Breviora

BREVIORA

Museum of Comparative Zoology

= ait I tha
— sila 2 sae

US ISSN 0006-9698

CAMBRIDGE, Mass. 8 OcTOBER 2010 NUMBER 520

THE ANOLES OF SOROA: ASPECTS OF THEIR ECOLOGICAL RELATIONSHIPS

LOURDES RODRIGUEZ SCHETTINO,! JONATHAN B. Losos,? PAUL E. HERTZ,? KEVIN DE QUEIROZ,*
ADA R. CHAmMiIzo,! MANUEL LEAL,° AND VILMA RIVALTA GONZALEZ!

AsstraAct. Most lizard communities are characterized by having one or two dominant species and a handful of
other species that occur at relatively low densities. However, Soroa, a site in the Sierra del Rosario of western Cuba,
is home to 11 sympatric species of Anolis, of which nine are found in high abundance. In this study, we evaluate how
interspecific differences in structural niche, thermal niche, body size, and behavior might allow the extraordinarily
high anole species diversity at this site. We found that all pairs of species differ in at least one of the following niche
axes: vegetation types occupied, substrates used, perch height, irradiance at occupied perch sites, and body
temperature. Interspecific differences across these axes might serve to reduce competition, allowing the 11 species to
live sympatrically within a relatively small geographic area.

Key worps: Anolis; Cuba; Soroa; community structure

INTRODUCTION
In their pioneering 1969 paper “The anoles

structural and climatic habitats of the seven
species. They also introduced a classification

of La Palma: Aspects of their ecological
relationships,’ Rand and Williams described
how seven sympatric Anolis species partition
habitat in the Cordillera Central of the
Dominican Republic. Following earlier work
by Rand (1964, 1967) and Schoener (1967,
1968), Rand and Williams examined the

of structural habitat use, dividing the species
into six categories that reflected where adult
males engage in most of their activities:
crown anoles, twig anoles, trunk-crown
anoles, trunk anoles, trunk-ground anoles,
and grass-bush anoles. Recognition of these
ecological differences soon led Williams to

‘Instituto de Ecologia y Sistematica, CITMA, La Habana, Cuba; e-mail: zoologia.ies@ama.cu

*Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University,
Cambridge, Massachusetts 02138, U.S.A.; e-mail: jlosos@oeb.harvard.edu

>Department of Biological Sciences, Barnard College, New York, New York, U.S.A.; e-mail: phertz@barnard.edu
-* National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.; e-mail: dequeirozk@si.edu
> Duke University, Durham, North Carolina, U.S.A.; e-mail: mleal@duke.edu

© The President and Fellows of Harvard College 2010.

i)

propose the ecomorph concept: the idea that
species within a community adapt morpho-
logically and behaviorally to using different
parts of the habitat and that, across com-
munities, similar sets of habitat specialists
evolve repeatedly (Williams, 1972, 1983).
Although resource partitioning and _ the
evolution of convergent ecomorphs have
been examined in great detail for three of
the four islands in the Greater Antilles, our
knowledge of Anolis communities in Cuba
has, until recently, been scant (Williams,
1972, 1983; see review in Losos, 2009). Our
goal in this paper is to provide a description
of the diverse anole fauna at a locality in the
Sierra del Rosario, near the town of Soroa in
western Cuba, from the perspective that
Rand and Williams developed. Rand and
Williams actually visited Soroa in October
1981 and June 1983—in the company of
Lourdes Rodriguez Schettino, Julio Novo,
Dale Marcellini, Noel Gonzalez, and Alberto
Coy Otero—and drafted a partial manu-
script that was never completed. We include
a copy of this manuscript as online supple-
mentary material.

Here, we report on parts of the lizard
community at Soroa, which includes 25
lizard species, 24 of which are diurnal
(Rodriguez Schettino et al., 2005). Remark-
ably, 11 of these species belong to the clade
Anolis; we include within Anolis the subclade
Chamaeleolis (Hass et al., 1993; Jackman et
al., 1999), which has traditionally been
treated as a separate genus (e.g., Williams,
1976). Thus, at the time we conducted our
study, Soroa was home to the highest
diversity of anoles yet discovered. More
recently, Diaz et al. (1998) and Garrido and
Hedges (2001) have described sites in eastern
Cuba that house 15 and 14 anole species,
respectively; some sites in Central America
might be equally species rich (Losos, 2009).
To date, however, none of these communities
has received detailed ecological study.

BREVIORA

No. 520

The literature on resource partitioning in
tropical lizard communities suggests that the
use of different perch sites and microhabitats
with different irradiance and thermal proper-
ties often allows sympatric species to avoid
intense interspecific competition (Schoener
and Gorman, 1968; Schoener, 1977; Heat-
wole, 1982; Williams, 1983; Jenssen et al,
1988; Hertz, 1991; Losos, 1992; Powell et al.,
1996; Leal and Fleishman, 2002). Martinez
Reyes (1995) found 11 anole species at Soroa
and noted that most of them perched on tree
trunks or shrubs. In a field trip undertaken in
May and June 1997, the authors of the current
contribution collected data on the type,
height, and diameter of the perches selected
for all 11 species in early summer. After
subjecting these data to a principal compo-
nents analysis, we published a short paper on
niche lability in this lizard community (Losos
et al., 2003). However, neither of the afore-
mentioned contributions presented the raw
data on which the analyses were based. In the
current paper, we provide qualitative and
quantitative descriptions of the natural histo-
ry and habitat use of the anole species at Soroa
to characterize more thoroughly the compo-
sition and structure of the community. Our
observations should be useful as baseline data
for detecting potential changes in the natural
history of these species as climate change over
the coming decades alters the environment
they occupy (Rodriguez Schettino and Rivalta
Gonzalez, 2007; Huey et al., 2009).

MATERIALS AND METHODS

The locality we identify as Soroa encom-
passes the grounds of the Horizontes Hotel
and the forested area at 700-800 m elevation
adjacent to a pathway along the Manantiales
River, ending at the Arco Iris waterfall. It is
located at 22°48'5”N, 83°0'53”W (WGS84)
near the western limit of the Biosphere
Reserve “Sierra del Rosario” in the eastern-

2010

most part of the Cordillera de Guaniguanico,
Pinar del Rio province, Cuba. Soroa has the
highest species richness of vertebrates of any
locality in the Reserve, mainly in_ the
evergreen and gallery forests (Rodriguez
Schettino et al, 1999).

Macrohabitat Types

Because Soroa is an actively used resort,
the site includes buildings, shrubby “living
fences,” chain-link fencing supported by
metal posts, and various other human
constructions (largely concrete) that some
Anolis species use as perch sites. The study
site also includes a modest patch of evenly
planted Ocuje trees (Callophylum antillarum),
as well as the three following natural
vegetation types described by Capote (1989).

Evergreen forest. The canopy layer has
trees 20-30 m tall. The most abundant
species are Erythrina poeppigiana (bucare),
Zanthoxilum martinicense (ayia), Prunus
occidentalis (cuajani), Hibiscus elatus (maja-
gua), C. antillarum (ocuje), Ficus aurea
(jaguey), Cedrela odorata (cedro), Bursera
simaruba (almacigo), Pseudolmedia spuria
(macagua), Oxandra lanceolata (yaya), Ma-
tayba apetala (macuriye), Roystonea regia
(palma royal), and Syzygium jambos (pomar-
ossa). The understory layer is 3-5 m in
height; the most abundant understory species
are Piper aduncum (platanillo de Cuba) and
Psychotria horizontalis (dagame cimarron).

Gallery forest. The canopy layer has trees of
15-20 m, with many lianas and grasses; the most
abundant species are S. jambos (pomarossa), C.
antillarum (ocuje), Lonchocarpus dominguensis
(guama), and R. regia (palma royal).

Secondary vegetation. There are canopy,
understory, and grass layers with many
individuals of H. elatus (majagua), R. regia
(palma royal), Cecropia peltata (yagruma),
- Muntingia calabura (capuli), Z. martinicense
(ayua), and Solanum umbellatum (pendejera).

