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Full text of "Breviora"

BREVIORA 



JVL ui s e m mn ox LompaFatiTe Zoology 




'*«««*...... 



US ISSN 0006-9698 



Cambridge, Mass. 



31 May 2012 



Number 530 



MORPHOLOGY, PHYLOGENY, AND BEHAVIOR OF ANOLIS PROBOSCIS 

Steven Poe, 1 Fernando Ayala, 2 Ian M. Latella, 1 Thomas L. Kennedy, 1 James A. Christensen, 3 
Levi N. Gray, 1 Natalie J. Blea, 1 Beeling M. Armtjo, 4 and Eric W. Schaad 1 

Abstract. Males of the three species of the Anolis laevis group, so-called proboscis anoles, display a remarkable 
appendage extending from the snout. All A. laevis group Anolis are poorly known and rarely collected. We redescribe 
Anolis proboscis based on the type specimen and male and female specimens we collected recently near the town of 
Mindo, Pichincha Province, Ecuador. We include color notes and additional morphological measures lacking from 
the original description, natural history data for 1 1 specimens, an estimate of the phylogenetic relationships of the 
species, and a description of display behavior and intraspecific interactions incorporating the proboscis. 

Key words: Anolis proboscis; Ecuador; redescription, behavior 



Among the myriad strange and varied 
Anolis, from semiaquatic forms such as A. 
barker i to a blue insular species A. gorgonae 
to a colorful dewlapless cliff dweller A. 
bartschi, the three species of proboscis anoles 



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, ilatella@ 
unm.edu, tkennedy@unm.edu, lngray@unm.edu, natblea505@ 
gmail.com, ewschaad@unm.edu. 

2 Laboratorio de Reptiles, Museo de Zoologia QCAZ, 
Pontificia Universidad Catolica del Ecuador, Quito, 
Ecuador; e-mail: fpayala2000@yahoo.com. 

3 P.O. Box 2259, Port Angeles, Washington 98362, 
U.S.A.; e-mail: james.a.christensen@gmail.com. 

4 School of Medicine, University of New Mexico, 
Albuquerque, New Mexico 87131, U.S.A.; e-mail: 
littlebee2506@hotmail.com. 



with their scaly anterior snout extensions 
stand out as especially curious. Originally 
described from few specimens and no fe- 
males (Scytomycterus [Anolis] laevis Cope 
1876, n = 1; A. phyllorhinus Myers and 
Carvalho 1945, n = 2; A proboscis Peters 
and Orces-V 1956, n = 6) and for many 
decades unreported in life since their descrip- 
tions, two of the three species of proboscis 
anoles have recently been rediscovered. 
Rodrigues et al. (2002) described the first 
female proboscis anoles of A. phyllorhinus 
from Amazonia. Almendariz and Vogt 
(2007) reported on a male A. proboscis cap- 
tured but released near Mindo, Ecuador, and 
Yanez-Munoz et al. (2010) described varia- 
tion in five specimens of A. proboscis. 



The President and Fellows of Harvard College 2012. 



BREVIORA 



No. 530 



ALTITU 


DE (m) 

0-500 
501 - 1000 
1001 -2000 
2001 - 3000 
3001 - 4000 
>4000 


1 1 


1 I 


■i 






Figure 1. Collecting localities for Anolis proboscis near 
Mindo, Pichincha Province, Ecuador (latitude, longitude, 
elevation in parentheses): (1) Tulipe (0°4'58.8"N, 
-78°46'58.8"W) (2) Las Tolas (0°2'45.5994"N, 
-78°47'5.9994"W), (3) Cunuco, 5 km NW Mindo 
(-0°1'58.8"S, -78°48'28.8"W), (4) 5 km E Mindo 
(-0°1'55.2"S, -78°45'36"W), (5) turnoff to Mindo, 1- 
5 km down from intersection (-0° 1 '55.2"S, -78°45'36"W). 



