A RE VI O R B Nl m s e Til inn ©i Comparative /^oology US ISSN Cambridge, Mass DESCRIPTIONS OF 0006-9698 Number June 2009 516 TWO NOMEN NUDUM SPECIES OF ANOLIS LIZARD FROM NORTHWESTERN SOUTH AMERICA Steven Poe, Abstract We Julian Velasco, Kenneth Miyata, - and Ernest E Williams ' describe two northern South American Pacific lowland species of Anolis that historically have been associated with names that are currently nomina nuda Anolis lyra new species differs primarily in possessing smaller dorsal scales on the head and body and is similar to Anolis vittigerus but in color pattern Anolis anchicayae new and color pattern We scored the new species for skeletal and external morphological phylogenetic characters and performed a parsimony analysis with 251 additional Anolis species using molecular and morphological data Anolis anchicayae is sister to A peraccae in a basal group of South American "alpha" Anolis ("dactyloids," "latifrons" group) Anolis lyra is placed with other lemurinus-group Anolis within the large clade of mainland "beta" Anolis {"Norops") species is similar to Anolis peraccae Key words: and Anolis fasciatus but differs primarily in size Anolis anchicayae; Anolis lyra; Colombia; Ecuador Resumen Describimos dos especies de Anolis de zonas bajas del Pacifico de Colombia y Ecuador que tradicionalmente han estado asociadas nombres en nomen nundum Anolis lyra sp nov es parecido a Anolis vittigerus pero se diferencia principalmente en que presenta pequenas, y en el patron de coloracion Anolis anchicayae sp las escamas del dorso de la cabeza y el cuerpo mas nov es parecido a Anolis peraccae y Anolis fasciatus se diferencia principalmente en el tamano y el patron de coloracion Se codificaron caracteres filogeneticos de morfologia externa y osteologia y se realizo un analisis de parsimonia 25 especies adicionales de Anolis usando una combination de datos morfologicos y moleculares Anolis anchicayae es taxon hermano de A peraccae dentro de pero un grupo basal de especies suramericanas de Anolis "alfa." Anolis lyra esta relacionado otras especies del grupo lemurinus dentro de un clado mas grande compuesto por especies continentales de Anolis "beta." Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131, U.S.A.; e-mail: anolis@unm.edu Grupo de Ecologia Animal, Departamento de Biologia, Universidad del Valle, A A 25360, Cali, Colombia Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A Williams and Miyata are deceased and are included as coauthors for the following contributions Poe was able to diagnose each species and identify MCZ specimens on the basis of Williams' unpublished notes, which clearly were influenced by Miyata's work on the second species described herein Additionally, Williams is responsible for Figure Subsequent to submission of this manuscript, the editor provided Poe with a chapter of Miyata's Ph.D thesis, never published, describing the second of the two species described herein Poe incorporated information from this manuscript into the diagnosis of this species ® The President and Fellows of Harvard College 2009 BREVIORA Ernest Williams and collaborators pub- from 1983 lished a series of papers to 1992 "new or problematic" Anolis from describing Colombia These papers reviewed and expanded the Colombian anoline fauna and established the country a as diversity for the Anolis clade center of The publica- and other taxonomic works on Colombian lizards (e.g., Ayala and Castro, 1983) prompted the need tion of these descriptions for a summary of information available No 516 the anterior of the cloaca measured from the the knee, with the limb bent at a 90-degree Head width was measured angle broadest of the part head, made with parisons were in quotes ("") and associated with authors to signal that descriptions were not yet published but were forthcoming Pub- the Com- preserved material and with published species, species descrip- tions, preserved material, or occur in Colombia This listed the of the putative closest relatives of the new terminology included eight at between posterolateral corners of the orbits distant relatives (online list was length anterior edge of the ear Femoral length was measured from the midline of the venter to Stephen Ayala (1986) obliged and produced a list of the 205 lizard species thought to species of Anolis (and species in other clades) Head of the snout to the tip and characters Williams of descriptions description et is anoline mainly by Williams 1996) al, Scale 1) used standards established follow (e.g., both of more Appendix species for Skeletal lizards given in terms of Poe's (1998, lished descriptions of Anolis species associated 2004) and Etheridge's (1959) characters See with four of these names soon followed (A those papers for danieli Williams 1988, A ruizi Rueda and Williams 1986, A lamari Williams 1992, A medemi Ayala and Williams 1988), but the other names have languished without descriptions as nomina nuda for over 20 years The aim of this paper is to rectify this situation with respect to two of the remaining names well known to Colombian herpetologists more detailed descriptions of skeletal conditions and alternative conditions in Anolis We scored the new mor- species for the phological phylogenetic characters of Poe (2004; Appendix mony analysis of these taxa and performed a parsiand 251 other of Anolis and eight outgroups with species the of use 1) 1,291 parsimony-informative some characters from published sources and of our additional unpublished morphological MATERIALS AND METHODS data This dataset includes 91 morphological We consider species to be evolutionary and lineages (Simpson, 1961; Wiley, operationalize concept by identifying species on the this basis of consistent differences between populations That that 1978) is, we hypothesize populations that are diagnosable by major differences in the frequencies of traits characters (Poe, 2004), 998 sites of (NADH dehydrogenase transfer RNAs; Nicholson 182 sites scribed Codes of nuclear tracer et DNA region; mtDNA subunit al , five 2, 2005), (internal Nicholson, and tran- 2002) for morphological data for A anchi- cayae and A lyra are in online Table PAUP* We are distinct evolutionary lineages or species used Wiens and Servedio, 2000) Measurements were made with digital calipers on preserved specimens and are given in millimeters (mm), usually to the nearest 0.1 mm Snout-vent length (SVL) was measured from the tip of the snout to perform a heuristic search with additions of taxa and treebranch swapping bisection-reconnection (see 2002) version 4.0b 10 (Swofford, to 100 random We also used PAUPrat (Sikes and Lewis, 2001) to perform several runs of the parsi- mony ratchet (Nixon, 1999) to find optimal NEW SOUTH AMERICAN 2009 ANOLIS Colombia, Valle, Rio Blanco, road between Cali and Anchicaya (where road crosses river), ca 500 m along river just above bridge on banana plant, 12 May 1980, H Carvajal, D Harris, S Ayala; MCZ 1597678, Colombia, Valle, Anchicaya Hydroelectric Central, June 1976, Figure Anolis anchicayae new species, male Photo by Julian Velasco DESCRIPTION OF NEW H Chin, F Castro; MCZ 160230, Colombia, Valle, Bajo Calima, 14 August 1977, H Chin; MCZ 160231-3, Colombia, Choco, Bahia Solano, near Jayita, 14-16 October 1977, F Castro; MCZ 160633, Colombia, Valle, Anchicaya, December 77, SPECIES Anolis anchicayae, new species Figures 1-5 H Chin, F Castro; ICN 6115, Colombia, Choco, Bahia Solano, near 160234, adult male, Co- lombia, Valle, San Isidro, 1978, August 1977, Helen Chin Paratypes Jayita, MCZ 112426-7, Colombia, Anolis anchicayae new species, Renjifo A peraccae and A fasciatus species share keeled dorsal double ** J Diagnosis Anolis anchicayae Choco, Rio San Juan, Carlo Docordo between Cucurrupi and Noanama, 24 December 1968, B Malkin; MCZ 158385-7, Figure Castillo, F Castro, Valle, San Isidro, Carton, and August 1977, H Chin; MCZ 160008-12, Colombia, Valle, Granje Bajo Central, 1976- (20%-35% of characters reweighted, 150-200 replicates per run) MCZ J 159997-8, 160234-9, Colom- bia, trees Holotype MCZ H Chin; MCZ is similar to These three head scales, row of middorsal caudal fSsk 145362 Tail is missing in this specimen a scales, BREVIORA Figure Anolis anchicayae, MCZ No 516 145362, dorsal head scales most absence of dewlap in females, absence of supralabial Anolis anchicayae caudal transverse vertebral processes poste- distinguished from A peraccae by larger size riorly, and an elongate anterior nasal scale that reaches the rostral but does not contact the sulcus between the rostral and first (to 54 mm SVL in A A anchicayae; Fig 6) anchicayae is is peraccae; to 63 easily mm and