Breviora

BREVIORA

Museum of Comparative Zoology

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 Pog,! JULIAN VELASCO,2 KENNETH MIYATA,3;4 AND ERNEST E. WILLIAMS?-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,” “‘/atifrons” group). Anolis lyra is placed with other
lemurinus-group Anolis within the large clade of mainland “beta” Anolis (“Norops’’).

Key worps: 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
pequefias, y en el patron de coloracion. Anolis anchicayae sp. nov. es parecido a Anolis peraccae y Anolis fasciatus
pero se diferencia principalmente en el tamafio y el patron de coloracion. Se codificaron caracteres filogenéticos de
morfologia externa y osteologia y se realizo un analisis de parsimonia con 251 especies adicionales de Anolis usando
una combinacion de datos morfoldgicos 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.”

‘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.

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.

i)

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

BREVIORA

No. 516

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 |. We
used PAUP* version 4.0b10 (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

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, Cano Docordo
between Cucurrupi and Noanama, 24 De-
cember 1968, B. Malkin; MCZ 158385-7,

NEW SOUTH AMERICAN ANOLIS 3

Colombia, Valle, Rio Blanco, road between
Cali and Anchicaya (where road crosses
river), ca. 500m 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, 1602349, Colom-
bia, Valle, San Isidro, Carton, 5 and 9
August 1977, H. Chin; MCZ 160008-12,
Colombia, Valle, Granje Bajo Central, 1976—
Wi, Is Chim, 1h, Castros lulCZ4 ioGow3so,
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,

Figure 2.

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

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absence of dewlap in females, absence of
caudal transverse vertebral processes poste-
riorly, and an elongate anterior nasal scale
that reaches the rostral but does not contact
the sulcus between the rostral and first

Anolis anchicayae, MCZ 145362, dorsal head scales.

supralabial. Anolis anchicayae is most easily
distinguished from A. peraccae by larger size
(to 54 mm SVL in A. peraccae; to 63 mm in
A. anchicayae; Fig. 6) and color pattern (A.
anchicayae is predominantly yellow-green on

. 516

2009 NEW SOUTH AMERICAN ANOLIS 5

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Figure 4. Anolis anchicayae, MCZ 145362, lateral 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 4A.
fasciatus by size (to 72mm 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

pret

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sen gnes : CY y the absence of transverse processes on

CHL A A posterior caudal vertebrae (present in beta
a SOO CHT tk 5 \ MI Anolis); from phenacosaur and _ ftigrinus-
i WO { group Anolis (A. heterodermus, A. inderenae,

A. nicefori, A. bellipeniculus, A. carlostoddi,
A. euskalerriari, A. neblininus, A. orcesi, A.

Figure 5. Anolis anchicayae, MCZ 145362, ventral ¢e¢aril, A. vanzolinii, A. ruizi, A. solitarius, A.
head scales. menta, A. nasofrontalis, A. pseudotigrinus, A.

6 BREVIORA

MALES

Anolis anchicayae
Valle, Chocd; Colombia

N= 20

%,

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

Figure 6.

Anolis peraccae
Narino; Colombia

Anolis peraccae
Pichincha; Ecuador

No. £16

FEMALES

SVL 42mm 44 46 48 50 52 54 56 58

SVL 42mm 44 46 48 50 52

4 56 58

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 Jatifrons- 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.
transversdalis, A. boettgeri, A. deltae, A.
dissimilis, A. santamartae) 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-group
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

pattern (solid bright green in A. chloris;
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,

NEW SOUTH AMERICAN ANOLIS i]

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 (2-4) 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

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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-

BREVIORA

No. 516

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, 10OA

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

Figure 7.

NEW SOUTH AMERICAN ANOLIS 9

200 Kilometers

a

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,
Narifio 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, | km N Buena Fe, 31 July 1975,
R. Webster; MCZ 1524334, 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 (¥ = 4.1 in A.
lyra, 2.4 in A. vittigerus; P = 0.001, Mann-
Whitney U test; Fig. 10). Also, A. Lyra tends
to display much weaker lateral striping (..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

lence SES

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

h i

A a Jae A

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

11

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2009

Figure 10. Head scales of (A) Anolis lyra, MCZ 77459, Lower Rio Calima, Valle, Colombia
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 BREVIORA

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. /yra.