The paucity of specimens of A. laevis and 
A. phyllorhinus in museum collections may 
be explained by the remote ranges of these 
species. The single specimen of A. laevis is 
from a mountain trail connecting two lower 
elevation Peruvian towns, and A. phyllorhi- 
nus is an Amazonian forest form with an 
apparently small distribution. The rarity of 
A. proboscis in collections is more surprising. 
The type locality of this species is a 
populated area within 3 hours drive of Quito 
along a well-traveled, paved road. The area 
in which the species has been found is a 
famous birding site with several hotels 
frequented by ecotourist groups and scien- 
tists alike. 



In August 2009, a joint expedition of the 
University of New Mexico and the Pontificia 
Universidad Catolica del Ecuador, Quito 
collected three males and two females of 
A. proboscis in approximately 8 hours of 
searching along the paved road leading from 
the major western Ecuadorean Highway, Via 
Calacali-Los Bancos, to Mindo (Fig. 1). 
This locality is within 5 km of Cunuco, the 
type locality of A. proboscis. A second 
expedition in December 2009 found six 
specimens of A. proboscis in approximately 
2 hours of searching at the same locality. 

The type specimen of A. proboscis was well 
described by Peters and Orces-V. (1956), but 
the description lacked information on some 
standard characters in Anolis, such as number 
of scales across the snout between the second 
canthals (a common measure of head scale 
size), Williams (1979) reviewed the proboscis 
anoles but had no information on them in life. 
Yanez-Mufioz et al. (2010) described external 
variation in their newly collected material but 
did not report on osteology, redescribe the type 
specimen, or compare their material to previ- 
ously collected specimens. Here we present a 
redescription of the holotype of A. proboscis 
(MCZ 54300); describe variation in newly 
collected and previously preserved specimens 
of the species, including characters not previ- 
ously scored in A. proboscis, such as toe length, 
size of ventral and dorsal body scales, and skull 
structure; perform a phylogenetic analysis to 
determine the placement of this species within 
Anolis\ and present data on the color, behav- 
ior, and ecology of A. proboscis in life. 

MATERIALS AND METHODS 

Measurements were made with digital 
calipers on preserved specimens and are given 
in millimeters (mm), usually to the nearest 
0.1 mm. Snout to vent length (SVL) was 
measured from the anterior tip of the lower 
jaw to the anterior margin of the cloaca. Head 



2012 



BIOLOGY OF ANOLIS PROBOSCIS 



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 knee bent at a 90° angle. 
Head width was measured at the broadest 
part of the head between the posterolateral 
corners of the orbits. Tail length was mea- 
sured from the anterior margin of the cloaca 
to the tip of the tail. Toe length was measured 
from the origin of the fourth (i.e., longest) toe 
to the tip of its claw. Scale terminology and 
characters used mainly follow standards 
estabflished by Williams (e.g., Williams et al, 
1995) for species descriptions of Ano lis lizards. 
Skull description is given in terms of Ether- 
idge's (1959) and Poe's (1998, 2004) charac- 
ters. See those papers for more detailed 
descriptions of skull conditions and alterna- 
tive conditions in Anolis. 

For phylogenetic analysis, we added 59 
species to the dataset of Poe (2004), which 
included A. proboscis. Species were added 
to obtain a better estimate of the closest 
relatives of A. proboscis because many 
potential sister species for A. proboscis were 
not included by Poe (2004). 

We generated a novel phylogenetic estimate 
of A. proboscis using a dataset of 91 morpho- 
logical characters (Poe, 2004; unpublished 
morphological data for 57 additional species), 
979 sites of mtDNA (NADH dehydrogenase 
subunit 2, five transfer RNAs; Nicholson et al. , 
2005), and 1,179 sites of nuclear DNA (internal 
transcribed tracer region; Nicholson 2002) for 
265 species of Anolis and eight outgroups. Data 
coverage varied from all 1,267 parsimony- 
informative characters scored to 52 parsimony- 
informative characters for some species scored 
only for external morphology. The matrix in- 
cludes large amounts of missing data; for 75 
species, only varying amounts of morpholog- 
ical data were scored. 

