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Full text of "Breviora"

B RE VI O R A 

Nl m s e Til inn ©i Comparative /^oology 



US ISSN 0006-9698 



Cambridge, Mass. 1 June 2009 Number 516 

DESCRIPTIONS OF TWO NOMEN NUDUM SPECIES OF ANOLIS LIZARD FROM 
NORTHWESTERN SOUTH AMERICA 

Steven Poe, 1 Julian Velasco, 2 Kenneth Miyata, 3 - 4 and Ernest E. Williams 3 ' 4 

Abstract. We 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 is similar to Anolis vittigerus but 
differs primarily in possessing smaller dorsal scales on the head and body and in color pattern. Anolis anchicayae new 
species is similar to Anolis peraccae and Anolis fasciatus but differs primarily in size 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"). 

Key words: 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 con nombres en nomen nundum. Anolis lyra sp. nov. es parecido a Anolis 
vittigerus pero se diferencia principalmente en que presenta las escamas del dorso de la cabeza y el cuerpo mas 
pequenas, y en el patron de coloracion. Anolis anchicayae sp. nov. es parecido a Anolis peraccae y Anolis fasciatus 
pero 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 con 25 1 especies adicionales de Anolis usando 
una combination de datos morfologicos y moleculares. Anolis anchicayae es taxon hermano de A. peraccae dentro de 
un grupo basal de especies suramericanas de Anolis "alfa." Anolis lyra esta relacionado con otras especies del grupo 
lemurinus dentro de un clado mas grande compuesto por especies continentales de Anolis "beta." 



1 Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, New 
Mexico 87131, U.S.A.; e-mail: anolis@unm.edu 

2 Grupo de Ecologia Animal, Departamento de Biologia, Universidad del Valle, A. A. 25360, Cali, Colombia 

3 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. 

4 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 6. 
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 



No. 516 



Ernest Williams and collaborators pub- 
lished a series of papers from 1983 to 1992 
describing "new or problematic" Anolis from 
Colombia. These papers reviewed and ex- 
panded the Colombian anoline fauna and 
established the country as a center of 
diversity for the Anolis clade. The publica- 
tion of these descriptions and other taxo- 
nomic works on Colombian lizards (e.g., 
Ayala and Castro, 1983) prompted the need 
for a summary of available information. 
Stephen Ayala (1986) obliged and produced 
a list of the 205 lizard species thought to 
occur in Colombia. This list included eight 
species of Anolis (and species in other clades) 
listed in quotes ("") and associated with 
authors to signal that descriptions were not 
yet published but were forthcoming. Pub- 
lished descriptions of Anolis species associated 
with four of these names soon followed (A. 
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 descrip- 
tions 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. 

MATERIALS AND METHODS 

We consider species to be evolutionary 
lineages (Simpson, 1961; Wiley, 1978) and 
operationalize this concept by identifying 
species on the basis of consistent differences 
between populations. That is, we hypothesize 
that populations that are diagnosable by 
major differences in the frequencies of traits 
are distinct evolutionary lineages or species 
(see 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 



the anterior of the cloaca. Head length was 
measured from the tip of the snout to the 
anterior edge of the ear. Femoral length was 
measured from the midline of the venter to 
the knee, with the limb bent at a 90-degree 
angle. Head width was measured at the 
broadest part of the head, between the 
posterolateral corners of the orbits. Com- 
parisons were made with preserved material 
of the putative closest relatives of the new 
species, and with published species descrip- 
tions, preserved material, or both of more 
distant relatives (online Appendix 1). Scale 
terminology and characters used mainly 
follow standards established by Williams 
(e.g., Williams et al, 1996) for species 
descriptions of anoline lizards. Skeletal 
description is given in terms of Poe's (1998, 
2004) and Etheridge's (1959) characters. See 
those papers for more detailed descriptions 
of skeletal conditions and alternative condi- 
tions in Anolis. 