THE ANOLES OF SOROA 3

Lizard Surveys

During our surveys, we searched for lizards by
walking slowly through disturbed habitats
adjacent to the hotel (buildings, fences, and a
planted woodland) and through a_ nearby
forested area (from the head of the trail across
the road from the hotel to the waterfall at the
bottom of that trail). We sampled lizards in
whatever habitats they occupied; for some
species, we report data separately for lizards
that occupied natural habitats and those that
lived on or around human constructions.
Whenever possible, we report data on males
and females separately; however, it was not
always possible to identify the sex of individual
Anolis barbatus and Anolis luteogularis observed
from a distance because these species exhibit
little sexual size dimorphism and because both
males and females have large dewlaps.

All observations were made between 0800
and 1800 hours over 19 days (May 21 to June
8, 1997). Because previous studies of West
Indian anoles have shown that perch sites
and sun/shade conditions are the resource
axes that sympatric species most commonly
partition (e.g., Rand, 1964; Schoener, 1974;
Roughgarden et al, 1981), during each
excursion, we collected data on the location
where each lizard was first spotted, including
the type of substrate (trunk, branch [includ-
ing twigs], leaf, vine, rock, ground, or
artificial substrate), the height above ground,
the diameter of the substrate (no measure-
ment for ground, walls, or leaves), and the
light conditions (full sun, full shade, or
filtered sun). Lizards that were moving in
response to our presence when first sighted
were excluded. Some lizards were captured
to obtain their body temperatures with a
quick-reading electronic thermocouple ther-
mometer (0.1°C resolution). We measured
activity by walking transects at 1.5-h inter-
vals throughout 2 days and noting every
lizard observed within the forest, as well as in

4 BREVIORA

disturbed habitats (planted ocuje trees, living
fence, and chain-link fence). For species that
we saw only rarely, the time of activity data
were augmented with observations made on
other days; in some cases, the precise time of
observation was not recorded, and some data
were allotted equally to two adjacent time
periods if they were recorded as exactly the
minute separating the periods. Whenever we
captured a lizard to record its body temper-
ature, we also measured its snout—vent length
(SVL). Losos et al. (2003) provides a more
comprehensive description of the methods
used for these surveys.

We report the frequencies with which lizards
used different substrates (listed above), as well
as the frequencies at which they perched in full
sun, filtered sun, or full shade during sunny
weather. We also report descriptive statistics
for perch heights and diameters, body temper-
atures, and SVLs of lizards sampled.

Behavioral Observations

We located adult male anoles by walking
slowly through the habitat until we found an
apparently undisturbed lizard. Animals were
watched for up to 20 minutes, although some
moved out of sight before the observation
period ended. We used data only from
individuals that we observed for at least
5 minutes and that moved at least five times,
following our previous methods (Losos, 1990).
We calculated moves per minute as the number
of individual movements divided by time.
Moves were classified as jumps, runs, or walks.
To be consistent with previous studies (e.g.,
Losos, 1990), data from only one observer
(JBL) were used to distinguish between runs
and walks. For that reason, percentage of
jumps is calculated on the basis of larger
sample sizes than the other two measures, and
the frequencies of the three movement types do
not sum to 1.0. Percent displaying is the
percentage of time a lizard was dewlapping,

No. 520

head bobbing, or performing some other social
display behavior. Data for A. barbatus are
taken from Leal and Losos (2000); these data
were collected for both males and females,
including some subadults, and do not include
information on moves per minute.

RESULTS
Body Size

The 11 anole species at Soroa vary greatly in
body size (Table 1). Two species—A. angusti-
ceps and A. alutaceus—are notably smaller
than most (mean adult male SVL < 45 mm).
Most species are of intermediate size (mean
adult male SVL = 50-70 mm). Three spe-
cies—A. barbatus, A. luteogularis, and A.
vermiculatus—are substantially larger than
the others (mean adult male SVL > 100 mm).
In most species, males appear to be larger than
females, but our sample sizes for some species
and sex combinations are too small to allow
firm conclusions. The trunk-ground species A.
homolechis and A. sagrei exhibit essentially no
differences in body size among sites.

Macrohabitat Distribution

Some of the 11 Anolis species at Soroa
exhibit substantial differences in the macro-
habitats and forest vegetation types that they
occupy (Table 2). Five species (A. allogus, A.
angusticeps, A. barbatus, A. homolechis, and
A. luteogularis) occur in all three forest types;
two species (A. porcatus and A. sagrei) occupy
two forest types; and four species (A. aluta-
ceus, A. loysianus, A. mestrei, and A. vermi-
culatus) are restricted to just one forest type.

Taking into account all the macrohabitats
available at Soroa (Table 2), A. homolechis is
the most widely distributed species, occurring
in disturbed habitats (e.g., the planted wood-
lands and living fences) and open forest edges;
however, it does not occupy chain-link fencing
(not distinguished as a separate category in

2010

TABLE |.

THE ANOLES OF SOROA 5

SNOUT-VENT LENGTH (SVL) OF 11 ANOLIS SPECIES FROM SOROA BY SITE AND SEX. LIZARDS WERE MEASURED AT

THE MOMENT OF CAPTURE.

Anolis Species Site N Mean
A. alutaceus Forest 3} 3917
A. angusticeps Forest —
A. loysianus Forest 5 42.8
A. sagrei Chain-link fence 24 50.6
Living fence 15 50.4
All 39 50.5
A. mestrei Forest 52 52.0
A. allogus Forest 86 52:5
A. homolechis Forest 49 52.0
Living fence 22 522
Planted woodland 26 51.8
All 97 52.0
A. porcatus Forest 2 67.2
A. barbatus Forest 3. 106.7
A. luteogularis Forest 1 183.0
A. vermiculatus Forest 11 =: 121.5

Table 2), the most exposed habitat available at
Soroa. Anolis luteogularis and A. porcatus have
similarly broad habitat distributions (five
macrohabitats each), but the former species
only rarely occurs on or around human
constructions, whereas the latter species is
abundant in that habitat. Anolis sagrei occupies
fairly sunny areas of the forest, and it is
abundant on living fences and human con-

SVL (mm)
Males Females
SD Range N Mean SD Range
7.39 34-48 1] 33
4 392 2.40 32-38
523 40-52. —
4.35 34-55 5 41.6 2.30 38-44
4.29 40-56 —
4.27 34-56 > 41.6 2.30 38-44
4.08 40-60 22 41.6 3.10 35-46
6.70 34-62 20 42.0 2.40 39-46
3.42 45-60 33 41.4 2.39 37-46
2.20 48-55 4 42.3 4.50 40-49
2.30 48-56 3 43.7 2.3) 41-45
2.88 45-60 40 41.7 2.63 37-49
11.70 59-76 —
20.50 83-119 2 131.0 12.73 122-140
6 = 154.7 4.50 149-160
5.56 116-128 13 78.3 6.83 70-90

structions. Four species (A. alutaceus, A.
loysianus, A. mestrei, and A. vermiculatus) have
the narrowest habitat distributions, each being
restricted to just one (forested) habitat type.

Structural Habitats

The anole species at Soroa can be divided
into two groups on the basis of mean perch

TABLE 2. MACROHABITAT DISTRIBUTIONS OF 11 ANOLIS SPECIES AT SOROA. FOREST VEGETATION TYPES FOLLOW
CAPOTE (1989).

Forest Vegetation

Anolis Species Evergreen

. allogus x x
. alutaceus

. angusticeps
. barbatus

. homolechis
loysianus

. luteogularis
. mestrei

. porcatus

. Sagrel

. vermiculatus

Gallery

AR AAA AA AA AA
mK mK mK OM
xx KM KKK MX

mm KM

Secondary Planted Woodland Living Fences

Artificial Structures

x x x

x x

x x x
x x

6 BREVIORA

TABLE 3.

No. 520

FREQUENCIES OF SUBSTRATE USE BY 11 ANOLIS SPECIES AT SOROA. DATA FOR MALES AND FEMALES OF A.

LUTEOGULARIS AND A. BARBATUS ARE COMBINED BECAUSE IT IS NOT POSSIBLE TO DETERMINE THEIR SEX AT A DISTANCE.