We used a partitioned Bayesian analysis 
with separate GTR + gamma models for the 
two DNA data sets and the "standard" 



model for the morphological data with 
"informative" coding and "ordered" and 
"unordered" coding for characters according 
to Poe (2004). Topology was assumed to be 
shared between each of the three partitions, 
whereas branch lengths were allowed to vary. 
We ran two independent analyses for 10 
million generations, sampling every 500 gen- 
erations, resulting in 20,001 trees. For each 
analysis, we ran three heated and one cold 
chain with heating temperature = 0.1. We 
checked for convergence by monitoring the 
standard deviation of split frequencies and the 
natural log likelihoods of each sampled 
generation. We discarded the first 19,000 trees 
as burn-in, leaving 1,001 trees for analysis. 

We also performed a parsimony analysis 
in PAUP* (Swofford, 2002) with all charac- 
ters weighted equally. We used the parsimo- 
ny ratchet (150-200 repetitions, 10-35% 
character removal; Nixon, 1999) to find an 
optimal tree and then swapped on this tree 
using the tree-bisection-reconnection search 
in PAUP*. Support for individual clades was 
evaluated using the bootstrap (Felsenstein, 
1985). For bootstrap analysis, 100 replicate 
matrices of the same size as the original were 
constructed by sampling characters with 
replacement. Replicate matrices were ana- 
lyzed with parsimony using one round of 
random taxon addition and TBR branch 
swapping with a time limit of 600 seconds 
per rep. Resulting trees were summarized by 
mapping bootstrap values greater than 50% 
onto the optimal parsimony tree. 

REDESCRIPTION OF A. PROBOSCIS 

Anolis proboscis Peters and Orces-V. 1956 
Figures 2-3 

Holotype. MCZ 54300, adult male, Ecua- 
dor, Cunuco, 5 km NW of Mindo, 27-29 
April 1953, Antonio Proano collector. 

Additional Material. USNM 207671, adult 
male, Ecuador, vicinity of Mindo, date of 



BREVIORA 



No. 530 




Figure 2. Anolis proboscis collected near Mindo, Ecuador. A, male; B, female. 



2012 



BIOLOGY OF ANOLIS PROBOSCIS 






Figure 3. Dewlap of Anolis proboscis. A, male; B, female. 



collection unknown, Rosero collector; 2009; QCAZ 9718 adult female, 5 

USNM 207672-3, adult males, Ecuador, August 2009; QCAZ 9734-35 adult males, 

vicinity of Mindo, August 1964, M. Olalla 7 August 2009; QCAZ 9753, adult female, 10 

collector; QCAZ 9717, adult male, 5 August August 2009; Ecuador, 1-4 km from turnoff 



BREVIORA 



No. 530 



toward Mindo (i.e., 2-5 km N of Mindo; 
approximately latitude — 0°1'58.8"S, longi- 
tude -78°45'36"W); Steven Poe, Fernando 
Ayala, Eric Schaad, Ian Latella, Thomas 
Kennedy, Natalie Blea collectors; QCAZ 
8061-63, 10209-11, same locality as other 
QCAZ specimens; Ecuador, 1-4 km from 
turnoff toward Mindo (i.e., 2-5 km N of 
Mindo), 28 December 2009, Steven Poe, Ian 
Latella, Levi Gray, and Beeling Armijo 
collectors. 

Description of External Morphology of the 
Holotype (MCZ 54300) of Anolis proboscis 
(Variation in Additional Material in Paren- 
theses, Based on USNM 207671-3, QCAZ 
9717-8, 9735, 9753). SVL 72.9 mm (males: 
61.1-75.1 mm; females: 70.4-72.6 mm); head 
length 22.3 mm, 0.31 ratio of head length to 
SVL (males: 0.26-0.31 SVL; females 0.26 
SVL), width 11.1 mm, 0.15 SVL (males: 
0.14-0.15 SVL; females 0.14 SVL); ear height 

