Provincia Chimborazo: 14.5 km N Tixan on Panamerican Highway, 3200 m, QCAZ 3659-3661. Provincia Cotopaxi: near Panamerican High- way on the Cotopaxi National Park, 3200 m, FHGO 629; Cotopaxi Na- tional Park, approx. 4000 m, QCAZ 1109. Provincia Imbabura: Atuntaqui, 2387 m, QCAZ 776; 7,5 km N Otavalo on Panamerican Highway, QCAZ 3761; Tabacundo-Mojanda road, 3150 m, QCAZ 3793. Provincia Pichincha: Cayambe, SMF 11162; Cayambe volcano, 3500 m, FHGO 1136; Guayllabamba, 2139 m, QCAZ 718, 777, 779, 782; Ilalo, Hacienda Chuspiyacu, QCAZ 722; Iliniza Sur, QCAZ 730; Jerusalem, 2578 m, QCAZ 1323; Lloa, 00°15' S, 78°35’W, 3060 m, QCAZ 4108; Machachi, 2940 m, QCAZ 720, 736, 758, 775, 778, 780, 781, 783, 784; Pintag-Antisana road, 2880 m, QCAZ 2808; Pusuqui, QCAZ 4108; Quito, 2810 m, EPN 5900, QCAZ 432, 728, 737, 2857, SMF 60592; San Antonio, 00°01' S, 78°27' W, QCAZ 713-716, 738, 740-754, 1357, 1400, 2163, 2199, BM 58.7.25.16, 58.7.25.16a, 58.7.25.18, 59.9.20.6, 60.6.16.18, 60.6.16.20—21 (syntypes); Uyumbicho, QCAZ 760. Provincia Tungurahua: Ambato, 01 AlSy Sy, Wy’ W, 2575 m, EPN 5898, 5899; Picaihua, EPN 5887-5889; Urbina, QCAZ 2858; no specific locality, FHGO 522, 852, 1493, SMF 53199. Stenocercus haenschi (1 specimen) Provincia Bolivar: Balsapamba, 01 °47' S, 79°10' W, 750 m, ZMB 16595 (holotype). Stenocercus humeralis (42 specimens) Provincia Loja: Catamayo-Jimbilla road, EPN 5824-5843; Loja, 04°00' S, 79°12’ W, 2064 m, BM 1946.8.11.76 (syntype), EPN 1343, 5807, 5808, 5810-5813, 5815, FHGO 1494, 1495, KU 121136, 121138, 134000, 134002; 5 km N Loja, 2150 m, KU 134003, 134005; 2 km E Loja, 2200 m, KU 121137; 2.7 km E Loja, 2135 m, KU 141162; 12.2 kmS Loja on road to Vilcabamba, 2275 m, KU 141163; Malacatos, EPN 1270, 5809. Stenocercus iridescens (42 specimens) Provincia Azuay: Tamarindo, FHGO 416. Provincia Cotopaxt: La Mana, 00°56' S, 79°13' W, 300 m, QCAZ 2767, 3052. Provincia El Oro: 15 km E Pasaje on road Pasaje-Pan de Azucar, 90 m, QCAZ 3620; Pinas, 03 “41'S, 79°41' W, 1410 m, FHGO 1089. Provincia Esmeraldas: Atacames, 00°52' N, 79°50' W, 8 m, EPN 5909; La Union, 00°49' N, 79°51' W, 10 m, FHGO 97; Rio Tiaone EPN 5906, 5908; Same, 00°51' N, 79°54' W, 0 m, QCAZ 721; Tonsupa, 00°53' N, 79°45' W, 0 m, QCAZ 762, 763, 804, 805. Provincia Guayas: Balzar, 01°22'S,79°54' W, 37 m, EPN 5925-5927, 5929-5931, QCAZ 735; Cerro Blanco, EPN 5005, 5007-5009. Provincia Los Rios: Centro Cientifico Rio Palenque, 00°35' S, 79°22' W, QCAZ 431, 2205, 2206, 2212; Jauneche, EPN 5004; Patricia Pilar, 00°33'S, 79°22' W, OCAZ 97; Ventanas, 01°27' S, 79°27' W, QCAZ 1655. Provincia Manabi: Cabo Pasado, 00°22' S, 80°29' W, 20 m, QCAZ 3322; Cerro San Sebastian, EPN 5014; Puerto Rico, 01°38' S, 80°49' W, 12 m, OQCAZ 1634; 27 km N San Vicente on road San Vicente-Pedernales, 00°22' S, 80°26' W, 50 m, OCAZ 3329, 3330, 3343; 32 km N San Vicente on road San Vicente-Pedernales, 00 °20' S, 80°21' W, 183 m, QCAZ 3314. Western Ecuador: BM 60.6.16.2-4 (syntypes). 