We scored the new species for the mor- 
phological phylogenetic characters of Poe 
(2004; Appendix 1) and performed a parsi- 
mony analysis of these taxa and 251 other 
species of Anolis and eight outgroups with 
the use of 1,291 parsimony-informative 
characters from published sources and some 
of our additional unpublished morphological 
data. This dataset includes 91 morphological 
characters (Poe, 2004), 998 sites of mtDNA 
(NADH dehydrogenase subunit 2, five 
transfer RNAs; Nicholson et al. , 2005), and 
182 sites of nuclear DNA (internal tran- 
scribed tracer region; Nicholson, 2002). 
Codes for morphological data for A. anchi- 
cayae and A. lyra are in online Table 1. We 
used PAUP* version 4.0b 10 (Swofford, 
2002) to perform a heuristic search with 
100 random additions of taxa and tree- 
bisection-reconnection branch swapping. 
We also used PAUPrat (Sikes and Lewis, 
2001) to perform several runs of the parsi- 
mony ratchet (Nixon, 1999) to find optimal 



2009 



NEW SOUTH AMERICAN ANOLIS 




Figure 1. Anolis anchicayae new species, male. 
Photo by Julian Velasco. 



trees (20%-35% of characters reweighted, 
150-200 replicates per run). 

DESCRIPTION OF NEW SPECIES 

Anolis anchicayae, new species 

Figures 1-5 

Holotype. MCZ 160234, adult male, Co- 
lombia, Valle, San Isidro, 5 August 1977, 
Helen Chin. 

Paratypes. MCZ 112426-7, Colombia, 
Choco, Rio San Juan, Carlo Docordo 
between Cucurrupi and Noanama, 24 De- 
cember 1968, B. Malkin; MCZ 158385-7, 



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 159767- 
8, Colombia, Valle, Anchicaya Hydroelectric 
Central, 3 June 1976, J. Castillo, F. Castro, 
H. Chin; MCZ 159997-8, 160234-9, Colom- 
bia, Valle, San Isidro, Carton, 5 and 9 
August 1977, H. Chin; MCZ 160008-12, 
Colombia, Valle, Granje Bajo Central, 1976- 
77, 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, 7 December 

1978, H. Chin, F. Castro; ICN 6115, 
Colombia, Choco, Bahia Solano, near 
Jayita, J. Renjifo. 

Diagnosis. Anolis anchicayae is similar to 
A. peraccae and A. fasciatus. These three 
species share keeled dorsal head scales, a 
double row of middorsal caudal scales, 







**. 



fSsk 



Figure 2. Anolis anchicayae new species, MCZ 145362. Tail is missing in this specimen. 



BREVIORA 



No. 516 




Figure 3. Anolis anchicayae, MCZ 145362, dorsal head scales. 



absence of dewlap in females, absence of supralabial. Anolis anchicayae is most easily 

caudal transverse vertebral processes poste- distinguished from A. peraccae by larger size 

riorly, and an elongate anterior nasal scale (to 54 mm SVL in A. peraccae; to 63 mm in 

that reaches the rostral but does not contact A. anchicayae; Fig. 6) and color pattern {A. 

the sulcus between the rostral and first anchicayae is predominantly yellow-green on 



2009 



NEW SOUTH AMERICAN ANOLIS 




Figure 4. Anolis anchicayae, MCZ 145362, lateral head scales. 




Figure 5. Anolis anchicayae, MCZ 145362, ventral 
head scales. 



its flanks with an elongate ocellus from eye 
to shoulder 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 anchi- 
cayae 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 
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 (= "Nor- 
ops"; with which it is unlikely to be confused) 
by the absence of transverse processes on 
posterior caudal vertebrae (present in beta 
Anolis); from phenacosaur and tigrinus- 
group Anolis {A. heterodermus, A. inderenae, 
A. nicefori, A. bellipeniculus, A. carlostoddi, 
A. euskalerriari, A. neblininus, A. orcesi, A. 
tetarii, A. vanzolinii, A. ruizi, A. solitarius, A. 
menta, A. nasofrontal, A. pseudotigrinus, A. 



BREVIORA 



No. 516 



MALES 



FEMALES 





SVL 42mm 44 46 48 50 52 54 56 58 60 



SVL 42mm 44 46 48 50 52 54 56 58 



30 
20 

10 



N = 30 



Anolis peraccae 
Norino; Colombia 




50 












40 


N»I8 






Anolis per 
Plchincha; 


30 










20 










10 






% 




K%4l%!*'&3kS£ 


^•i.!':.:.:.:.**:'*':'*.* 


. . 