1.3 mm, 0.01 SVL (males: 0.01-0.02 SVL; 
females: 0.02); femoral length 14.4 mm, 0.20 
SVL (males 0.20-0.22 SVL; females 0.21); 
tail length not recorded in Type (males 1.1- 

1.4 SVL; females 1.3 SVL); fourth toe length 
not recorded in Type (males 0.13-0.14 SVL, 
females 0.12-0.13 SVL). 

Dorsal head scales smooth; frontal depres- 
sion absent (weak); rostral extends over 
mental, reaching anterior triangular peak 
inferior to proximal aspect of proboscis; 
eight (8-10) scales across snout between 
second canthals; supraorbital semicircles 
separated by two scales; suboculars in con- 
tact with supralabials; elongate supraciliary 
scales not evident, only small undifferentiat- 
ed scales present; four to five loreal rows; 
circular nasal scale contacts first supralabial, 
separated from rostral by two scales; inter- 
parietal length 2.6 mm (0.04 SVL; 0.03- 
0.04); ratio of length of interparietal to 
length of second largest scale lateral to in- 
terparietal 3.7 (2.1-2.4); two scales separate 
interparietal and supraorbital semicircles; 



preoccipital absent; very large sublabials, 
two to four in contact with supralabials; 10 
(9-12) supralabials to center of eye; 8-13 
postrostrals; four (3-4) postmentals; some 
slightly enlarged scales present in supraocu- 
lar disc, decreasing gradually in size; mental 
partially divided posteriorly, extending pos- 
terolaterally beyond rostral, with posterior 
border in concave arc (or concave); five (2-6) 
greatly enlarged sublabials, three (2-3) in 
contact with infralabials; dewlap reaches just 
posterior to axillae (same in males and 
females); scales on dewlap closely packed, 
not in regular rows; no axillary pocket; a pair 
of greatly enlarged postcloacal scales (absent 
in females); nuchal, dorsal, and caudal crests 
present as raised spikes separated by undif- 
ferentiated scales (Lazell's [1969] type 5); 
dorsal scales smooth; middorsal scales equal 
in size to lateral scales (except crest), 10 (9- 
11) longitudinal rows in 5% of SVL; ventral 
scales in transverse rows, smooth, nine (6-9) 
scales in 5% of SVL; limb and supradigital 
scales smooth; toepads expanded; 19 (20-22) 
expanded lamellae under fourth toe. 

The proboscis (Fig. 2A; absent in fe- 
males) is a laterally compressed, pliable 
appendage covered with somewhat elongate 
scales that are slightly smaller than the 
dorsal headscales. The posterior aspect of 
the proboscis rises longitudinally from the 
midsnout region and extends forward with 
a partially serrated dorsal edge, eventually 
coming to a blunt anterior end. Neither the 
rostral nor the supralabials form significant 
parts of the proboscis, although the supra- 
labial/loreal scale demarcation extends an- 
teriorly as a line along the ventrolateral edge 
of the appendage. The proboscis extends 
anteriorly 85% of the head length beyond 
the snout (mean 77-89.5% for three adult 
males; 62% for one juvenile male of SVL 53 
mm; Table 1). 

Skull Description (Based on Dry Skeletons 
USNM 207673, QCAZ 9734). Parietal roof 



2012 



BIOLOGY OF AN O LIS PROBOSCIS 



Table 1 . Measurements of specimens of Anolis 

proboscis showing length of proboscis relative to 

snout to vent length (svl) and head length. 

Measurements are in millimeters. 