color pattern in {A predominantly yellow-green on NEW SOUTH AMERICAN ANOLIS 2009 Figure Anolis anchicayae, MCZ 145362, lateral head scales from eye and a yellow-green dewlap in males; A peraccae is predominantly brown on its flanks with no shoulder ocellus and a cream-colored male dewlap) Anolis anchicayae is most easily distinguished from A fasciatus by size (to 72 mm SVL in A fasciatus; to 63 mm in A anchicayae) and male dewlap color (white with gray scales in its flanks with an elongate ocellus to shoulder A fasciatus, yellow-green in A anchicayae) Anolis anchicayae can be distinguished from other South American Anolis as follows (groupings are informal according to Savage and Guyer [1989] and others and used only for organizational purposes; no relationships are implied): from "beta" anoles (= "Norops"; with which it is unlikely to be confused) by the absence of transverse processes on posterior caudal vertebrae (present in beta from phenacosaur and tigrinusgroup Anolis {A heterodermus, A inderenae, Anolis); A nicefori, A bellipeniculus, A carlostoddi, A euskalerriari, A neblininus, A orcesi, A Figure head scales Anolis anchicayae, MCZ 145362, ventral tetarii, A vanzolinii, A ruizi, A solitarius, A menta, A nasofrontal, A pseudotigrinus, A BREVIORA No 516 MALES SVL 42mm 44 46 FEMALES 50 48 56 54 52 30 N = SVL 42mm 44 60 58 46 48 50 52 54 50 52 54 56 58 Anolis peraccae Norino; Colombia 30 20 10 50 Anolis per Plchincha; N»I8 40 N = 28 30 20 10 % K%4l%!*'&3kS£ ^•i.!':.:.:.:.**:'*':'*.* SVL 42mm 44 Figure 46 48 50 54 52 56 58 SVL 42mm 44 60 46 48 56 58 body sizes within populations for samples of A peraccae and A anchicayae, showing The specimens used to construct this table are unknown because they were not listed in Distribution of larger size of A anchicayae Williams' notes menta, A lamari, A vaupesianus, scales separating the supraorbital semicircles A jacare, A umbrivagus, A paravertebralis, A (supraorbital semicircles in contact in these tigrinus, A by williamsmitteremeierorum) possessing species); from other punctatus- and aequa- strongly keeled head scales superior to the torialis-group Anolis (A chocorum, A cali- smooth or pustulose in and tigrinus-group Anolis); from latifrons- and some aequatorialis-group mae, A antioquiae, A Anolis (A latifrons, A frenatus, A fraseri, these species); A proboscis Anolis (A proboscis, A orbits (mostly phenacosaur danieli, A apollinaris, A casildae, A ialis, dewlap fitchi, A aequator- megalopithecus) in the lack of a A in females (female dewlap present in from A punctatus and the laevis, A on squamulatus, A princeps, A purpurescens, phyllorhinus) by the absence of swelling A agassizi, A eulaemus, A maculigula) by the snout of the male (swelling present in much (A mm SVL these males of these species); from mirus-gvoup Anolis {A mirus, A parilis) by the presence of a broad, raised toepad (toepad narrow and continuous with first phalanx in mirus group); from A chloris in body color and A A in soinii, A huilae, A boettgeri, A deltae, A A cuscoensis, transversalis, dissimilis, (> 100 from some punctatus-group Anolis smaller size species); sanlamartae) by displaying NEW SOUTH AMERICAN ANOLIS 2009 Moris; especially posteriorly; interparietal length 1.3 banded pattern of browns and greens in A anchicayae); from A festae and A nigroli- (1.2-1.6); three (3-4) scales separate interpa- pattern name neatus (the is (dark blotch these in anchicayae); from A absence in synonym of 1982) in male dewlap elongate species; absent in A caquetae A in a probable A festae; Williams, pattern green bright (solid between of contact and supraorbital semicircles; from A propinqus in number of subdigital interparietal lamellae (27 in A propinqus, from anchicayae); of condition rostral and nasal first in the (elongate scale ital and supraorbital semicircles; preoccip- absent; six (5-6) supralabials to center of eye; nine (8-10) postrostrals; six (6-9) post- some mentals; in slightly enlarged scales present supraocular disc, decreasing gradually in bordered medially by a partial row of size, small scales; mental partially divided poste- extending riorly, rostral, arc; beyond posterolaterally with posterior border in concave two (2-A) enlarged sublabials in contact with infralabials (or posteriorly separated by between small scales); dewlap reaches