N = 28 



SVL 42mm 44 46 48 50 52 54 56 58 60 



SVL 42mm 44 46 48 50 52 54 56 



58 



Figure 6. Distribution of body sizes within populations for samples of A. peraccae and A. anchicayae, showing 
larger size of A. anchicayae. The specimens used to construct this table are unknown because they were not listed in 
Williams' notes. 



tigrinus, A. menta, A. lamari, A. vaupesianus, 
A. jacare, A. umbrivagus, A. paravertebralis, A. 
williamsmitteremeierorum) by possessing 
strongly keeled head scales superior to the 
orbits (mostly smooth or pustulose in 
phenacosaur and tigrinus-group Anolis); 
from latifrons- and some aequatorialis-group 
Anolis (A. latifrons, A. frenatus, A. fraseri, 
A. danieli, A. apollinaris, A. casildae, A. 
squamulatus, A. princeps, A. purpurescens, 
A. agassizi, A. eulaemus, A. maculigula) by 
much smaller size (> 100 mm SVL in these 
species); from some punctatus-group Anolis 
(A. cuscoensis, A. soinii, A. huilae, A. 
transversalis, A. boettgeri, A. deltae, A. 
dissimilis, A. sanlamartae) by displaying 



scales separating the supraorbital semicircles 
(supraorbital semicircles in contact in these 
species); from other punctatus- and aequa- 
torialis-group Anolis (A. chocorum, A. cali- 
mae, A. antioquiae, A. fitchi, A. aequator- 
ialis, A. megalopithecus) in the lack of a 
dewlap in females (female dewlap present in 
these species); from A. punctatus and the 
proboscis Anolis (A. proboscis, A. laevis, A. 
phyllorhinus) by the absence of swelling on 
the snout of the male (swelling present in 
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 



2009 



NEW SOUTH AMERICAN ANOLIS 



pattern (solid bright green in A. Moris; 
banded pattern of browns and greens in A. 
anchicayae); from A. festae and A. nigroli- 
neatus (the name is a probable synonym of 
A. festae; Williams, 1982) in male dewlap 
pattern (dark elongate blotch in these 
species; absent in A. anchicayae); from A. 
caquetae in absence of contact between 
interparietal and supraorbital semicircles; 
from A. propinqus in number of subdigital 
lamellae (27 in A. propinqus, 16-20 in A. 
anchicayae); from A. gemmosus in the 
condition of the nasal scale (elongate 
anterior nasal contacts sulcus between 
rostral and first supralabial in A. gemmosus; 
is separated from sulcus by small scale 
inferior to naris in A. anchicayae). 

External Description of Holotype (paratype 
variation in parentheses; description is based 
on adults in good condition: MCZ 160234, 
158386-7, 159767, 159997-8, 160009-10, 
160230, 160232-3, 160238). Snout-vent 
length 52.0 (50.0-53.6); head length 12.0 
(11.8-13.2; 0.23-0.25 SVL), width 7.1 (7.0- 
7.8; 0.14-0.15 SVL); ear height 1.3 (1.1-1.6; 
0.02-0.03 SVL); femoral length 12.9 (13.6- 
16.0; 0.25-0.30 SVL); tail length 115 (102- 
116; 1.96-2.26 SVL); fourth toe length 10.5 
(10.7-11.9; 0.21-0.22 SVL), width 1.0 (1.0- 
1.3; 0.02 SVL). 

Dorsal head scales uni- to multicarinate 
above orbits, mostly smooth in frontal area, 
unicarinate on snout; frontal depression 
present; rostral slightly overlaps mental 
anteriorly; nine (8-12) scales across snout 
between second canthals; supraorbital semi- 
circles separated by two (1-2) scales; sub- 
oculars in contact with supralabials; one (1-2) 
elongate supraciliary scale followed by small 
undifferentiated scales; seven (5-7) loreal 
rows; elongate anterior nasal scale contacts 
rostral, is separated from sulcus between 
rostral and first supralabial by one small 
scale inferior to naris; supraorbital semicir- 
cles evident but not strongly differentiated, 