Head 


Proboscis 


Specimen 


SVL 


Length 


Length 


QCAZ9717 


59.9 


16.9 


13.9 


QCAZ 9734 


NA 


19.7 


17.6 


QCAZ 9735 


69.3 


19.4 


15.0 


QCAZ 8061 


53.4 


16.4 


10.1 


QCAZ 8063 


73.1 


22.8 


19.6 



Note: Snout to vent length for specimen QCAZ 9734 
was unavailable because the body of the specimen was 
not intact. 



flat, with convex V- or Y-shaped crests, with 
slight casquing, lacking crenulation on edges, 
extending slightly posteriorly over supraoc- 
cipital; anterolateral corners flush with pos- 
terolateral edges of frontal; pineal foramen at 
parietal-frontal suture; dorsal skull bones 
smooth; postfrontal present; prefrontal con- 
tacts nasal between frontal and maxilla; 
frontal sutures anteriorly with nasals; no 
parallel crests on nasals; external nares 
bordered posteriorly by nasals; dorsal aspect 
of jugal terminates on lateral surface of 
postorbital; jugal contacts squamosal; pos- 
terodorsal ramus of squamosal long, may 
contact parietal above supratemporal; poste- 
rior aspect of jugal mostly straight; epipter- 
ygoid short, does not contact parietal dorsal- 
ly; pterygoid and palatine teeth absent; lateral 
edge of vomer smooth, without posteriorly 
directed lateral processes; maxilla extends 
posteriorly beyond ectopterygoid on ventral 
surface of skull; crest between basipterygoid 
processes of basisphenoid absent; lateral shelf 
of quadrate absent; some black pigment on 
the dorsal surface of parietal and frontal; 
nasals overlap premaxilla dorsally or are 
flush with it; dorsal surface of supraoccipital 
with single central process; posteriormost 
mandibular tooth partially anterior to anteri- 
or mylohyoid foramen; large splenial present; 



ventral aspect of anteromedial process of 
coronoid extends posteriorly; labial process 
of coronoid present; external opening of 
surangular foramen is entirely within suran- 
gular; posterior suture of dentary is pronged; 
anteriormost aspect of posterior border of 
dentary anterior to mandibular fossa; jaw 
sculpturing absent; angular absent; angular 
process of articular present, small. 

Color in Life ( Adapted from Field Notes). 
Below we describe two color morphs each 
in males and females. QCAZ 9717 (male; 
Fig. 2A): dorsum white laterally with three 
broad peach blotches outlined with black (one 
above axilla, two around midbody); middor- 
sum green becoming orangeish at pelvis, with 
orange extending back onto tail; top of head 
brown, lacking interorbital bar; white from 
under eye back to body; peach blotch above 
axilla; limbs mottled rust and green; belly white 
with brown; tail strongly banded; dewlap 
white with white scales, with some scales faint 
yellow-green or peach (Fig. 3A); throat off- 
white; tongue tan; iris brown; eyelids white; 
scales on proboscis orange-brown. 

In the second male morph (QCAZ 9734), 
the lateral blotches (midbody and axillary) 
are entirely black rather than peach with 
black edges. In at least one specimen (QCAZ 
9735), the proboscis is greenish yellow 
anteriorly. 

QCAZ 9718 (female) dorsum brown, 
almost no green, largely patternless but 
somewhat mottled with lighter brown, black, 
and peach; top of head orange-brown, 
peach-brown laterally; belly cream with 
peach tint; tail faintly banded; iris brown; 
dewlap skin white, scales white to peach 
(Fig. 3B). 

The larger female (QCAZ 9753; Fig. 2B) 
was recorded as more greenish-brown than 
QCAZ 9718. 

Neither males nor females exhibited much 
color change while alive beyond slight 
darkening and lightening. 



BREVIORA 



No. 530 



Ecology, Behavior, and Habitat. The 6.3- 
km spur road connecting Mindo to the Via 
Calacali-Los Bancos Highway descends 
steeply through pasture and good patches 
of secondary forest from an altitude of 
1,679 m at the highway to 1,275 m at the 
bridge, signaling the entrance to Mindo. 
Despite at least equal effort in other areas 
around Mindo, including some with better 
forest at similar elevations, all of our 
specimens of A. proboscis were collected 
along this road. Anolis gemmosus, A. fraseri, 
and A. aequatorialis are sympatric with A. 
proboscis here. We found perhaps 50 indi- 
viduals of A. gemmosus per A. proboscis 
record, with most of these on ferns less than 
3 m above the ground. Anolis aequatorialis 
was less abundant than A. gemmosus, but 
still common, and was extremely abundant 
in nearby forest. We found only two A. 
fraseri along the road to Mindo but collected 
several specimens of this species at a slightly 
lower elevation along the Rio Mindo just 
northwest of town. 