well posterior supralabial in A gemmosus; to axillae in males, absent in females; scales contacts nasal anterior gemmosus A the 16-20 in A rietal sulcus on dewlap in inferior to naris in A anchicayae) scales with some External Description of Holotype (paratype variation in parentheses; description is based pocket; pair of enlarged postcloacal scales in is separated on adults in from sulcus by small good condition: MCZ scale 160234, no axillary contact (or separated by small scales; absent in females); nuchal and dorsal crests absent; 159997-8, 160009-10, dorsal scales keeled; approximately 10 (5-12) 160232-3, 160238) Snout-vent enlarged middorsal rows, 160230, head length 12.0 (11.8-13.2; 0.23-0.25 SVL), width 7.1 (7.07.8; 0.14-0.15 SVL); ear height 1.3 (1.1-1.6; 0.02-0.03 SVL); femoral length 12.9 (13.616.0; 0.25-0.30 SVL); tail length 115 (102116; 1.96-2.26 SVL); fourth toe length 10.5 (10.7-11.9; 0.21-0.22 SVL), width 1.0 (1.01.3; 0.02 SVL) 52.0 (50.0-53.6); Dorsal head scales uni- to multicarinate above orbits, mostly smooth in frontal area, unicarinate present; scattered scales; 159767, 158386-7, length rows of approximately three on rostral snout; frontal slightly anteriorly; nine (8-12) depression overlaps scales; seven (5-7) loreal 5% six (6-8) longitu- of SVL; ventral scales in mostly transverse but some diagonal rows, smooth, eight (7-9) scales in 5% of SVL; anterior thigh scales large, keeled, overlapping, becoming smaller and nonoverlapping posteriorly; supradigitals multicarinate; toe- pads expanded; 19 (16-20) expanded lameallae under third and fourth phalanges of fourth toe (second and third phalanges of Williams [e.g., Williams et al, 1995]); tail with a double row of middorsal scales Skeletal Description (Based on mental scales across snout between second canthals; supraorbital semicircles separated by two (1-2) scales; suboculars in contact with supralabials; one (1-2) elongate supraciliary scale followed by small undifferentiated dinal rows in tons MCZ 112426-7, ICN 6115) Parietal roof dal with crests, a Dry 159767, flat, very Skele- 160012, with trapezoi- narrow posterior border, with no casquing, lacking crenulation on edges, not extending posteriorly over supraoccipital, with anterolateral corners rows; elongate anterior nasal scale contacts flush with posterolateral edges of frontal; from sulcus between pineal foramen at parietal-frontal suture (n rostral, rostral is and separated first supralabial by one small scale inferior to naris; supraorbital semicircles evident but not strongly differentiated, = 4) or in parietal (n = 1); dorsal skull bones smooth; postfrontal present; prefrontal separated from nasal by anterior extension of BREVIORA = frontal (n sutures anteriorly with frontal 2); crests parallel with nasal (n 2) or in contact on nasals = nasals; external absent; nares bordered posteriorly by nasals; dorsal on aspect of jugal terminates lateral surface No 516 ually (Etheridge's posteriorly lost [1959] "alpha" condition) Color in Life (Adapted from Field Notes and Color Photos by Velasco) Males brown middorsally interrupted by approximately 10 of postorbital; jugal does not contact squa- dark blotches that extend mosal; posterodorsal ramus of squamosal greenish-yellow with narrow diagonal rows laterally; flanks shorter than posteroventral ramus, separated of partially connected small brown spots; from elongate by supratemporal; posterior aspect of jugal mostly straight; epipterygoid contacts parietal dorsally; pterygoid and parietal palatine teeth absent; lateral edge of is vomer smooth, without posteriorly directed lat- processes; maxilla extends posteriorly eral beyond ectopterygoid on ventral surface of = skull (n 3) or reaches ectopterygoid (n = between basipterygoid processes of basisphenoid absent; lateral shelf of quadrate 2); crest no black pigment on surface of skull; nasals not overlap premaxilla dorsally; eye, even with vertical suture; approximately is level of parietal-frontal posteriormost mandibular tooth is = = or partially anterior partially posterior (n (n mandibles) mandibles) to anterior mylohyoid foramen; large splenial brown, with yellow on head brown; lateral ocular border yellow; yellow with brown dewlap yellow green with and tail banded Females with broad middorsal stripe bordered laterally by black; flanks