especially posteriorly; interparietal length 1.3 
(1.2-1.6); three (3-4) scales separate interpa- 
rietal and supraorbital semicircles; preoccip- 
ital absent; six (5-6) supralabials to center of 
eye; nine (8-10) postrostrals; six (6-9) post- 
mentals; some slightly enlarged scales present 
in supraocular disc, decreasing gradually in 
size, bordered medially by a partial row of 
small scales; mental partially divided poste- 
riorly, extending posterolaterally beyond 
rostral, with posterior border in concave 
arc; two (2-A) enlarged sublabials in contact 
with infralabials (or posteriorly separated by 
small scales); dewlap reaches well posterior 
to axillae in males, absent in females; scales 
on dewlap in rows of approximately three 
scales with some scattered scales; no axillary 
pocket; pair of enlarged postcloacal scales in 
contact (or separated by small scales; absent 
in females); nuchal and dorsal crests absent; 
dorsal scales keeled; approximately 10 (5-12) 
enlarged middorsal rows, six (6-8) longitu- 
dinal rows in 5% 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, overlap- 
ping, becoming smaller and nonoverlapping 
posteriorly; supradigitals multicarinate; toe- 
pads expanded; 19 (16-20) expanded lameal- 
lae 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 Dry Skele- 
tons MCZ 112426-7, 159767, 160012, 
ICN 6115). Parietal roof flat, with trapezoi- 
dal crests, with a very narrow posterior 
border, with no casquing, lacking crenula- 
tion on edges, not extending posteriorly over 
supraoccipital, with anterolateral corners 
flush with posterolateral edges of frontal; 
pineal foramen at parietal-frontal suture (n 
= 4) or in parietal (n = 1); dorsal skull bones 
smooth; postfrontal present; prefrontal sep- 
arated from nasal by anterior extension of 



BREVIORA 



No. 516 



frontal (n = 2) or in contact with nasal (n = 
2); frontal sutures anteriorly with nasals; 
parallel crests on nasals absent; external 
nares bordered posteriorly by nasals; dorsal 
aspect of jugal terminates on lateral surface 
of postorbital; jugal does not contact squa- 
mosal; posterodorsal ramus of squamosal 
shorter than posteroventral ramus, separated 
from parietal by supratemporal; posterior 
aspect of jugal mostly straight; epipterygoid 
contacts parietal dorsally; pterygoid and 
palatine teeth absent; lateral edge of vomer 
is smooth, without posteriorly directed lat- 
eral processes; maxilla extends posteriorly 
beyond ectopterygoid on ventral surface of 
skull (n = 3) or reaches ectopterygoid (n = 
2); crest between basipterygoid processes of 
basisphenoid absent; lateral shelf of quadrate 
absent; no black pigment on surface of skull; 
nasals do not overlap premaxilla dorsally; 
posterior edge of parietal is approximately 
even with vertical level of parietal-frontal 
suture; posteriormost mandibular tooth is 
partially anterior (n = 3 mandibles) or 
partially posterior (n = 7 mandibles) to 
anterior mylohyoid foramen; large splenial 
present; ventral aspect of anteromedial pro- 
cess of coronoid extends posteriorly (n = 4) 
or slopes anteriorly (n = 1); external opening 
of surangular foramen is entirely within 
surangular (n = 1) or bordered by dentary 
and surangular (n = 3); posterior suture of 
dentary is pronged; anteriormost aspect of 
posterior border of dentary is anterior to (n 
= 3 mandibles) or within (n = 7 mandibles) 
mandibular fossa; labial process of coronoid 
present; coronoid does not extend postero- 
laterally beyond surangular foramen; jaw 
sculpturing absent; angular absent; angular 
process of articular present; teeth unicuspid 
anteriorly, tricuspid posteriorly; nine (n = 3) 
or 11 (n = 1) premaxillary teeth. 

Four postxiphisternal ribs attached dor- 
sally, none free (4:0 rib formula); transverse 
processes on anterior caudal vertebrae grad- 



ually lost posteriorly (Etheridge's [1959] 
"alpha" condition). 

Color in Life (Adapted from Field Notes 
and Color Photos by Velasco). Males brown 
middorsally interrupted by approximately 10 
dark blotches that extend laterally; flanks 
greenish-yellow with narrow diagonal rows 
of partially connected small brown spots; 
elongate brown mark extending back from 
eye, bordered by black; upper head part 
yellow with brown, lateral head brown; 
ocular border yellow; yellow with brown 
blotches on lips; dewlap yellow green with 
brown spots; iris blue; limbs and tail banded. 
Females with broad middorsal stripe bor- 
dered laterally by black; flanks yellow green 
to yellow with longitudinal very narrow dark 
brown stripes similar to middorsal stripe. 