Anolis Species N Trunk
Lizards sampled in natural areas
A. allogus o 139 0.56
A. allogus 9 20 0.45
A. alutaceus 0 23 0.22
A. alutaceus 9 8 0.25
A. angusticeps & 4 0.50
A. angusticeps Q 8 0.38
A. barbatus 9 0.56
A. homolechis & 83 0.52
A. homolechis 9 36 0.39
A. loysianus 0 11 0.91
A. loysianus 9 5 1.00
A. luteogularis 28 0.39
A. mestrei 0 96 0.25
A. mestrei Q 22 0.14
A. porcatus 9 8 0.38
A. porcatus 9 6 0.83
A. sagrei % 45 0.18
A. sagrei 9 5 0.40
A. vermiculatus 0 18 0.44
A. vermiculatus 9 17 0.12
Lizards sampled around human

constructions

A. homolechis & 79 0.27
A. homolechis Q 36 0.25
A. porcatus 0 13 0.62
A. porcatus Q 6

A. sagrei Oo 44 0.09
A. sagrei 9 7

Frequency
Branch Leaf Vine Rock Ground Artificial
0.08 0.04 0.25 0.04 0.04
0.15 0.10 0.10 0.10 0.10
0.35 0.17 0.13 0.13
0.26 0.13 0.13 0.25
0.50
0.50 0.13
0.44
0.19 0.01 0.12 0.07 0.08
0.17 0.06 0.11 0.14 0.14
0.09
0.39 0.04 0.11 0.04 0.04
0.08 0.07 0.50 0.05 0.04
0.09 0.68 0.09
0.38 0.25
0.17
0.07 0.02 0.07 0.67
0.60

0.22 0.28 0.06
0.06 0.71 0.12
0.03 0.01 0.04 0.66

0.03 0.03 0.69
0.08 0.15 0.15
0.17 0.67 0.17
0.05 0.02 0.07 0.02 0.75
0.28 0.14 0.57

heights of males (Tables 3, 4a): those that
perch closer to the ground (on average,
< 2 m) and those that perch higher in the
vegetation (on average, = 3 m). Perch heights
of females are similar to those of males in some
species, but quite different in others (Table 4a).

Three species that use lower perches—A.
allogus, A. homolechis, and A. mestreitend
to use broad natural surfaces, such as tree
trunks or rock walls; A. mestrei, in particu-
lar, is always found on or near rock walls in
the forest. Anolis sagrei, a trunk-ground
anole like the other three, frequently perches
on human constructions, primarily walls or

fences. Another species found low to the
ground is A. alutaceus, which usually perches
on narrow vegetation, including thin branch-
es, vines, and ferns. Anolis vermiculatus was
almost invariably found within 5 m of
streams (maximum reported distance is
15 m; Rodriguez Schettino et al, 1987),
often perching as high as 4 m on trunks, on
branches that hung over the water, or on
rocks or walls along the streamside.

The remaining species were normally
found higher in the vegetation, at or above
3 m. Four species (A. angusticeps, A.
loysianus, A. barbatus, and A. luteogularis)

2010

TABLE 4A.

THE ANOLES OF SOROA 1

PERCH HEIGHTS FOR 11 ANOLIS SPECIES AT SOROA. SPECIES ARE LISTED IN ASCENDING ORDER OF MEAN PERCH

HEIGHTS OF MALES. DATA FOR MALE AND FEMALE A. LUTEOGULARIS ARE COMBINED BECAUSE THE SEXES CANNOT BE
DISTINGUISHED FROM A DISTANCE. DATA FOR A. BARBATUS ARE FROM SIX INDIVIDUALS, SOME OF WHICH PROBABLY WERE
SUBADULTS, AND REPRESENT THE MEAN OF THE MIDDLE OF THE HEIGHT RANGES USED BY EACH INDIVIDUAL (DATA FROM LEAL
AND Losos, 2000); RANGES ARE THE LOWEST AND HIGHEST POINT ACROSS ALL INDIVIDUALS. GIVEN THE COMPOSITE NATURE OF
THESE VALUES, NO STANDARD DEVIATIONS ARE REPORTED.

Anolis Species N
Lizards sampled in natural areas

A. sagrei 46
A. allogus 139
A. mestrei 93
A. homolechis 84
A. alutaceus 35
A. vermiculatus 17
A. barbatus 3
A. loysianus 11
A. luteogularis (0 + 9) 28
A. angusticeps 3
A. porcatus 8

Lizards sampled around human constructions

A. homolechis 158
A. sagrei 88
A. porcatus 13

spend most of their time on trunks, branches,
or both in the subcanopy and canopy. Anolis
porcatus, which frequently perches on trunks
and branches and occasionally on leaves,
occupies substantially higher perches in the
forest, often near the canopy, than it does
around human constructions. By contrast,
we found A. angusticeps primarily on
branches, whereas A. barbatus occupies
trunks and branches with nearly equal
frequency. With regard to the diameter of
the perches used, the average size of sub-
strates used by most species was moderate (in
the range of 10-18 cm; Table 4b). Two
species, A. alutaceus and A. barbatus, used
much narrower supports; A. mestrei also
used narrow perches when it perched on
vegetation. Surprisingly, A. angusticeps,
commonly categorized as a twig anole
(Irschick and Losos 1996), used relatively

Perch Diameter (mm)

Males Females

Mean SD Range N Mean SD Range
0.9 0.4 0.2-1.9 5 0.9 0.6 0.3-1.8
1.0 0.6 0.0-3.0 16 0.5 0.4 0.1-1.5
1.4 1.1 0.0-7.0 21 0.8 0.9 0.0-4.0
1.5 2.4 0.0-17.0 23 0.6 0.5 0.1-2.5
1.5 1.1 0.2-5.0 10 2D 1.9 0.3-6.0
1.6 1.3 0.0-4.0 10 0.5 0.7 0.0-1.5
2.8 — 0.5-4.0 3 5 — 03-19
3.1 Sy ees) 6 1.6 1.1 0.3-3.0
5.9 5.1 0.3-20.0
6.3 3.8 2.0-9.0 9 Dep) 1 1.0-4.0
12.0 9.7 3.0-30.0 7 7.0 4.7 1.0-14.0
1.0 0.8 0.1-9.0 55) 0.6 0.4 0.0-1.5
1.0 0.6 0.0-1.8 23 0.5 0.6 0.0-1.8
2.0 1.1 1.0—-5.0 6 2.4 1.4 1.5=5.0

large supports on average, although it was
sometimes seen on narrow surfaces (dis-
cussed further below). The three species that
were commonly observed in both natural
areas and around human constructions used
similar perches in both areas.

Climatic Habitat and Body Temperatures

The 11 Anolis species at Soroa also differ
in their use of perch sites in full sun, filtered
sun, or shade (Table 5). Within the forest, six
species (A. alutaceus, A. angusticeps, A.
barbatus, A. loysianus, A. luteogularis, and
A. mestrei) perched primarily in fully shaded
microhabitats (> 60% of observations). The
remaining species used a more even mix of
sunny and shaded perch sites. Notably, only
one species, A. porcatus, spent a majority of
its time perched in full sun. All three species
that we routinely sampled in the warmer

8 BREVIORA

TABLE 4B.

No. 520

DIAMETERS OF PERCHES (ROCK AND GROUND PERCHES EXCLUDED) USED BY 11 ANOLIS SPECIES AT SOROA. SPECIES

LISTED AS IN TABLE 4A. DATA FOR MALE AND FEMALE A. LUTEOGULARIS ARE COMBINED BECAUSE THE SEXES CANNOT BE
DISTINGUISHED FROM A DISTANCE. DATA FOR A. BARBATUS ARE FROM SIX INDIVIDUALS, SOME OF WHICH PROBABLY WERE
SUBADULTS AND REPRESENT ALL SURFACES USED DURING BEHAVIORAL OBSERVATIONS; THE SAMPLE SIZES REPORTED ARE
NUMBER OF OBSERVATIONS, NOT NUMBER OF INDIVIDUALS (DATA FROM LEAL AND Losos [2000]; No STANDARD DEVIATION
BECAUSE THE VALUES ARE BASED ON WEIGHTED MEANS FOR INDIVIDUALS). SAMPLE SIZES FOR DIAMETER ARE LOWER THAN FOR
PERCH HEIGHT BECAUSE DIAMETER WAS NOT RECORDED FOR LIZARDS ON THE GROUND, ON WALLS, OR ON LEAVES.