yellow green to yellow with longitudinal very narrow dark blotches lips; brown spots; brown stripes similar to absent; posterior edge of parietal brown mark extending back from bordered by black; upper head part iris Distribution blue; limbs middorsal Anolis anchicayae stripe is found throughout the Pacific coastal lowlands of Colombia and Ecuador in primary and secondary humid forest (Fig were collected between and m imens Etymology The species 7) day by Many named is spec- on trunks for its type locality, the Anchicaya River valley present; ventral aspect of anteromedial process of coronoid extends posteriorly (n = 1); foramen is or slopes anteriorly (n of surangular surangular (n = 1) is 3); mandibles) or within (n is = anterior to (n coronoid beyond surangular foramen; jaw sculpturing absent; angular absent; angular process of articular present; teeth unicuspid anteriorly, tricuspid posteriorly; nine (n or 11 (n = 1) = 3) premaxillary teeth Four postxiphisternal ribs attached dornone free (4:0 rib formula); transverse processes on anterior caudal vertebrae grad- sally, 8, 9, 10A MCZ Holotype adult 80955, female, Ecuador, Pichincha, Finca Victoria, 37 SE of Santo Domingo, 16 July 1964, km F Vuilleumier mandibles) fossa; labial process of present; coronoid does not extend posterolaterally Figures within posterior suture of posterior border of dentary = entirely pronged; anteriormost aspect of mandibular Anolis lyra, new species 4) external opening or bordered by dentary and surangular (n = dentary = Paratypes Valle, MCZ 77458-60, Lower Rio Calima, Colombia, tributary of Rio San Juan, 1960, I Cabrera; MCZ 80954, Ecuador, Pichincha, Finca Victoria, 37 km SE of Santo Domingo, 16 July 1964, F Vuilleumier; MCZ 124407, Ecuador, Pichincha, Santo Domingo de los Colorados, August 1968, J Lynch; MCZ 145263, Ecuador, 15 km N Pichincha, Hotel Santo Domingo de November los Tinalandia, Colorados, 1964, A Kiester, E Williams; NEW SOUTH AMERICAN ANOLIS 2009 200 Kilometers Figure Map of coastal regions of Colombia and Ecuador showing collecting and Anolis sites of Anolis anchicayae (filled and personal observations of authors Nuqui, Choco Department, Colombia; Pueblo Rico, Risaralda Department, Colombia; Cafio Docordo between Cucurrupi and Noanama, San Juan River, Choco, Colombia; Bajo Calima, Buenaventura, Valle del Cauca Department, Colombia; Cisneros, Buenaventura, Valle del Cauca Department, Colombia; Bajo Anchicaya, Buenaventura, Valle del Cauca Department, Colombia; Cajambre river, Buenaventura, Valle del Cauca Department, Colombia; Malaga Bay, Buenaventura, Valle del Cauca Department, Colombia; Tangareal, Nariiio Department, Colombia; 10 Centro Cientifico Rio Palenque, Los Rios Province, Ecuador; 11 Finca La Victoria, southeast from Santo Domingo de los Colorados, Pichincha Province, Ecuador; 12 Puerto Quito, triangles) lyra (open squares) according to type series Pichincha Provice, Ecuador; 13 Hotel Tinalandia, approx 15 km N of Santo Domingo de los Colorados, Pichincha Province, Ecuador MCZ 145867, Ecuador, Los Rios, Centro Rio Palenque, 23 June 1974, K Cientifico Miyata; MCZ chincha, 146994-5, km N Buena R Webster; MCZ Ecuador, Fe, 31 July 1975, 152433-4, Ecuador, Pi- chincha, Hotel Tinalandia, 15 Domingo de K Miyata; los Pin- km N Santo Colorados, 27 April 1977, MCZ 164416, 165209-10, 171160, Ecuador, Pinchincha, Puerto Quito, April and September 1983, December 1984, MCZ 164420-1, Ecuador, PinchG Onore; incha, Santo Domingo de los Colorados, April 1983, G Onore Diagnosis Anolis lyra differs from all other South American Anolis except A vittigerus in possessing a small red dewlap with a dark central symbol spot in males and a lyre-shaped in the occipital region (Fig 8) Anolis lyra differs from A vittigerus in possessing smaller middorsal scales (mean number of 5% SVL for A lyra = = 5.8; P = 0.006, Mann- longitudinal scales in 8.5, A vittigerus Whitney U test), more scales between supra(mean = 2.3 in A lyra, 0.6 P = 0.001, Mann-Whitney U orbital semicircles in A vittigerus; and more scales between interparietal and supraorbital semicircles (x = 4.1 in A test), lyra, 2.4 in A vittigerus; Whitney P = 0.001, Mann- f/test; Fig 10) Also, A lyra tends to display much weaker lateral striping (i.e., broken lines only extending to the axillae versus broad stripes to