Distribution. Anolis anchicayae is found 
throughout the Pacific coastal lowlands of 
Colombia and Ecuador in primary and 
secondary humid forest (Fig. 7). Many spec- 
imens were collected by day on trunks 
between 2 and 5 m. 

Etymology. The species is named for its 
type locality, the Anchicaya River valley. 

Anolis lyra, new species 

Figures 8, 9, 10A 

Holotype. MCZ 80955, adult female, 
Ecuador, Pichincha, Finca Victoria, 37 km 
SE of Santo Domingo, 16 July 1964, F. 
Vuilleumier. 

Paratypes. MCZ 77458-60, Colombia, 
Valle, Lower Rio Calima, 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, Pi- 
chincha, Santo Domingo de los Colorados, 
2 August 1968, J. Lynch; MCZ 145263, 
Ecuador, Pichincha, Hotel Tinalandia, 
15 km N Santo Domingo de los Colorados, 
6 November 1964, A. Kiester, E. Williams; 



2009 



NEW SOUTH AMERICAN ANOLIS 




200 Kilometers 



Figure 7. Map of coastal regions of Colombia and Ecuador showing collecting sites of Anolis anchicayae (filled 
triangles) and Anolis lyra (open squares) according to type series and personal observations of authors. 1. Nuqui, 
Choco Department, Colombia; 2. Pueblo Rico, Risaralda Department, Colombia; 3. Cafio Docordo between 
Cucurrupi and Noanama, San Juan River, Choco, Colombia; 4. Bajo Calima, Buenaventura, Valle del Cauca 
Department, Colombia; 5. Cisneros, Buenaventura, Valle del Cauca Department, Colombia; 6. Bajo Anchicaya, 
Buenaventura, Valle del Cauca Department, Colombia; 7. Cajambre river, Buenaventura, Valle del Cauca 
Department, Colombia; 8. Malaga Bay, Buenaventura, Valle del Cauca Department, Colombia; 9. 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, 
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 
Cientifico Rio Palenque, 23 June 1974, K. 
Miyata; MCZ 146994-5, Ecuador, Pin- 
chincha, 1 km N Buena Fe, 31 July 1975, 
R. Webster; MCZ 152433-4, Ecuador, Pi- 
chincha, Hotel Tinalandia, 15 km N Santo 
Domingo de los Colorados, 27 April 1977, 
K. Miyata; MCZ 164416, 165209-10, 
171160, Ecuador, Pinchincha, Puerto Quito, 
April and September 1983, December 1984, 
G. Onore; MCZ 164420-1, Ecuador, Pinch- 
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 spot in males and a lyre-shaped 
symbol in the occipital region (Fig. 8). Anolis 



lyra differs from A. vittigerus in possessing 
smaller middorsal scales (mean number of 
longitudinal scales in 5% SVL for A. lyra = 
8.5, A. vittigerus = 5.8; P = 0.006, Mann- 
Whitney U test), more scales between supra- 
orbital semicircles (mean = 2.3 in A. lyra, 0.6 
in A. vittigerus; P = 0.001, Mann-Whitney U 
test), and more scales between interparietal 
and supraorbital semicircles (x = 4.1 in A. 
lyra, 2.4 in A. vittigerus; P = 0.001, Mann- 
Whitney 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 



10 



BREVIORA 



No. 516 




Figure 8. Anolis lyra new species, male, from 2 km N of Boca Ana, Valle, Colombia. Photo by Steven Poe. 




figure 9. Anolis lyra new species, female, from 2 km N of Boca Ana, Valle, Colombia. Photo by Steven Poe. 



2009 



NEW SOUTH AMERICAN ANOLIS 



11 











Figure 10. Head scales of (A) Anolis lyra, MCZ 77459, Lower Rio Calima, Valle, Colombia, and (B) Anolis 
vittigerus, LACM 42200, upper Rio Napipi, 45 minutes by canoe below mouth of Rio Merendo. These localities are 
approximately 250 km apart in western Colombia. Note larger scales (e.g., around interparietal) in A. vittigerus. 



12 



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No. 516 



occur, A. lyra and A. vittigerus may be most 
easily separated by the condition of the 
supraorbital semicircles — in broad contact 
in A. vittigerus; usually separated by 2-3 
scales in A. lyra. 