38 ScIENTIFIC Papers, NATURAL History Museum, THE UNIVERSITY OF KANSAS Stenocercus ornatus (41 specimens) Provincia Loja: Catamayo, 2280 m, KU 141167; Cerro Uritusinga, 3000. m, QCAZ 2020; Cerro Villonaco, EPN 3540; Loja, 04°00' S, 79°12' W, 2064 m, KU 121126; 4.6 km N Loja, 2065 m, KU 141168-141170; 5 km N Loja, KU 134150, 134151, 134153, 134154; 2 km E Loja, 2200 m, KU 121127, 121129-121134; 6 km S Loja on road Loja-Vilcabamba, 2300 m, FHGO 585; 3 km W Loja, 2150, KU 134127, 134129-134131, 134134, 134138, 134139; 12 km W Loja, KU 134148; 15 km W Loja, KU 134140-134144, 134149; 10.6 km S Yangana, 2190 m, QCAZ 3790; Vilcabamba, 1500 m, FHGO 405, 679, 1161; no specific locality, EPN 5877-5880. Stenocercus rhodomelas (32 specimens) Provincia Azuay: 4.8 km W Abdon Calderon, 1435 m, KU 152183; 1.1-2.7 km SW Catavina, 1310 m, KU 152188; 2.7-3.5 km SW Catavina, 1250 m, KU 152185, 152187; 5 km S Nabon on road Nabén-San Isidro, 03°20'S, 79°04' W, UDAR 10; Ona, BM 1946.8.29.77-78 (syntypes); N Ona, 1885 m, KU 141164, 141166; 50.5 km E Pasaje, 730 m, KU 152177; Rio Herp aLess. L25 T67 2000 Fea can a ee a | i HN 4 062 4 genus Ste in Le6én, 11.8 km N Buenos Aires, 1940 m, KU 202945, 202946; Rio Leon, 12.5 km N Ona, 1920 m, KU 142699-142701; 1.6 km W Minas at Rio Minas, 1410 m, KU 152178, 152179; Santa Isabel, 03°16' S, 79°19' W, QCAZ 3076; Valle de Yunguilla, Chalcapac, 1550 m, QCAZ 3663. Provincia Loja: Valle de Casanga, EPN 5910-5921. Stenocercus simonsii (1 specimen) Provincia Azuay: Ona, 6500 ft (1981.2 m), BM 1946.8.11.73 (syntype). Stenocercus varius (25 specimens) Provincia Cotopaxi: Penas Coloradas, QCAZ 1695; Reserva Integral de Bosque Nublado Otonga, 00°44'S, 78°59' W, 2000-2200 m, QCAZ 3118, 3845; San Francisco de las Pampas, 00°26' S, 78°57' W, QCAZ 86-91, 2015. Provincia Pichincha: Estacion Forestal La Favorita, 00°14' S, 78°46' W, 1900 m, FHGO 354, 412, 424, 445; Reserva Floristica Ecologica Rio Guajalito, 00°14' S, 78°48' W, 1840 m, QCAZ 717, 719, 1334, 3046, FHGO 337; Rio Blanco, EPN 5932; Tandapi, 00°25’S, 78°47' W, 1460 m, QCAZ 590, 591, 593-596. \ PUBLICATIONS OF THE NATURAL HISTORY MUSEUM, THE UNIVERSITY OF KANSAS The University of Kansas Publications, Museum of Natural History, beginning with Volume 1 in 1946, was discontinued with Volume 20 in 1971. Shorter research papers formerly published in the above series were published as The University of Kansas Natural History Museum Occasional Papers until Number 180 in Decem- ber 1996. 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