groin in A vittigerus) and a more prominent lyre-shaped posterior head marking (Fig 8; absent to weak in A vittigerus) In western Colombia, the only area in which both species can be expected to BREVIORA 10 Figure Anolis lyra new species, male, figure Anolis lyra new species, female, from km N from No 516 of Boca Ana, Valle, Colombia Photo by Steven Poe km N of Boca Ana, Valle, Colombia Photo by Steven Poe NEW SOUTH AMERICAN ANOLIS 2009 Figure vittigerus, 10 Head scales LACM 42200, approximately 250 km of (A) Anolis lyra, MCZ 77459, Lower Rio Calima, 11 Valle, Colombia, and (B) Anolis upper Rio Napipi, 45 minutes by canoe below mouth of Rio Merendo These apart in western Colombia Note larger scales (e.g., around localities are interparietal) in A vittigerus BREVIORA 12 and A vittigerus may be most by the condition of the supraorbital semicircles in broad contact in A vittigerus; usually separated by 2-3 occur, A lyra easily separated — External Description of Holotype (paratype variation in parentheses; description based on adults good in is MCZ condition: 80955, 77458-9, 145267, 145867, 146994-5, 164420-1) 164416, Snout-vent head length 17.8 (14.8-19.6; 0.23-0.26 SVL), width 11.8 (10.1-12.8; 0.16-0.17 SVL); ear height 1.9 (1.2-2.1; 0.02-0.03 SVL); femoral length 22.8 (19.2-24.2; 0.29-0.32 SVL); tail length approximately 165 (128-165; 2.21-2.39 SVL) length 76.7 (60.7-75.2); Dorsal head scales keeled; frontal depression present; rostral slightly overlaps mental scales across snout anteriorly; nine (7-11) between second canthals; supraorbital semicircles oculars separated by two (1^1) scales; sub- with contact in scales keeled; zero (0-20) enlarged middorsal 5% rows, (8-9) longitudinal rows in SVL; ventral ly keeled, of scales in diagonal rows, strong- 5% five (5-7) scales in of SVL; supradigitals multicarinate; toepads expand- scales in A lyra 152433^, No 516 supralabials (or row of scales); two elongate separated by one supraciliary scales (or 1-2 slightly elongate expanded lamellae under third and fourth phalanges of fourth toe (second and third phalanges of Williams [e.g., Wil- ed; 18 (18-21) liams et al, 1995]); fourth toe length 15.6 (14.9-15.6; 0.21-0.23 SVL); scales of anterior thigh large, keeled, overlapping, becoming much smaller posteriorly; row of middorsal tail with a single scales Skeletal Description (Based on ton MCZ 77460) Parietal roof shaped parietal crests, Dry flat, Skele- with Y- with no casquing, lacking crenulation on edges, with anterolateral corners flush with posterolateral edges of frontal; pineal foramen at parietal-frontal suture; dorsal skull tal bones smooth; postfron- present as sliver at rim of orbit; prefrontal separated from nasal by anterior extension of frontal or in contact with nasal; frontal sutures anteriorly with nasals; parallel crests slightly on nasals absent; external nares bordered enlarged scales; eight (6-10) loreal rows; two posteriorly by nasals; dorsal aspect of jugal and sulcus between terminates on lateral surface of postorbital; by scales) followed scales separate parallel naris rows of interparietal posterodorsal ramus of squamosal smaller length 2.0 (1.7-2.3); three (3-5) scales sepa- than posteroventral ramus, separated from rostral and supralabial; first rate interparietal and supraorbital semicir- cles; preoccipital absent; seven (6-8) supra- labials center to of eye; (6-10) six postmentals; eight (6-9) postrostrals; some enlarged scales present in supraocular disc, parietal by supratemporal; posterior aspect of jugal mostly straight; epipterygoid contacts parietal dorsally; pterygoid tine teeth absent; lateral smooth, without posteriorly directed decreasing gradually in processes; ally yond ectopterygoid on size, bordered mediby a complete (or partial) row of small scales; mental partially divided posteriorly, extending posterolaterally with transverse (i.e., beyond rostral, not concave) posterior and pala- edge of vomer maxilla extends posteriorly ventral is lateral be- surface of between basipterygoid processes absent; lateral shelf of quadrate absent; black pigment is present skull; crest of basisphenoid border; no (0-1) enlarged sublabials; dewlap on just reaches axillae in males, nearly reaches slightly overlap premaxilla dorsally; posteri- axillae in females; single scales; dewlap weak scales