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,
1524334, 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 (1-4) 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

No. 516

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

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.

NEW SOUTH AMERICAN ANOLIS 13

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 (1.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. 11.
Anolis anchicayae is sister species to A.
peraccae among an unresolved group of
South American alpha Anolis (approximate-
ly equivalent to Etheridge’s [1959] Jatifrons
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 fortunensis
A trachyderma
other Anolis tolimensis
aeneus medemi
roquet maculiventris
richardi granuliceps
67 bonairensis poecilopus
67, extremus tropidogaster
riseus lionotus
rinitatus oxylophus
luciae macrolepis
esperanzai : concolor
wiliamsmittermeierorum 92 pinchoti
Weel ial limifrons
roboscis Zeus
Inderenae 92 g2 92;— bicaorum
nicefori lyra <—_
solitarius femurinus.
rulzi if galpenten ant
aequatorialis ocelloscapularis
cadildae ee humilis a
71 fitchi . notopholis
fraseri compressicauda
74 jacare magnaphallus
microtus pachypus
insignis 92 tropidolepis
squamulatus capito
frenatus cuprinus
latifrons tropidonotus
ventrimaculatus 92 cobanensis
apollinaris parvicirculatus
agassizi polylepis
chioris cupreus
festae 92 dollfusianus
punctatus altae
USCOeNSIS kemptoni
uilae townsendi
transyersalis Moreuthes
anchicayae <—=— 92 anton
peraccae isthmicus
iasciatle microlepidotus
sericeus
Boe tigeri intermedius
outgroups laeviventris
pentaprion
vociferans
92 ortoni :
megapholidotus
utilensis
subocularis
nebuloides
quercorum
nebulosus
crassulus
sminthus
liogaster
dunni
adovi
aylori
viltigerus
we Webel
92 aquaticus
92 biporcatus
petersi
92 gracilipes
uniformis
92 bitectus _.
polyrhachis
loveridgei_ .
purpurgularis
92 nitens chrysolepis

nitens scypheus
meridionalis
auratus
annectens

onca |

barkeri

lineatus

Figure 11. Relationships of (A) Anolis anchicayae and (B) A. lyra 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 lato, ““dactyloids”); Fig. 9B shows
relationships of mainland beta Anolis (mainland “Norops’’). Other analyzed species are omitted for brevity.

2009

lemurinus within the mainland beta Anolis
(Etheridge, 1959; = “Norops’’). Anolis lyra
shares several /emurinus-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 Jemurinus group of Anolis
might be nonmonophyletic (Nicholson,
2002). In the current estimate, this group is
composed of three putative clades: A. /emur-
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. malkini,’ 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 fasciatus MCZ

143961, 147007, 154126, 151645, 151647.
Anolis peraccae MCZ_ 170463, 170465,
1704735, 170479, 170486—-7. Anolis vitti-

NEW SOUTH AMERICAN ANOLIS 15

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,
Andrés Quintero, Mario Yanez-Munioz, Erik
Hulebak, and Eric Schaad for helping in the
field and providing photos. Four reviewers
provided helpful comments.

LITERATURE CITED

AYALA, S. C. 1986. Saurios de Colombia: lista actuali-
zada y distribucion de ejemplares colombianos en
los museos. Caldasia, 15: 555-575.

, AND F. Castro. 1983. Dos nuevos gecos (Sauria:

Gekkonidae, Sphaerodactylinae) para Colombia:

Lepidoblepharis xanthostigma (Noble) y descripcion

de una nueva especie. Caldasia, 8: 743-53.

, AND E. E. WILLIAMS. 1988. New or problematic
Anolis fro Colombia, VI. Two fuscoauratoid anoles
from the Pacific lowlands, A. maculiventris Boulen-
ger, 1898 and A. medemi, a new species from
Gorgona Island. Breviora, 490: 1-16.

ETHERIDGE, R. E. 1959. The relationships of the anoles
(Reptilia: Sauria: Iguanidae): an interpretation
based on skeletal morphology Univ. Microfilms:
Ann Arbor, Michigan, xiii + 236 pp.