Anolis Species N
Lizards sampled in natural areas
A. sagrei 37
A. allogus 139
A. mestrei 40
A. homolechis 64
A. alutaceus 28
A. vermiculatus 12
A. barbatus 27
A. loysianus 11
A. luteogularis (0 + 9) 32
A. angusticeps 4
A. porcatus 7
Lizards sampled around human constructions
A. homolechis 149
A. sagrei 76
A. porcatus 9

habitats around human constructions (A.
homolechis, A. porcatus, and A. sagrei) used a
mixture of sunny and shaded perch sites.

Mean body temperatures divide the 11
species into three groupings that reflect their
macrohabitat preferences and their structur-
al and thermal niches (Table 6). Among the
species sampled in natural areas, only A.
sagrei exhibited mean body temperatures
> 30°C; three species (A. allogus, A. barba-
tus, and A. mestrei) had mean body temper-
atures < 28°C; and six species (A. alutaceus,
A. angusticeps, A. homolechis, A. loysianus,
A. luteogularis, and A. vermiculatus) exhibit-
ed mean body temperatures within a narrow
intermediate range (28.7—29.6°C).

Among the three species sampled around
human constructions, mean body temperatures
were slightly lower than those recorded in

Perch Diameter (mm)

Males Females
Mean SD Range N Mean SD Range
6.2 4.5 0.2-20 4 6.3 7.1 0.2-16
11.4 139 05-80 13 100 12.7 0.439
10 12.9 1.0-50 5 11.0 7.6 2.5-22
12.1 13.2 0.6-70 26 109 13.4 0.7-60
1.1 1.4 0.2-7.0 7 2.1 2.8 0.3-8.0
14.6 17.5 1.5-67 3 10.0 13.0 2.0-25
1.7 —- 0.54.0 27 2.0 — 0.3-19
18: Jae 47 = 210=50 O27 ay 2057, 8-60
12.5 12.7 1.0-50
10.6 9.9 2.5-25 9 14.1 12.0 0.8-30
11.4 9.7 1.0-20 7 146 10.6 5.0-33
15.8 94 1.3-60 52 13.1 6.8 1.3-30
6.6 4.8 0.5-25 11 5.0 4.2 0.5-15
11.9 11.3 1.5-40 4 6.6 5.6 0.5-14.0

natural areas for A. homolechis and A. sagrei;
A. porcatus, which was not sampled in the
forest, had a very high mean body temperature.

Activity Times

Activity times varied among species in
concert with their thermal biology (Table 7).
Among the four trunk-ground anoles in the
forest, the two shade-dwelling species with low
body temperatures (A. allogus and A. mestrei)
were most active from early morning through
midafternoon. By contrast, the heliothermic
species with high body temperature (A. sagrei)
was most active from midday through late
afternoon, and the most broadly distributed
species (A. homolechis) exhibited a fairly con-
sistent level of activity from early morning until
sundown. For most of the other forest-dwelling
species, activity also peaked in midafternoon.

2010

TABLE 5.

THE ANOLES OF SOROA 9

FREQUENCY AT WHICH 11 ANOLIS SPECIES AT SOROA PERCHED IN SUN, FILTERED SUN, OR SHADE DURING SUNNY

WEATHER. DATA FOR MALE AND FEMALE A. LUTEOGULARIS AND A. BARBATUS ARE COMBINED BECAUSE THE SEXES CANNOT BE
DISTINGUISHED FROM A DISTANCE.

Frequency
Males Females
Full Filtered Full Full Filtered Full
Anolis Species N Shade Sun Sun N Shade Sun Sun
Lizards sampled in natural areas
A. allogus 65 0.43 0.43 0.14 9 0.56 0.33 0.11
A. alutaceus 12 1.00 3 0.33 0.33 0.33
A. angusticeps 1 1.00 5 0.80 0.20
A. barbatus 6 0.83 0.17
A. homolechis D2 0.21 0.33 0.46 20 0.35 0.30 0.35
A. loysianus 9 0.67 0.22 0.11 4 1.00
A. luteogularis 19 0.63 0.26 0.11
A. mestrei 79 0.66 0.23 0.11 16 0.69 0.31
A. porcatus 3 0.33 0.67 3 0.33 0.67
A. sagrei 38 0.58 0.03 0.40 2 1.00
A. vermiculatus 12 0.17 0.58 0.25 17 0.24 0.59 0.18
Lizards sampled around human

constructions
A. homolechis 94 0.37 0.39 0.23 17 0.53 0.35 0.12
A. porcatus 9 0.78 0.22 4 0.25 0.75
A. sagrei 4] 0.42 0.44 0.15 6 0.33 0.67

In the areas sampled outside the forest, both
species that occupy open habitats (A. porcatus
and A. sagrei) were active in the morning, an
unsurprising result, given that these habitats
heat up much earlier in the day and reach
higher temperatures than do locations within
the forest. By contrast, A. homolechis showed
high levels of activity over a narrower range of
times (midday) than it does in the forest.

Behavior

The species varied fivefold in their rates of
movement (Table 8). The more terrestrial
species tended to be relatively inactive,
whereas some of the more arboreal species
(A. angusticeps, A. loysianus, and A. porcatus)
moved at much higher rates, observations
that parallel those seen for anoles on other
islands in the Greater Antilles (Johnson et al.,
2008). Anolis alutaceus jumped twice as much
~ as any other species. Trunk-ground anoles (A.

allogus, A. homolechis, A. mestrei, and A.
sagrei) and A. loysianus tended to run more
than other species, whereas the twig (A.
angusticeps) and large arboreal species (A.
luteogularis), as well as A. alutaceus, ran
relatively infrequently. Anolis angusticeps
and A. barbatus walked considerably more
often than other species. Percentage of time
spent displaying also varied greatly among
species, with trunk-ground anoles, A. barba-
tus, and A. loysianus displaying more fre-
quently than other species.

Complementarity of Resource Use
among Species

At Soroa, species that are ecologically
similar in one niche dimension often differ in
another. For example, A. allogus and A.
homolechis perch low on trunks (Table 4a) in
all three types of forest vegetation (Table 2).
Although they are similar in size (Table 1)

10 BREVIORA

No. 520

TABLE 6. BODY TEMPERATURES FOR I|1 ANOLIS SPECIES AT SOROA. SPECIES ARE LISTED IN DESCENDING ORDER OF MALE
MEAN BODY TEMPERATURES.

Males
Anolis Species N Mean SD
Lizards sampled in natural areas
A, sagrei 9 32.1 1.8
A, vermiculatus 7 29.6 1.7
A. luteogularis 8 29.4 1.1
A. loysianus 3 29.3 1.6
A. angusticeps —
A. homolechis 51 29.2 2.4
A. alutaceus 28.7 1.3
A. mestrei 53 27.5 1.9
A. barbatus 3 Dips 0.2
A. allogus 82 26.9 Pep
Lizards sampled around human
constructions
A. porcatus 3 32.7 2.19
A, sagrei 87 30.4 3.36
A. homolechis 133 28.5 2.02

and exhibit similar patterns of substrate use
(Table 3), A. homolechis occupies sunnier
areas of the forest (Table 5), and this
behavioral difference is reflected in a differ-
ence in their mean body temperatures
(Table 6). A third species, A. mestrei, which
is similar in size and habitus to the other two,
also perches low in the gallery forest, but it
frequently uses rock faces, which differenti-
ates it ecologically from the other two
species. Finally, A. sagrei, which is similar
in size and closely related to the three
aforementioned species, uses similar perch
heights and substrates, but it is most
abundant in nonforested habitats, and its
mean body temperature is among the highest
of all the anoles we sampled at Soroa.

A different pattern of ecological differen-
tiation is seen among the three species that
occupy high, relatively narrow perches with-
in the forest (A. angusticeps, A. barbatus, and
A. luteogularis): A. angusticeps is much
smaller than the other two species, which

Body Temperature (°C)

Females
Range N Mean SD Range

29.6-34.9 5 B29, 1.8 31.6-33.9
26.7—32.0 9 29.8 1.8 27.0-32.1
28.0-30.7 _
27.7-30.8 2 28.6 0.3 28.4-28.8

3 29) 0.4 28.7-29.4
24.2-33.5 34 28.7 2.5 22.3-33.4
27.6-30.6 3 29.1 1.0 28.0-29.9
22.8-30.8 23 27.6 2.0 22.1-29.9
27.1-27.5 —
21.3-31.1 18 26.9 IES. 24.2-29.9
30.8-35.1 7 31.5 1.4 29.8-33.3
24.5-36.8 16 28.2 2.1 24.4-30.9
24.1-32.5 54 27.6 2.1 24.6-32.7

also differ substantially in size; these differ-
ences in SVL almost certainly enable the
species to consume different foods (Schoener
1967). Indeed, we observed only A. luteogu-
laris feeding on smaller species of anoles.