in axillary rows of pocket (not parietal, prefrontals, and nasals; nasals ormost mandibular tooth is posterior to anterior mylohyoid foramen; splenial appar- tubelike); enlarged postcloacal scales absent; ently absent (difficult to nuchal and dorsal crests absent; some dorsal of anteromedial process of coronoid extends tell); ventral aspect NEW SOUTH AMERICAN ANOLIS 2009 were 13 posteriorly; external opening of surangular lyra foramen secondary humid forest on twigs or leaves entirely is surangular within shares border between dentary lar; posterior suture of dentary or and suranguis blunt, not 1-5 labial process does is of coronoid present; coronoid extend not within mandibular fossa; posterolaterally high by day pronged; anteriormost aspect of posterior border of dentary m collected in Some specimens were teeth unicuspid eight posteriorly; tricuspid Etymology This species premaxillary back extends from area posterior to eye, becoming broken between by dark brown; limbs, bordered ventrally light for the its head DISCUSSION yellow extends posteriorly from snout Many Relationships few as ters occipital region (lyre "opens" anteriorly); dark band between orbits dorsally; laterally, stripe named (Fig 8) anteriorly, Color in Life (Adapted from Field Notes and Color Photos by Poe) Male basically dark brown with pale yellow pattern; middorsum with four light blotches breaking up dark background; butterfly-shaped marking above pelvis; posterior of dorsal head light gray with dark lyre-shaped marking around light is beyond tooth positions a 3% (i.e., taxa are scored for as of parsimony informative charac- only external morphological data) Inclusion of such poorly scored taxa will tend to increase the number of most parsi- monious trees and thus lower support levels such as bootstrap frequencies (Huelsenbeck, 1991) We nevertheless include these taxa to an achieve estimate of relative to A anchicayae their and A phylogeny lyra and ment of A anchicayae and A lyra (Gauthier et al, 1988; Hendy and Penny, 1989) Because support values are uniformly low and many optimal trees resulted, as expected, we summarize results with a majority-rule consensus of optimal brown band, broken by dark blotches below orbit; limbs and tail banded with curved anchicayae and A lyra are shown in Fig lines; dorsal markings less distinct, melding together in some specimens (perhaps in response to stress); venter pale yellow dark flecks most evident laterally; with underside to increase the accuracy of phylogenetic place- along dorsal edge of mouth below dark yellow in collected lyre-shaped marking on the back of angular absent; angular process of articular large; night at lower parts of trees and on ground surangular foramen; jaw sculpturing absent; present, sleeping trees Results of the phylogenetic analysis of A anchicayae Anolis peraccae is among an sister species to unresolved group 1 A of South American alpha Anolis (approximateequivalent to Etheridge's [1959] latifrons ly tongue tan, throat light gray; eyes reddish brown; dewlap skin red with orange tint, and Guyer and Savage's [1986] "DacThe "latifrons" group usually has been found to be monophyletic (including " Phenacosaurus' Etheridge, 1959; Jackman with discrete dark central spot; dewlap scales et al., 1999; Poe, 2004, fig 2; of head pale yellow washed with red, with some dark reticulations anterior to orbits; light distally, dark centrally Female coloration male but with like spot Distribution Anolis lyra Ecuador '; ' found from the Colombia south Most A (Fig 7) 2005) but (Poe, 2004, fig Nicholson et occasionally paraphyletic is 5; studies Obviously, is is Pacific lowlands of central to northwestern tyloa") al., bluish-gray dewlap skin with faint red tint and dark central series needed on the this study) in recent more phylogenetic work South American alpha Anolis Anolis lyra is grouped with some other lemurinus-group species A bicaorum and A BREVIORA 14 No 516 fortunensis B trachyderma tolimensis other Anolis medemi aeneus roquet j— maculiventris granuliceps — poecilopus richardi bonairensis tropidogaster extremus r— lionotus -P— oxylophus griseus irinitatus macrolepis luciae i— concolor esperanzai wiliamsmittermeierorum heterodermus proboscis inderenae P— pinchoti ~\ limifrons — zeus J- 92c— bicaorum -P— nicefori solitarius lyra