All 11 of our specimens of A. proboscis 
were collected sleeping at night horizontally 
on twigs, except QCAZ 8061. QCAZ 8061 is 
a juvenile male that was found sleeping in a 
vertical position on the twig of a tall tree 3 m 
above ground in steep, highly disturbed 
terrain in a grove of sparsely vegetated trees 
by the road. QCAZ 9717 was 3.5 m above- 
ground in a tangle of bare twigs below a 
dense higher canopy in a patch of good 
forest. QCAZ 9718 was 8 m aboveground on 
a bare twig in a disturbed area adjacent to 
good forest. QCAZ 9734 was approximately 
9 m above the side of a ravine (approximate- 
ly a meter above eye level from the edge of 
the ravine) on a bare twig in a disturbed area. 
QCAZ 9735 was 4 m aboveground on a leafy 
twig in a highly disturbed area of pasture 
with isolated brush and small trees. QCAZ 
9753 was 6 m aboveground on a leafy twig in 
the same area. 



The sleeping behaviors of QCAZ 8062-63, 
10209-11 were recorded as follows. Two 
specimens (male and female) were collected 
approximately 8 m aboveground, within 2.5 m 
of each other in a sparsely vegetated tree on 
the edge of good forest by the road. Two 
specimens were collected on twigs of small (~ 
4 m) trees at 2 and 2.5 m height, respectively, 
along the road bordering pasture. One adult 
female was 3.5 m aboveground on a bare twig 
in pasture approximately 30 m from the road. 

The sleeping posture of A. proboscis is 
reminiscent of twig anoles of the Caribbe- 
an (Thomas, 1965) and Phenacosaurus 
anoles of South America. Captive behav- 
ior suggests the tail is prehensile, as spec- 
imens periodically wrapped the tail around 
twigs. 

Subsequent to our visits to where the 
species was initially collected, one of us 
(J.C.) was able to observe the behavior of 
A. proboscis in life. He provided the follow- 
ing notes: 

"I had an opportunity to photograph a full 
courtship to copulation sequence, but things 
proceeded so quickly that I was left only with a 
few shots of the copulating pair. Notwith- 
standing, I was at least able to observe the 
employment of the proboscis in the courtship 
display. Predictably, before closing the distance 
the male dewlap displayed, followed by a rush 
toward the female, during which the proboscis 
was flourished quickly from side to side in 
several stereotyped, rather 'mechanical look- 
ing' sweeps of the head. The male then elevated 
the proboscis to 45 degrees and then appeared 
to seize the female by the nape in typical Anolis 
fashion (my view was partially obstructed at 
times). The male moved very rapidly and 
aggressively throughout, and copulation oc- 
curred within seconds of the initiation of 
courtship behavior, in contrast to the much 
lighter touch that I have observed in, for 
example, A. gemmosus."" 



2012 



BIOLOGY OF AN O LIS PROBOSCIS 



"Having in my possession two males of 
similar size, I decided to place them together 
on a branch and attempt to record any 
resulting behavior. To my surprise the indi- 
vidual that had seemed more robust, and that 
had fed more freely in captivity, immediately 
deferred to a male that appeared slighter of 
build. The submissive male flattened himself 
against the limb, adopted a dark brown 
coloration, and remained immobile while 
being threatened by the other. The aggressor 
assumed bright colours, laterally compressed 
its body, gaped, flared its dewlap while 
bobbing its head slowly, and approached very 
deliberately, with proboscis arrow straight. As 
the dominant male neared the submissive one 
it pointed its head downward at a shallow 
angle and brought its proboscis parallel with 
the head of the other male, sometimes 
touching it lightly. It maintained this stance 
until the submissive male broke and ran. 
When these two animals were released togeth- 
er at the capture site, hours later, the 
submissive male remained terrified of the 
other and climbed rapidly away from it." 