GautTHieR, J.. A. G. KLuGE, AND T. Rowe. 1988.
Amniote phylogeny and the importance of fossils.
Cladistics, 4: 105-209.

Guyer, C., AND J. M. SAvAGE. 1986. Cladistic relation-
ships among anoles (Sauria: Iguanidae). Systematic
Zoology, 35: 509-531.

Henpy, M. D., AND D. Penny. 1989. A framework for
the quantitative study of evolutionary trees. Sys-
tematic Zoology, 38: 297-309.

HUELSENBECK, J. P. 1991. When are fossils better than
extant taxa in phylogenetic analysis? Systematic
Zoology, 40: 458-469.

16 BREVIORA

JACKMAN, T. R., A. LARSON, K. DE QUEIROZ, AND J. B.
Losos. 1999. Phylogenetic relationships and tempo
of early diversification in Anolis lizards. Systematic
Biology, 48: 254-285.

Kouter, G. 1999. Eine neue saumfingerart der gattung
Norops von der Pazifikseite des nordlichen Mittel-
amerika. Salamandra, 35: 37-52.

NICHOLSON, K. E. 2002. Phylogenetic analysis and a test of
the current infrageneric classification of Norops (beta
Anolis). Herpetological Monographs, 16: 93-120.

, R. E. Gtor, J. J. Koise, A. Larson, S. B. HEDGES,
AND J. B. Losos. 2005. Mainland colonization by
island lizards. Journal of Biogeography, 32: 929-938.

Nixon, K. C. 1999. The Parsimony Ratchet, a new
method for rapid parsimony analysis. Cladistics, 15:
407-414.

Por, S. 1998. Skull characters and the cladistic
relationships of the Hispaniolan dwarf twig Anolis.
Herpetological Monographs, 12: 192-236.

2004. Phylogeny of anoles. Herpetological
Monographs, 18: 37-89.

Rueba, A. J. V., AND E. E. WILLIAMs. 1986. Una nueva
especie de saurio para Cordillera Oriental de
Colombia (Sauria, Iguanidae). Caldasia, 13: 511-524.

SAVAGE, J. M., AND C. Guyer. 1989. Infrageneric
classification and species composition of the anole
genera, Anolis, Ctenonotus, Dactyloa, Norops, and
Semiurus (Sauria: Iguanidae). Amphibia-Reptilia,
10: 105-116.

Sikes, D. S., AND P. O. Lewis. 2001. Beta software,
version 1. PAUPRat: PAUP* implementation of
the parsimony ratchet. Distributed by the authors.
Department of Ecology and Evolutionary Biology,
University of Connecticut, Storrs.

No. 516

Simpson, G. G. 1961. Principles of Animal Taxonomy.
New York: Columbia University Press.

SworrorD, D. L. 2002. Phylogenetic analysis using
parsimony (*and other methods). Version 4 Sinauer
Associates: Sunderland, Massachusetts.

Wiens, J. J.. AND M. R. Servepio. 2000. Species
delimitation in systematics: inferring diagnostic
differences between species. Proceedings of the
Royal Society of London Series B, 267(1444):
631-636.

Witey, E. O. 1978. The evolutionary species concept
reconsidered. Systematic Zoology, 27: 17-26.
WILLIAMS, E. E. 1982. Three new species of the Anolis
punctatus complex from Amazonian and _ inter-
Andean Colombia, with comments on the eastern
members of the punctatus species group. Breviora,

467: 1-38.

. 1988. New or problematic Anolis from Colom-

bia. V. Anolis danieli, a new species of the latifrons

species group and a reassessment of Anolis apolli-

naris Boulenger, 1919. Breviora, 489: 1—25.

. 1992. New or problematic Anolis from Colom-

bia. VII. Anolis lamari, a new anole from the

Cordillera Oriental of Colombia, with discussion of

tigrinus and punctatus species group boundaries.

Breviora, 495: 1—24.

, H. RANpb, A. S. RAND, AND R. J. O'HARA. 1995.

A computer approach to the comparison and

identification of species in difficult taxonomic

groups. Breviora, 502: 1-47.

, M. J. PRADEIRO, AND S. GorzuLa. 1996. A

phenacosaw from Chiamata Tepui, Venezuela.

Breviora, 506: 1—15.