External Description of Holotype (para- 
type variation in parentheses; description is 
based on adults in good condition: MCZ 
80955, 77458-9, 145267, 145867, 146994-5, 
152433^, 164416, 164420-1). Snout-vent 
length 76.7 (60.7-75.2); 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 ap- 
proximately 165 (128-165; 2.21-2.39 SVL). 

Dorsal head scales keeled; frontal depres- 
sion present; rostral slightly overlaps mental 
anteriorly; nine (7-11) scales across snout 
between second canthals; supraorbital semi- 
circles separated by two (1^1) scales; sub- 
oculars in contact with supralabials (or 
separated by one row of scales); two elongate 
supraciliary scales (or 1-2 slightly elongate 
scales) followed by parallel rows of slightly 
enlarged scales; eight (6-10) loreal rows; two 
scales separate naris and sulcus between 
rostral and first supralabial; interparietal 
length 2.0 (1.7-2.3); three (3-5) scales sepa- 
rate interparietal and supraorbital semicir- 
cles; preoccipital absent; seven (6-8) supra- 
labials to center of eye; six (6-10) 
postmentals; eight (6-9) postrostrals; some 
enlarged scales present in supraocular disc, 
decreasing gradually in size, bordered medi- 
ally by a complete (or partial) row of small 
scales; mental partially divided posteriorly, 
extending posterolaterally beyond rostral, 
with transverse (i.e., not concave) posterior 
border; no (0-1) enlarged sublabials; dewlap 
just reaches axillae in males, nearly reaches 
axillae in females; dewlap scales in rows of 
single scales; weak axillary pocket (not 
tubelike); enlarged postcloacal scales absent; 
nuchal and dorsal crests absent; some dorsal 



scales keeled; zero (0-20) enlarged middorsal 
rows, 9 (8-9) longitudinal rows in 5% of 
SVL; ventral scales in diagonal rows, strong- 
ly keeled, five (5-7) scales in 5% of SVL; 
supradigitals multicarinate; toepads expand- 
ed; 18 (18-21) expanded lamellae under third 
and fourth phalanges of fourth toe (second 
and third phalanges of Williams [e.g., Wil- 
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; tail with a single 
row of middorsal scales. 

Skeletal Description (Based on Dry Skele- 
ton MCZ 77460). Parietal roof flat, with Y- 
shaped parietal crests, with no casquing, 
lacking crenulation on edges, with anterolat- 
eral corners flush with posterolateral edges 
of frontal; pineal foramen at parietal-frontal 
suture; dorsal skull bones smooth; postfron- 
tal 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 
on nasals absent; external nares bordered 
posteriorly by nasals; dorsal aspect of jugal 
terminates on lateral surface of postorbital; 
posterodorsal ramus of squamosal smaller 
than posteroventral ramus, separated from 
parietal by supratemporal; posterior aspect 
of jugal mostly straight; epipterygoid con- 
tacts parietal dorsally; pterygoid and pala- 
tine teeth absent; lateral edge of vomer is 
smooth, without posteriorly directed lateral 
processes; maxilla extends posteriorly be- 
yond ectopterygoid on ventral surface of 
skull; crest between basipterygoid processes 
of basisphenoid absent; lateral shelf of 
quadrate absent; black pigment is present 
on parietal, prefrontals, and nasals; nasals 
slightly overlap premaxilla dorsally; posteri- 
ormost mandibular tooth is posterior to 
anterior mylohyoid foramen; splenial appar- 
ently absent (difficult to tell); ventral aspect 
of anteromedial process of coronoid extends 



2009 



NEW SOUTH AMERICAN ANOLIS 



13 



posteriorly; external opening of surangular 
foramen is entirely within surangular or 
shares border between dentary and surangu- 
lar; posterior suture of dentary is blunt, not 
pronged; anteriormost aspect of posterior 
border of dentary is within mandibular fossa; 
labial process of coronoid present; coronoid 
does not extend posterolaterally beyond 
surangular foramen; jaw sculpturing absent; 
angular absent; angular process of articular 
present, large; teeth unicuspid anteriorly, 
tricuspid posteriorly; eight premaxillary 
tooth positions. 