NATURAL HISTORY OF
SOROA ANOLES

In this section we briefly describe our
natural history observations on the 11 Anolis
species at Soroa. These descriptions include
information about the colors of their dewlaps
because previous research has established that
the diversity of Anolis dewlap patterns and
behavioral displays, as well as redundancy in
dewlap information content, allows them to
communicate species identity unambiguously
(Rand and Williams 1970; Losos and Chu
1998; Nicholson et al. 2007). Our observa-
tions of syntopic species (e.g., the four trunk-
ground anoles) at Soroa are consistent with
this hypothesis. All photographs are males
and, except where noted, from Soroa.

2010 THE ANOLES OF SOROA 1]

TABLE 7. RELATIVE ACTIVITY TIMES FOR 11 ANOLIS SPECIES AT SOROA. FOR EACH SPECIES, ENTRIES IN THE TABLE RECORD
THE PERCENTAGE OF ALL LIZARDS OBSERVED (N) THAT WERE SPOTTED DURING EACH OF EIGHT TIME PERIODS. CENSUS DATA
WERE COLLECTED WHILE WALKING TRANSECTS ON EACH OF TWO DAYS. DATA WERE AUGMENTED BY OBSERVATIONS MADE ON
OTHER DAYS FOR SPECIES LESS COMMONLY SEEN; FOR THESE OBSERVATIONS, SOME POINTS WERE ALLOTTED EQUALLY TO TWO

ADJACENT TIME PERIODS IF THEY WERE RECORDED AS EXACTLY THE MINUTE SEPARATING THE PERIODS.

Relative Activity Time
0700— 0830— 1000— 1130— 1300—- 1430- 1600- = 1730-

Anolis Species N 0830 1000 1130 1300 1430 1600 1730 1900

Lizards sampled in the forest

A. allogus 161 0.08 0.09 0.15 0.21 0.15 0.12 0.12 0.08

A. alutaceus 44 0.02 0.16 0.09 0.27 0.21 0.07 0.18

A. angusticeps 10 0.35 0.45 0.20

A. barbatus 5 0.20 0.20 0.60

A. homolechis 119 0.03 0.08 0.17 0.13 0.15 0.18 0.16 0.10

A. loysianus 20 0.15 0.45 0.20 0.15 0.05

A. luteogularis 31 0.18 0.23 0.21 0.19 0.16 0.03

A. mestrei 119 0.03 0.08 0.13 0.24 0.18 0.13 0.11 0.11

A. porcatus 15 0.20 0.07 0.03 0.50 0.13 0.07

A. sagrei 5) 0.07 0.39 0.04 0.26 0.19 0.05

A. vermiculatus 8 0.50 0.44 0.06
Lizards sampled outside of the forest

A. homolechis 210 0.07 0.17 0.40 0.13 0.21 0.03 0.01

A. porcatus 21 0.05 0.57 0.10 0.10 0.10 0.10

A. sagrei 114 0.04 O11 0.20 0.23 0.09 0.17 0.17 0.00

TABLE 8. AVERAGE MOVES PER MINUTE, FREQUENCIES OF THREE LOCOMOTOR BEHAVIORS, AND FREQUENCY OF DISPLAY
BEHAVIOR IN 11 ANOLIS SPECIES AT SOROA. SAMPLE SIZE IS THE NUMBER OF INDIVIDUAL LIZARDS OBSERVED. ANIMALS THAT
MOVED < 5 TIMES WERE NOT INCLUDED IN THE ANALYSIS OF THE FREQUENCIES OF THE LOCOMOTOR BEHAVIORS. IN ADDITION,
ONLY INDIVIDUALS OBSERVED BY JBL WERE INCLUDED IN CALCULATIONS OF PERCENTAGES OF MOVES THAT WERE RUNS VERSUS
WALKS TO PROVIDE CONSISTENCY WITH PREVIOUS RESEARCH. AS A RESULT, THE FREQUENCIES OF RUNS, WALKS, AND JUMPS DO
NOT SUM TO 1.0. DATA WERE COLLECTED ONLY FROM ADULT MALES, EXCEPT FOR A. BARBATUS, FOR WHICH DATA (FROM LEAL
AND Losos, 2000) WERE COLLECTED FROM SUBADULT AND ADULT MALES AND FEMALES; PERCENT TIME DISPLAYING FOR THIS
SPECIES IS BASED ON TWO SMALL MALES. DATA ON MOVEMENTS PER MINUTE FOR ALL SPECIES EXCEPT A. VERMICULATUS WERE

PREVIOUSLY PUBLISHED IN JOHNSON ET AL. (2008).

Anolis Species N Moves/Min Jump Run Walk Display

A. allogus 15 0.51 0.14 0.46 0.07 0.037
A. alutaceus 8 0.64 0.56 0.13 0.35 0.008
A. angusticeps D 2.49 0.09 0.09 0.82 0.020
A. barbatus 7 — ~ 0.001 ~ 0.10 0.89 0.078
A. homolechis 21 0.67 0.24 0.44 0.45 0.075
A. loysianus 6 1.83 0.06 0.41 0.52 0.065
A. luteogularis 6 0.62 0.14 0.15 0.72 0.021
A. mestrei 15 0.59 0.28 0.27 0.45 0.068
A. porcatus 11 1.52 0.23 0.21 0.59 0.037
A. sagrei 19 0.57 0.19 0.27 0.49 0.125
A. vermiculatus 7 0.88 0.23 0.22 0.55 0.023

12 BREVIORA

Anolis allogus

Photo by Kevin de Queiroz.

This species fits the classic definition of
a trunk-ground anole: a stocky lizard with
long hindlimbs, often found perching on
broad surfaces (usually tree trunks) rela-
tively close to the ground. Anolis allogus is
found primarily in deep forest and has a
lower body temperature than some of the
other trunk-ground anoles. It is one of
four sympatric trunk-ground species at
Soroa (the others being A. homolechis, A.
mestrei, and A. sagrei), which differ in the
color of the dewlap and, to a lesser extent,
of the body. The dewlap pattern of A.
allogus exhibits a significant amount of
geographic variation across Cuba. At
Soroa, the dewlap is primarily yellow, with
three to four well-delimited transverse
brick-colored bars. The bars are usually
located on the upper to middle region of
the dewlap.

No. 520

Anolis alutaceus

Photos courtesy of Luke Mahler.

This species is a small anole usually
found on narrow diameter substrates, such
as bushes, ferns, or vines, from near the
ground to several meters high. The species
moves by slow walks and fast jumps. Like
most grass-bush anoles, it sports an ex-
traordinarily long tail, as much as 2.5 times

2010

the SVL, as well as long hind limbs. Its
irises are blue, and its dewlap is a solid
yellow.

Anolis angusticeps

Lizard photograph by Kevin de Queiroz; dewlap
photograph courtesy of Luke Mahler.

Small and with short legs, this species fits
the classic morphological description of a
twig anole. Anolis angusticeps has been
studied intensively in the Bahamas, where it
primarily uses narrow-diameter vegetation

THE ANOLES OF SOROA 13

(Schoener, 1968; Irschick and Losos, 1996).
However, despite our extensive searching of
narrow surfaces, in 30.4% of our observa-
tions it perched on trunks and in relatively
few observations was it found on narrow
surfaces. Whether these observations repre-
sent a true difference between Cuban and
Bahamian populations of this species, or
whether we simply failed to observed these
lizards on narrow surfaces—perhaps out of
sight high in the canopy—remains to be
determined. Like other twig anoles, the A.
angusticeps we observed moved slowly,
rarely running or jumping (except when
fleeing perceived predators). The dewlap of
this species is peach in color.

Anolis barbatus

Photos courtesy of Luis Diaz.