"A male, some time after being removed 
from its terrarium and placed on branches, 
spontaneously dewlap displayed while scanning 
its surroundings. I never observed any appar- 
ent use of the proboscis as a long-range signal." 

"Captive animals fed readily on small 
Acrididae. A male fed on a small caterpillar 
and a leaf beetle while at liberty during a photo 
shoot. A female, released at its capture site, 
attacked and began to consume a polydesmid 
millipede within minutes of being set free. 
Males elevated the proboscis to roughly 45 
degrees when stalking and seizing prey." 

Relationships. The Bayesian analysis failed 
to converge after 45 days of computer time 
(standard deviation of split frequencies, ~ 
0.07). The resulting majority-rule consensus 
of post-burn-in trees placed A. proboscis in 
a large basal polytomy with other members 



of the Dactyloa clade (Etheridge, 1959; 
Guyer and Savage, 1986; Castaneda and 
de Queiroz, 2011) or latifrons-senQS (Ether- 
idge, 1959) species. The parsimony results 
were better resolved, and we thus present 
these results to communicate a tentative 
hypothesis for the relationships of A. 
proboscis. 

A part of the strict consensus of most 
parsimonious trees is shown in Figure 4. All 
relationships shown were present in fewer 
than 50% of bootstrap replicates. Anolis 
proboscis appears as a member of a clade 
of Phenacosaurus-gvoup Anolis. These high- 
elevation South American species share large 
smooth headscales and short limbs and tail. 
Morphologically, they resemble twig-ecomorph 
Anolis of the Caribbean (Losos, 2009). We 
were unable to obtain specimens of the other 
proboscis anoles A. laevis and A. phyllorhinus, 
and future work will include these species 
in phylogenetic analyses. The external mor- 
phology of A. laevis (Williams, 1979) suggests 
it also is a member of the phenacosaur 
clade, but A. phyllorhinus appears more similar 
to A. punctatus (Rodriguez et al. , 2002), which 
was placed outside of the clade shown in 
Figure 4. 

Concluding Comments. The recent redis- 
covery of Anolis proboscis and subsequent 
flood of information on it has offered a 
window into the life of a remarkable species, 
but much remains to be learned of A. 
proboscis and the other proboscis anoles. 
The Peruvian proboscis anole A. laevis 
awaits rediscovery, and the phylogenetic 
relationships of all proboscis anoles remain 
poorly known. The diurnal behavior of these 
species is only beginning to be understood, 
and the evolution of the proboscis (both 
within Anolis and in comparison with distantly 
related forms such as Ceratophora tennenti); 
its use in natural breeding, territorial displays 
in conjunction with or independent of the 
dewlap, or both; and its internal anatomy are 



10 



BREVIORA 



No. 530 



rC 



heterodermus 

proboscis 

inderenae 

nicefori 
solitarius 



C 



new sp. 

williamsmittermeierorum 

ruizi 



roquet group 



Figure 4. Estimated relationships of Anolis proboscis 
according to a strict consensus of optimal trees from 
parsimony analyses. Clade shown is nested within the basal 
alpha group of Anolis (i.e., Etheridge's [1959] latifrons series). 

all fertile subjects for future research. We 
hope the present work helps to lay a 
foundation for the understanding of these 
cryptic and fantastic forms. 

ACKNOWLEDGMENTS 

We thank Omar Torres and the Pontificia 
Universidad Catolica Quito del Ecuador for 
facilitating our fieldwork in Ecuador. Funding 
for this project was provided by National 
Science Foundation Grant DEB-0844624 to 
Steven Poe, and Secretaria Nacional de Ciencia 
y Tecnologia del Ecuador PIC-08-0000470 to 
Omar Torres. Specimens were collected under 
collection permit No. 008-09 IC-FAU-DNB/ 
MA issued by Ministerio del Ambiente del 
Ecuador. We thank the reviewers for comments 
that improved the manuscript. 