Color in Life (Adapted from Field Notes 
and Color Photos by Poe). Male basically 
dark brown with pale yellow pattern; mid- 
dorsum 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 
occipital region (lyre "opens" anteriorly); 
dark band between orbits dorsally; laterally, 
a light stripe extends back from area 
posterior to eye, becoming broken between 
limbs, bordered ventrally by dark brown; 
light yellow extends posteriorly from snout 
along dorsal edge of mouth below dark 
brown band, broken by dark blotches below 
orbit; limbs and tail banded with curved 
yellow lines; dorsal markings less distinct, 
melding together in some specimens (perhaps 
in response to stress); venter pale yellow with 
dark flecks most evident laterally; underside 
of head pale yellow washed with red, with 
some dark reticulations anterior to orbits; 
tongue tan, throat light gray; eyes reddish 
brown; dewlap skin red with orange tint, 
with discrete dark central spot; dewlap scales 
light distally, dark centrally. 

Female coloration like male but with 
bluish-gray dewlap skin with faint red tint 
and dark central spot. 

Distribution. Anolis lyra is found from the 
Pacific lowlands of central Colombia south 
to northwestern Ecuador (Fig. 7). Most A. 



lyra were collected sleeping at night in 
secondary humid forest on twigs or leaves 
1-5 m high. Some specimens were collected 
by day in lower parts of trees and on ground. 
Etymology. This species is named for the 
lyre-shaped marking on the back of its head 
(Fig. 8). 

DISCUSSION 

Relationships. Many taxa are scored for as 
few as 3% of parsimony informative charac- 
ters (i.e., 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 
achieve an estimate of their phylogeny 
relative to A. anchicayae and A. lyra and to 
increase the accuracy of phylogenetic place- 
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 trees. 

Results of the phylogenetic analysis of A. 
anchicayae and A. lyra are shown in Fig. 1 1 . 
Anolis anchicayae is sister species to A. 
peraccae among an unresolved group of 
South American alpha Anolis (approximate- 
ly equivalent to Etheridge's [1959] latifrons 
series and Guyer and Savage's [1986] "Dac- 
tyloa"). The "latifrons" group usually has 
been found to be monophyletic (including 
" ' Phenacosaurus' '; Etheridge, 1959; Jackman 
et al., 1999; Poe, 2004, fig. 2; Nicholson et 
al., 2005) but is occasionally paraphyletic 
(Poe, 2004, fig. 5; this study) in recent 
studies. Obviously, more phylogenetic work 
is needed on the South American alpha 
Anolis. 

Anolis lyra is grouped with some other 
lemurinus-group species A. bicaorum and A. 



14 



BREVIORA 



No. 516 



B 




other Anolis 

aeneus 

roquet 

richardi 

bonairensis 

extremus 

griseus 

irinitatus 

luciae 

esperanzai 

wiliamsmittermeierorum 

heterodermus 

proboscis 

inderenae 

nicefori 

solitarius 

ruizi 

aequatorialis 

casildae 

fitchi 

fraseri 

jacare 

microtus 

insignis 

squamulatus 

frenatus 

latifrons 

ventrimaculatus 

apollinaris 

agas$izi 

cnlons 

festae 

punctatus 

cuscoensis 

nuilae 

soinii 

transversalis 

anchicayae 

peraccae 

fasciatus 

gemmosus 

boettgeri 

outg roups 




fortunensis 
trachyderma 
tolimensis 
medemi 
maculiventris 
j— granuliceps 
1 — poecilopus 

tropidogaster 
r— lionotus 
-P— oxylophus 

1 macrolepis 

i— concolor 
P— pinchoti 

~\ limifrons 

J- — zeus 
92c— bicaorum ^ 
-P— lyra < — 

1 lemurinus 

r— carpenteri 

■— ocelloscapularis 
humilis 



92 



" I— magnaphallu 
I— P— pachypus 

1 tropidolepis 

i— capita 
— P — cuprinus 

1 tropidonotus 

r— cobanensis 
■— P— parvicirculatus 

— ' polylepis 

1 cupreus 

dollfusianus 

i— altae 
,—P— kemptoni 

— ' townsendi 

' fuscoauratus 

antonii 

mariarum 

r— isthmicus 
—H— microlepidotus 

J ' sericeus 

I r— intermedius 

L - laeviventris 
r— pentaprion 
vociferans 
ortoni 

megapholidotus 
utilensis 
subocularis 
nebuloides 
quercorum 
nebulosus 
j— crassulus 
1 — sminthus 
— liogaster 
i— dunni 
-P— gadovi 