A member of the Chamaeleolis subclade
of anoles, A. barbatus is large and heavy-
bodied. It is a slow-moving animal that
often adopts a rocking motion, much like

14 BREVIORA

that seen in chameleons. It is usually found
high in trees, but descends to the ground to
feed on hard-bodied prey such as mollusks
and beetle pupae. This species is often found
on surfaces that are relatively narrow for its
large size; for this and other reasons, such as
the rarity of jumping or running, it may be
considered a large twig anole (Leal and
Losos, 2000; Losos, 2009). Observations on
this species at Soroa were reported in Leal
and Losos (2000). The dewlap of A.
barbatus is white with a pink- to peach-
colored edge. Males and females have
dewlaps of equivalent size, an unusual
characteristic in Anolis that is shared with
A. luteogularis.

Anolis homolechis

Photo by Kevin de Queiroz.

This trunk-ground anole is found in light
shade in the forest and in patches of
woodland and vegetated perches in open
areas. It has a lower body temperature than
A. sagrei, but a higher body temperature
than the deep-forest trunk-ground anoles A.
allogus and A. mestrei. In all respects, it is a
typical trunk-ground anole, usually perch-
ing close to the ground on tree trunks or
other broad structures. Anolis homolechis
has a solid white dewlap that is brighter
than that of A. barbatus. On close exami-
nation, the dewlap of A. homolechis has
numerous small black dots between the
scales, most likely pockets of melanin. In

No. 520

some populations, the dewlap appears gray,
possibly because of a higher melanin con-
centration.

Anolis loysianus

Lizard photograph by Jonathan Losos; dewlap photo
courtesy of Luis Diaz.

This small-bodied species is found only on
very broad trunks with rugose bark. Like the
trunk anoles of Hispaniola, the only other
Greater Antillean Island on which the trunk
ecomorph occurs, it moves frequently, rarely
jumps, and sometimes ascends to great
heights. Its crypticity, heightened by fleshy
projections on its body, might also contrib-
ute to its apparent rarity. It has a pale
orange-red dewlap; individual variation is
considerable in the number of orange spots,
which are scattered across the dewlap and
might contribute to the dewlap’s saturated
appearance.

2010

Anolis luteogularis

Lizard photograph by Kevin de Queiroz; dewlap
photograph courtesy of Luke Mahler.

This large-bodied lizard is the longest and
heaviest anole at Soroa. It will eat anything it
can catch and has been observed stalking
other anoles. Normally found high in trees
on broad substrates such as trunks and large
branches, it descends to lower heights to
forage. Most likely, individuals maintain a
large home range that encompasses many
trees. This species generally moves by walk-
ing but will jump or run as necessary. Anolis
luteogularis also shows the most pronounced
response to large avian predators, moving to
the underside of a branch and remaining
immobile for a few minutes when a red-tailed
hawk (Buteo jamaicensis) flies overhead. The

THE ANOLES OF SOROA 15

dewlap is a pale yellow, sometimes so pale
that it almost looks white. Males and females
have dewlaps of equivalent size.

Anolis mestrei

Photo by Kevin de Queiroz.

Morphologically similar to the other
trunk-ground members of the A. sagrei
clade at Soroa, A. mestrei is associated with
large boulders and limestone walls in the
forest. When not found on rocky surfaces, it
is usually found on tree trunks. On all
surfaces, it is found relatively low to the
ground, like other trunk-ground anoles.
This species is only found in deep forest
and, like A. allogus, exhibits relatively low
body temperatures. The dewlap pattern of
A. mestrei exhibits a significant amount of
geographic variation across Cuba. At
Soroa, the dewlap is two-toned, with a
bright white edge and a relatively large
brick-red center. Individual variation on the
size and saturation of the brick-red center is
substantial.

16 BREVIORA

Anolis porcatus

Lizard photograph by Kevin de Queiroz; dewlap
photograph of a specimen from the vicinity of Mariel,
Cuba, courtesy of Luke Mahler.

This trunk-crown anole is commonly
found in human-disturbed habitats, where
it is seen on walls, palms, and other trees,
= 2 m above the ground. This species also
moves frequently on leafy vegetation. Anolis
porcatus is less commonly seen in the forest,
where it is often in the canopy. This species
usually has a high body temperature and is
often seen basking; individuals readily
change their body color between green and
brown. It has a relatively small, one-toned
dewlap with reddish to dirty pink coloration.

No. 520

Anolis sagrei

Lizard photograph by Kevin de Queiroz; dewlap
photograph courtesy of Luke Mahler.

The body temperature of this sun-loving
species is only rivaled at Soroa by that of the
equally sun loving A. porcatus. Anolis sagrei is
extremely common in human-disturbed areas.
Like the other trunk-ground anoles, it is found
near the ground, but it perches on practically
any surface, from broad tree trunks to

2010

relatively narrow chain link fences. It rarely
gets far into the forest, though it might
occasionally be seen in open areas within the
forest. The dewlap pattern of A. sagrei exhibits
substantial geographic variation across Cuba.
At Soroa, the dewlap is a bright reddish-
orange, with prominent yellow scales.

Anolis vermiculatus

Lizard photograph by Jonathan Losos; dewlap photo-
graph courtesy of Luke Mahler.

One of the most remarkable of the more
than 360 anole species, A. vermiculatus is
always found near rivers (Leal et al, 2002),
into which it dives to escape predators or to
pursue prey, including small fish and shrimps.
In a survey of stomach contents, Rodriguez

THE ANOLES OF SOROA V9

Schettino and Novo Rodriguez (1985) found
insects, crustaceans, and plant matter (flowers
and fruits of palms, primarily in the stomachs
of adult males). The species will run bipedally
across the river surface; individuals can be seen
crisscrossing as they chase each other in
territorial disputes. When not in the water,
these lizards perch on vertical rock walls and
on tree branches, from which they dive into the
water if threatened, as well as on rock outcrops
within the stream (Rodriguez Schettino et al,
1987). This is one of two anoles completely
lacking a dewlap (the other is its sister species,
A. bartschi). Individuals use shallow head bobs
to signal to each other; their complex body
coloration might aid in communication (Fitch
and Henderson 1987). The skin of this species
has a velvety texture that might have some
relationship to its aquatic habitats.

DISCUSSION

Natural selection often favors ecological
divergence that minimizes the intensity of
negative interspecific interactions among
sympatric species in a community (Hutch-
ison, 1959; MacArthur, 1972; Losos et al,
2003). The lizards of Soroa likely have a long
history of ecological interactions (Losos et
al., 2003). Our previous work, based on data
reported and elaborated on here, revealed

18 BREVIORA

that species are more dissimilar than expect-
ed by chance; in particular, species that are
similar along one niche axis tend to be
dissimilar along another (Losos et al,
2003). The evolution of ecological differences
among the anoles living in the same com-
munity was predicted by Rand and Williams
(1969) on the basis of their study of another
complex community of Anolis lizards. Wil-
liams (1972) coined the term ‘“‘ecomorph”’
for groups of species with similar morphol-
ogy and ecology; he also proposed six
ecomorphs for Anolis lizards from the
Greater Antilles.

The Anolis community of Soroa is com-
posed of species representing Williams’ six
ecomorph classes (see Losos [2009] for a
review of the ecomorph concept): crown-
giant, A. luteogularis; trunk-crown, A. por-
catus; trunk, A. loysianus; trunk-ground, A.
allogus, A. homolechis, A. mestrei, and A.
sagrei,; twig, A. angusticeps (but see qualifi-
cations above); and grass-bush, A. alutaceus.
Anolis barbatus, although much larger than
other species classified as twig anoles, is
similar to twig anoles in many morphological
and behavioral respects; moreover, it often
uses surfaces that are narrow relative to its
size. For this reason, A. barbatus might be
considered a twig anole (Losos, 2009). Anolis
vermiculatus does not fit neatly into any of
the six standard Greater Antillean ecomorph
categories, although it selects perch sites
similar to those used by _ trunk-ground
anoles. However, other anoles, both in the
Greater Antilles (A. eugenegrahami from
Hispaniola) and on the mainland (a number
of species) are similar to A. vermiculatus in
being found only near streams. The two
Greater Antillean species, however, are not
similar morphologically, and neither is sim-
ilar to those on the mainland. For this
reason, these “‘semiaquatic’’ anoles do not
constitute an ecomorph class sensu Williams
(Leal et al., 2002).

- No. 520

Our results indicate that, regardless of
whether competition is an important factor
in the structuring of the anole communities,
differences in the use of resources among the
species from Soroa are sufficient to allow 11
species to coexist in what seems to be a stable
community.