LITERATURE CITED 

AlmendAriz, A. C, and C. Vogt. 2007. Anolis proboscis 
(Squamata: Polychrotidae), Una lagartija rara pero 
no extinta. Politecnica 27(4) Biologia, 7: 133-135. 



Castaneda, M. R., and K. de Queiroz. 2011. Phyloge- 
netic relationships of the Dactyloa clade of Anolis 
lizards based on nuclear and mitochondrial DNA 
sequence data. Molecular Phylogenetics and Evolu- 
tion, doi: 1 0.1 01 6/j.ympev.20 11.07.004. 

Cope, E. D. 1876. Report on the reptiles brought by 
Professor James Orton from the middle and upper 
Amazon and western Peru. Journal of the Academy 
of Natural Sciences of Philadelphia N. S. , 8: 1 59- 
183. 

Etheridge, R. E. 1959. The relationships of the 
anoles (Reptilia: Sauria: Iguanidae): an interpre- 
tation based on skeletal morphology. Ph.D. 
Dissertation. Ann Arbor, Michigan, University 
of Michigan. 

Felsenstein, J. 1985. Phylogenies and the Comparative 
Method. The American Naturalist, 125: 1-15. 

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

Lazell, J. D., Jr. 1969. The genus Phenacosaurus 
(Sauria, Iguanidae). Breviora, 325: 1-24. 

Losos, J. B. 2009. Lizards in an Evolutionary Tree: 
Ecology and Adaptive Radiation of Anoles. Berkeley, 
California, University. of California Press. 

Myers, G. S., and A. L. Carvalho. 1945. A strange 
leaf-nosed lizard of the genus Anolis from Amazo- 
nia. Boletim do Museu Nacional-Zoologia, 43: 1^4. 

Nicholson, K. E. 2002. Phylogenetic analysis. Phyloge- 
netic analyses and a test of the current infrageneric 
classification of Nor ops (beta Anolis). Herpetologi- 
cal Monographs, 16: 93-120. 

, R. E. Glor, J. J. Kolbe, A. Larson, S. B. 

Hedges, and J. B. Losos. 2005. Mainland coloni- 
zation by island lizards. Journal of Biogeography, 
32: 1-10. 

Nixon, K. C. 1999. The parsimony ratchet, a new 
method for rapid parsimony analysis. Cladistics, 15: 
407^14. 

Peters, J. A., and G. V. Orces. 1956. A third leaf-nosed 
species of the lizard genus Anolis from South 
America. Breviora, 62: 1-8. 

Poe, 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. 

Rodrigues, M. T., V. Xavier, G. Sk.uk, and D. Pavan. 
2002. New specimens of Anolis phyllorhinus (Squamata, 
Polychrotidae): the first female of the species and 
of proboscid anoles. Papeis Avulsos de Zoologia. Museu 
de Zoologia da Universidade de Sao Paulo, 42: 363- 
380. 



2012 



BIOLOGY OF ANOLIS PROBOSCIS 



11 



Swofford, D. 2002. PAUP*. Phylogenetic Analysis 
Using Parsimony (*and Other Methods). Version 
4. Sunderland, Massachusetts, Sinauer Associates. 

Thomas, R. 1965. A new anole (Sauria: Iguanidae) from 
Puerto Rico. Part II. Field observations of Anolis 
occultus Williams and Rivero. Breviora, 231: 10- 
18. 

Williams, E. E. 1979. South American anoles: the 
species groups. 2. The proboscis anoles {Anolis 
laevis group). Breviora, 449: 1-19. 



— , H. Rand, 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. 
Yanez-Munoz, M. H., M. A. Urgiles, M. Altamirano B, 
and S. R. CAceres S. 2010. Redescripcion de Anolis 
proboscis Peters & Orces (Reptilia: Polychrotidae), 
con el descubrimiento de las hembras de la especie y 
comentarios sobre su distribution y taxonomia. 
Avances en Ciencias e Ingenierias, 2: B7-B15.