1 laylori 

Q 2r _r— vittigerus 
ooi-p- woodi 

v±\ ' aguaticus 

92 I r— biporcatus 

r- i— petersi 

I I r— gracilipes 

' — uniformis 

r— bitectus 

■— polyrhachis 
_r— loveridgei 
•— purpurgularis 
i— nitens chrysolepis 
'— n/tens scypheus 
meridionalis 
auratus 
j— annectens 
•— onca 
bar/cer/ 
//'neafL/s 

Figure 1 1. Relationships of (A) Anolis anchicayae and (B) /I. /rra based on parsimony analysis of 1,271 informative 
characters and 253 species of Anolis and eight outgroups. Shown are two portions of the majority-rule consensus of 4,961 
maximum parsimony trees (i.e., this is not a bootstrap tree). Relationships present in less than 100% of optimal trees are 
identified with percentages above those clades or with clade collapsed if present in less than 50% of optimal trees. All taxa are 
Anolis. Figure 9A shows relationships of basal alpha Anolis ("latifrons" group sensu lalo, "dactyloids"); Fig. 9B shows 
relationships of mainland beta Anolis (mainland "Noropf). Other analyzed species are omitted for brevity. 




92 




2009 



NEW SOUTH AMERICAN ANOLIS 



15 



lemurinus within the mainland beta Anolis 
(Etheridge, 1959; = "Norops"). Anolis lyra 
shares several lemurinus-group characters 
with these species (e.g., narrow toepads, a 
dark interorbital bar, lack of enlarged post- 
cloacal scales, dorsal color pattern; see 
Kohler, 1999). The lemurinus group of Anolis 
might be nonmonophyletic (Nicholson, 
2002). In the current estimate, this group is 
composed of three putative clades: A. lemur- 
inus, A. bicaorum, and A. lyra; the large- 
hemipenes group, A. tropidolepis, A. pachy- 
pus, and A. magnaphallus; and A. vittigerus. 
Anolis vittigerus is superficially similar to A. 
lyra but displays an unusual combination of 
characters, including, for example, nasal 
overlap of the premaxilla. Future studies 
will examine whether these unusual condi- 
tions signal independent evolution or some 
less interesting phenomenon such as limited 
sample size. 

Remarks. With the description of these 
two species, the remaining nomen nudum 
Anolis from Ayala (1986) are A. "jericoen- 
sis" "A. maikinir and A. "urraoi." We have 
examined specimens of putative A. "jericoen- 
sis" and are unable to distinguish these from 
A. eulaemus. We are unable to distinguish 
preserved specimens of putative "A. malkini" 
from A. pentaprion. Specimens of putative A. 
"urraoi" are likely to represent a distinct 
species similar to A. fuscoauratus and A. 
maculiventris. Anolis fuscoauratus clearly is 
composed of multiple undescribed species 
(authors' personal observations); thus, de- 
scription of A. "urraoi" could be challenging. 

COMPARATIVE 
MATERIAL EXAMINED 

Museum of Comparative Zoology, Har- 
vard, U.S.A. = MCZ. Anolis fas ciatus MCZ 
143961, 147007, 154126, 151645, 151647. 
Anolis peraccae MCZ 170463, 170465, 
170473-5, 170479, 170486-7. Anolis vitti- 



gerus MCZ 115725, 154549, 158391, 
165184, 17190-1. See Poe (2004) for list of 
additional specimens examined. 

ACKNOWLEDGMENTS 

Thanks to Jim Hanken and the Museum 
of Comparative Zoology at Harvard for 
financial support of this work. Jose Rosado 
and the MCZ department of Herpetology 
were helpful and hospitable during work 
performed at Harvard. Joe Martinez and 
Emily Becker provided pertinent literature. 
Laszlo Meszoly rendered the drawings. We 
thank Juan D. Daza, Alexandra Herrera, 
Andres Quintero, Mario Yanez-Munoz, Erik 
Hulebak, and Eric Schaad for helping in the 
field and providing photos. Four reviewers 
provided helpful comments. 

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