Comparisons to Previous Work on
These Species

Although the body temperatures of several
Anolis species occurring at Soroa have been
reported previously (Rodriguez Schettino,
1999a), only the data for A. mestrei (Rodriguez
Schettino and Chamizo Lara, 2001) and those
for A. vermiculatus (Gonzalez Bermudez and
Rodriguez Schettino, 1982; Rodriguez Schet-
tino et al., 1987; Rodriguez Schettino and
Martinez Reyes, 1989) had actually been
collected at Soroa, as opposed to elsewhere in
Cuba. Data for A. homolechis and A. allogus
(Silva Rodriguez, 1981) were collected at
nearby locations in the Sierra del Rosario.
Anolis mestrei body temperatures at Soroa in
June 1995 (Rodriguez Schettino and Chamizo
Lara, 2001) were similar to those that we
recorded (means of 28.2°C vs. 27.5°C, respec-
tively). Body temperatures for A. homolechis
elsewhere in the Sierra del Rosario generally
were higher than those reported here (means of
32.1°C vs. 29.2°C, respectively), but the values
were similar for A. allogus (means of 27.5°C vs.
26.9°C, respectively). Anolis vermiculatus body
temperatures were taken by Gonzalez Bermt-
dez and Rodriguez Schettino (1982) during the
dry season, and it is not possible to compare
their data with ours; however, Rodriguez
Schettino et al. (1987) and Rodriguez Schettino
and Martinez Reyes (1989) obtained their data
for June 1983 (mean 26.0°C) and June 1986
(mean 28.6°C), respectively. Both values are
lower than the mean we report (29.4°C),
probably reflecting differences in the weather
among the 3 years in which data were gathered.

2010

Ruibal (1961) reported body temperatures
for three of the species at various sites in
eastern Cuba. He collected data in July and
August, recording mean body temperatures
Oim29 25 tor A. allogus, 3\:8°C for A.
homolechis, and 33.1°C for A. sagrei. These
values are all somewhat higher than the
means for these species reported here; we
assume that the differences reflect seasonal
or geographical variation.

With regard to the vegetation types that the
11 anole species occupy, our results partially
mirror those of Martinez Reyes (1995), al-
though we found several species in a wider
range of habitats than she did. In both studies,
A. homolechis was found in the planted
woodland, as well as in all three forest
vegetation types; A. mestrei and A. vermicula-
tus only in the gallery forest; A. Joysianus only
in the evergreen forest; and A. alutaceus only in
secondary vegetation. However, Martinez
found A. allogus only in the evergreen forest,
whereas we discovered it in all three vegeta-
tions types. Similarly, although Martinez
found A. sagrei only around human construc-
tions, we also found it in the gallery forest. In
addition, Martinez Reyes (1995) found A.
porcatus, A. luteogularis, and A. angusticeps
only in secondary vegetation, whereas we
found the former species also in the gallery
forest and the latter two species in all three
vegetation types. She did not detect A. barbatus
in the gallery forest, whereas we did. Although
Martinez did not find the grass-bush anole
Anolis ophiolepis (Fig. 1) at Soroa, Rodriguez
Schettino et al. (2005) found two individuals in
secondary vegetation, and we found one
individual in grassy secondary vegetation along
a roadside near Soroa while collecting at night.

With regard to substrates used, our results
more closely resemble those of Martinez
Reyes (1995), but we found a wider range
of microhabitat use in some species. She did
not find A. homolechis, A. sagrei, or A.
vermiculatus on rocks, nor did she encounter

THE ANOLES OF SOROA 19

Figure 1. Anolis ophiolepis. Photograph by Kevin de
Queiroz of a specimen collected in Havana, Cuba.

A. homolechis, A. mestrei, or A. allogus on or
around human constructions. In addition,
she found A. alutaceus on grasses, whereas
we only observed this species in the forest,
where it primarily perched on narrow
branches, lianas, and ferns. Martinez Reyes
did not report data on perch height or body
temperature. Our results on perch height
generally coincide with the findings of
Rodriguez Schettino (1999a, b) and Rodri-
guez Schettino and de Queiroz (2002a, b). All
of the species at Soroa were observed in the
sun or shade at frequencies similar to those
reported by Rodriguez Schettino (1999a).

CONCLUSIONS

Research on anoles has played an impor-
tant role in the development of community
ecological theory (e.g., Schoener, 1968, 1974,
1977). Until now, no comprehensive study
had been conducted on Cuban anole com-
munities, even though Cuba hosts the richest
and most diverse anole fauna in the West
Indies. Our data indicate that at least one
Cuban anole community follows the same
patterns documented in less species-rich
communities on other islands in the West
Indies. Nonetheless, further work is needed

20 BREVIORA

on localities elsewhere in Cuba; indeed, little
is known about how anole communities
differ among localities within a single island.
In addition, the anole fauna of mainland
Central and South America is as diverse as
that on the islands, yet many differences exist
between mainland and island anoles (An-
drews, 1979; Pinto et al, 2008; reviewed in
Losos, 2009). Detailed studies of mainland
anole communities could prove very infor-
mative with regard to understanding how
these differences have arisen.

ACKNOWLEDGMENTS

We thank Adela Torres Barboza and
Angel Daniel Alvarez for their assistance
with field work and Luis Diaz and Luke
Mahler for allowing us to use their photo-
graphs. We received funds for this work from
the “Programa Nacional de Ciencia y
Técnica de Los Cambios Globales y la
Evolucion del Medio Ambiente Cubano,”
the National Geographic Society (grants
5639-96 and 6981-01), the National Science
Foundation, and a faculty grant from
Barnard College.

LITERATURE CITED

ANpreEws, R. M. 1979. Evolution of life histories: A
comparison of Anolis lizards from matched island
and mainland habitats. Breviora 454: 1-51.

Capote, R. P. 1989. La vegetacion de la Sierra del
Rosario. Ph.D. Dissertation, La Habana, Cuba,
Instituto de Ecologia y Sistematica.

Diaz, L. M., N. NAVARRO, AND O. H. Garripo. 1998.
Nueva especie de Chamaeleolis (Sauria: Iguanidae)
de la Meseta de Cabo Cruz, Granma, Cuba.
Avicennia 8/9: 27-34.

Fitcu, H. S., AND R. W. HENDERSON. 1987. Ecological
and ethological parameters in Anolis bahorucoensis,
a species having rudimentary development of the
dewlap. Amphibia-Reptilia 8: 69-80.

Garripo, O. H., AND S. B. Hepces. 2001. A new anole
from the northern slope of the Sierra Maestra in
eastern Cuba (Squamata: Iguanidae). Journal of
Herpetology 35: 378-383.

No. 520

GONZALEZ BERMUDEZ, F. M., AND L. RopRiGUEZ
SCHETTINO. 1982. Datos etoecologicos sobre Anolis
vermiculatus (Sauria: Iguanidae). Poeyana 245: 1-18.

Hass, C. A., S. B. HepGes, AND L. R. Maxson. 1993.
Molecular insights into the relationships and
biogeography of West Indian anoline lizards.
Biochemical Systematics and Ecology 21: 97-114.

Heatwote, H. 1982. A review of structuring in
herpetofaunal assemblages. U. S. Fish and Wildlife
Service Wildlife Research Report 13: 1-19.

Hertz, P. E. 1991. Evaluating thermal resource parti-
tioning by sympatric lizards Anolis cooki and A.
cristatellus: A field test using null hypotheses.
Oecologia 90: 127-136.

Huey, R. B., C. Deutscn, J. J. Tewkssury, L. J. Vitt, P.
E. Hertz, H. J. ALVAREZ PEREZ, AND T. GARLAND,
Jr. 2009. Why tropical forest lizards are vulnerable
to climate warming. Proceedings of the Royal
Society B 276: 1939-1948.

Hutcuison, G. E. 1959. Homage to Santa Rosalia or
why are there so many kinds of animals? The
American Naturalist 93: 145-159.

IrscHick, D. J., AND J. B. Losos. 1996. Morphology,
ecology, and behavior of the twig anole, Anolis
angusticeps, pp. 291-301. In R. Powell and R. W.
Henderson (eds.), Contributions to West Indian
Herpetology: A Tribute to Albert Schwartz. Ithaca,
New York, Society for the Study of Amphibians
and Reptiles.

JACKMAN, T., 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.

JENSSEN, T. A., D. L. MARCELLINI, AND E. P. SMITH.
1988. Seasonal micro-distribution of sympatric
Anolis lizards in Haiti. Journal of Herpetology 22:
266-274.

Jounson, M. A., M. LEAL, L. RopRiGUEZ SCHETTINO, A.
Cuamizo Lara, L. J. REVELL, AND J. B. Losos. 2008.
A phylogenetic perspective on foraging mode
evolution and habitat use in West Indian Anolis
lizards. Animal Behaviour 75: 555-563.

LEAL, M., AND L. FLEISHMAN. 2002. Evidence for habitat
partitioning based on adaptation to environmental
light in a pair of sympatric lizard species. Proceed-
ings of the Royal Society of London B 269:
351-359.

, AND J. B. Losos. 2000. Behavior and ecology
of the Cuban “chipojos bobos’” Chamaeleolis
barbatus and C. porcus. Journal of Herpetology
34: 318-322.

———.,, A. K. Knox, anp J. B. Losos. 2002. Lack of
convergence in aquatic Anolis lizards. Evolution 56:
785-791.

2010

Losos, J. B. 1990. Concordant evolution of locomotor
behaviour, display rate, and morphology in Anolis
lizards. Animal Behaviour 39: 879-890.

. 1992. The evolution of convergent structure in

Caribbean Anolis communities. Systematic Biology

41: 403-420.

. 2009. Lizards in an Evolutionary Tree: Ecology

and Adaptive Radiation of Anoles. Berkeley, Cali-

fornia, University of California Press.

, AND L.-R. Cuu. 1998. Examination of factors

potentially affecting dewlap size in Caribbean

anoles. Copeia 1998: 430-438.

, M. Leat, R. E. Gior, K. DE Queiroz, P. E.
Hertz, L. RopriGuez ScHETTINO, A. CHAMIZO
Lara, T. R. JACKMAN, AND A. Larson. 2003. Niche
lability in the evolution of a Caribbean lizard
community. Nature 424: 542-545.

MacArtuur, R. 1972. Geographical Ecology: Patterns
in the Distribution of Species. Princeton, New
Jersey, Princeton University Press.

MartTiNEz Reyes, M. 1995. Saurios de la Reserva de la
Biosfera “Sierra del Rosario” Pinar del Rio, Cuba.
Evaluacion ecologica de tres comunidades. Investi-
gaciones Geograficas Boletin 30: 50-77.

NicHo.son, K. E., L. J. HARMON, AND J. B. Losos. 2007.
Evolution of Anolis lizard dewlap diversity. PLoS
One 2(3): e274.

Pinto, G., D. L. MAHLER, L. J. HARMON, AND J. B.
Losos. 2008. Testing the island effect in adaptive
radiation: rates and patterns of morphological
diversification in Caribbean and mainland Anolis
lizards. Proceedings of the Royal Society B 275:
2749-2757.

PoweELL, R., J. S. PARMERLEE, JR., AND D. D. Situ.
1996. Evidence of spatial niche partitioning
by a Hispaniolan lizard community in a xeric
habitat, pp. 317-326. In R. Powell and R. W.
Henderson (eds.), Contributions to West Indian
Herpetology: A tribute to Albert Schwartz. Ithaca,
New York, Society for the Study of Amphibians
and Reptiles.

Rano, A. S. 1964. Ecological distribution in
anoline lizards of Puerto Rico. Ecology 465:
745-752.

. 1967. Ecology and social organization in the
iguanid lizard Anolis lineatopus. Proceedings of the
United States National Museum 122: 1-79.
, AND E. E. WILLIAMS. 1969. The anoles of La
Palma: aspects of their ecological relationships.
Breviora 327: 1-17.

, AND . 1970. An estimation of redundancy
and information content of anole dewlaps. The
American Naturalist 104: 99-103.

THE ANOLES OF SOROA 21

RopDRiGUEZ SCHETTINO, L. 1999a. Systematic accounts of
the species, pp. 104-380. In L. Rodriguez Schettino
(ed.), The Iguanid Lizards of Cuba. Gainesville,
Florida, University of Florida Press.

1999b. Morfometria, distribucion geografica y

microhabitat de los lagartos del género Anolis

(Lepidoasuria: Iguania). Ph.D. Dissertation. La

Habana, Cuba, Instituto de Ecologia y Sistematica.

, AND A. CHAmIzo LARA. 2001. Anolis mestrei

(NCN). Thermal habitat. Herpetological Review

32: 253.

, V. RIVALTA GONZALEZ, C. MANCINA
GONZALEZ, AND A. HERNANDEZ MArRERO. 2005.
Herpetofauna de la Reserva de la Biosfera ‘‘Sierra
del Rosario”, Cuba. Poeyana 493: 12-16.

, AND K. DE QuetrRoz. 2002a. Anolis homolechis

homolechis (NCN). Thermal biology and microhab-

itat. Herpetological Review 33: 304.

, AND . 2002b. Anolis sagrei sagrei (Cuban
brown lizard; Chino; Torito). Habitat use and
thermal biology. Herpetological Review 33: 305.

, C. A. MANCINA GONZALEZ, E. PEREZ MENA, A.

CHAMIzO Lara, A. MARTELL GaArciA, AND A.

HERNANDEZ Marrero. 1999. Bases para el futuro

manejo y conservacion de los vertebrados terrestres

de la Reserva de la Biosfera Sierra del Rosario.

Scientific Report, La Habana, Cuba, Instituto de

Ecologia y Sistematica.

, D. L. MARCELLINI, AND J. Novo RopriGuEz.
1987. Algunos aspectos ecoldgicos sobre Anolis
vermiculatus (Sauria: Iguanidae) en Soroa, Pinar del
Rio, Cuba. Poeyana 343: 1-9.

—, AND M. Martinez Reyes. 1989. Algunos
aspectos ecologicos sobre cuatro especies endémicas
del género Anolis (Sauria: Iguanidae). Scientific
Report, La Habana, Cuba, Instituto de Ecologia y
Sistematica.

, AND J. Novo RopriGuez. 1985. Nuevos dato

etoecologicos sobre Anolis vermiculatus (Sauria:

Iguanidae). Poeyana 296: I-11.

, AND V. RIVALTA GONZALEZ. 2007. Efectos
probables del aumento del nivel del mar sobre
la herpetofauna de la Reserva de la Biosfera
Ciénaga de Zapata, Matanzas, Cuba. Poeyana
495: 8-13.

ROUGHGARDEN, J., W. Porter, AND D. HECcKEL. 1981.
Resource partitioning of space and its relationship
to body temperatures in Anolis lizard populations.
Oecologia 50: 256-264.

Rua, R. 1961. Thermal relations of five species of
tropical lizards. Evolution 15: 98-111.

ScCHOENER, T. W. 1967. The ecological significance of
sexual dimorphism in size in the lizard Anolis
conspersus. Science 155: 474-477.

tO
i)

. 1968. The Anolis lizards of Bimini: resource

partitioning in a complex fauna. Ecology 49:

704-726.

1974. Resource partitioning in

communities. Science 185: 27-39.

. 1977. Competition and the niche, pp. 35-136. In

C. Gans and D. W. Tinkle (eds.), Biology of the

Reptilia, volume 7, New York, Academic Press.

AND G. C. GorMAN. 1968. Some _ niche
differences among three species of Lesser Antillean
anoles. Ecology 49: 819-830.

SitvA RopriGuez, A. 1981. Estudio del nicho estruc-
tural, climatico y temporal de dos especies del
género Anolis (Sauria: Iguanidae), p. 185. In

ecological

BREVIORA

No. 520

Abstracts of the First National
Biology, La Habana, Cuba.
WILLiAMS, E. E. 1972. The origin of faunas: evolution of
lizard congeners in a complex island fauna: a trial
analysis. Evolutionary Biology 6: 47-89.
. 1976. West Indian anoles: a taxonomic and
evolutionary summary. |. Introduction and a
species list. Breviora 440: 1-21.
1983. Ecomorphs, faunas, island size, and
diverse end points in island radiation of Anolis,
pp. 326-370. In R. B. Huey, E. R. Pianka and T. W.
Schoener (eds.), Lizard Ecology: Study of a Model
Organism. Cambridge, Massachusetts, Harvard
University Press.

Congress of