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Full text of "Breviora"

BREVI 






Jyi Hi s e tq mi o i Lomparative 




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US ISSN 0006-9698 



Cambridge, Mass. 



8 October 2010 



Number 520 



THE ANOLES OF SOROA: ASPECTS OF THEIR ECOLOGICAL RELATIONSHIPS 

Lourdes Rodriguez Schettino, 1 Jonathan B. Losos, 2 Paul E. Hertz, 3 Kevin de Queiroz, 4 
Ada R. Chamizo, 1 Manuel Leal, 5 and Vilma Rivalta GonzAlez 1 

Abstract. 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 1 1 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 1 1 species to 
live sympatrically within a relatively small geographic area. 



Key words: Anolis; Cuba; Soroa; community structure 

INTRODUCTION 

In their pioneering 1969 paper "The anoles 
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 



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



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

2 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 

3 Department of Biological Sciences, Barnard College, New York, New York, U.S.A.; e-mail: phertz@barnard.edu 

4 National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.; e-mail: dequeirozk@si.edu 

5 Duke University, Durham, North Carolina, U.S.A.; e-mail: mleal@duke.edu 



The President and Fellows of Harvard College 2010. 



BREVIORA 



No. 520 



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, 1 1 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 (200 1 ) 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. 



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 1 1 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 



THE ANOLES OF SOROA 



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,' 1 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 (ayua), Prunus 
occidentalis (cuajani), Hibiscus elatus (maja- 
gua), C. antillarum (ocuje), Ficus aurea 
(jagiiey), Cedrela odorata (cedro), Bursera 
simaruba (almacigo), Pseudolmedia spuria 
(macagua), Oxandra lanceolata (yaya), Ma- 
tayba apetala (macurije), 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 
1 5-20 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). 



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 



B REV 10 R A 



No. 520 



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, 



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 1 1 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 



THE ANOLES OF SOROA 



Table 1. Snout-vent length (SVL) of 1 1 Anolis species from Soroa by site and sex. Lizards were measured at 

THE MOMENT OF CAPTURE. 





Site 








SVL(i 


mm) 














Males 






] 


Females 




Anolis Species 


N 


Mean 


SD 


Range 


N 


Mean 


SD 


Range 


A. alutaceus 


Forest 


3 


39.7 


7.37 


34-48 


1 


33 






A. angusticeps 


Forest 


— 








4 


35.2 


2.40 


32-38 


A. loysianus 


Forest 


5 


42.8 


5.23 


40-52 


— 








A. sagrei 


Chain-link fence 


24 


50.6 


4.35 


34-55 


5 


41.6 


2.30 


38-44 




Living fence 


15 


50.4 


4.29 


40-56 


— 










All 


39 


50.5 


4.27 


34-56 


5 


41.6 


2.30 


38^14 


A. mestrei 


Forest 


52 


52.0 


4.08 


40-60 


22 


41.6 


3.10 


35^16 


A. allogus 


Forest 


86 


52.5 


6.70 


34-62 


20 


42.0 


2.40 


39^6 


A. homolechis 


Forest 


49 


52.0 


3.42 


45-60 


33 


41.4 


2.39 


37-46 




Living fence 


22 


52.2 


2.20 


48-55 


4 


42.3 


4.50 


40-49 




Planted woodland 


26 


51.8 


2.30 


48-56 


3 


43.7 


2.31 


41^15 




All 


97 


52.0 


2.88 


45-60 


40 


41.7 


2.63 


37^19 


A. porcatus 


Forest 


2 


67.2 


11.70 


59-76 


— 








A. barbatus 


Forest 


3 


106.7 


20.50 


83-119 


2 


131.0 


12.73 


122-140 


A. luteogularis 


Forest 


1 


183.0 






6 


154.7 


4.50 


149-160 


A. vermiculatus 


Forest 


11 


121.5 


5.56 


116-128 


13 


78.3 


6.83 


70-90 



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- 



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 




Planted Woodland 


Living Fences 




Anolis Species 


Evergreen 


Gallery 


Secondary 


Artificial Structures 


A. allogus 


X 




X 




X 








A. alutaceus 










X 








A. angusticeps 


X 




X 




X 








A. barbatus 


X 




X 




X 








A. homolechis 


X 




X 




X 


X 


X 


X 


A. loysianus 


X 
















A. luteogularis 


X 




X 




X 


X 


X 




A. mestrei 






X 












A. porcatus 






X 




X 


X 


X 


X 


A. sagrei 






X 




X 




X 


X 


A. vermiculatus 






X 













BREVIORA 



No. 520 



Table 3. Frequencies of substrate use by 1 1 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. 





N 








Frequency 








Anolis Species 


Trunk 


Branch 


Leaf 


Vine 


Rock 


Ground 


Artificial 


Lizards sampled in natural areas 


















A. allogus o* 


139 


0.56 


0.08 




0.04 


0.25 


0.04 


0.04 


A. allogus 9 


20 


0.45 


0.15 




0.10 


0.10 


0.10 


0.10 


A. alutaceus o" 


23 


0.22 


0.35 


0.17 


0.13 


0.13 






A. alutaceus 9 


8 


0.25 


0.26 


0.13 


0.13 


0.25 






A. angusticeps Cf 


4 


0.50 


0.50 












A. angusticeps 9 


8 


0.38 


0.50 




0.13 






- 


A. barbatus 


9 


0.56 


0.44 












A. homolechis o* 


83 


0.52 


0.19 




0.01 


0.12 


0.07 


0.08 


A. homolechis 9 


36 


0.39 


0.17 




0.06 


0.11 


0.14 


0.14 


A. loysianus o* 


11 


0.91 


0.09 












A. loysianus 9 


5 


1.00 














A. luteogularis 


28 


0.39 


0.39 


0.04 


0.11 


0.04 




0.04 


A. mestrei o* 


96 


0.25 


0.08 




0.07 


0.50 


0.05 


0.04 


A. mestrei 9 


22 


0.14 


0.09 






0.68 




0.09 


A. porcatus o* 


8 


0.38 


0.38 


0.25 










A. porcatus 9 


6 


0.83 




0.17 










A. sagrei o* 


45 


0.18 


0.07 


0.02 




0.07 




0.67 


A. sagrei 9 


5 


0.40 












0.60 


A. vermiculatus o* 


18 


0.44 


0.22 






0.28 




0.06 


A. vermiculatus 9 


17 


0.12 


0.06 






0.71 


0.12 




Lizards sampled around human 


















constructions 


















A. homolechis o* 


79 


0.27 


0.03 


0.01 




0.04 




0.66 


A. homolechis 9 


36 


0.25 








0.03 


0.03 


0.69 


A. porcatus o* 


13 


0.62 


0.08 


0.15 








0.15 


A. porcatus 9 


6 




0.17 


0.67 


0.17 








A. sagrei cr 


44 


0.09 


0.05 


0.02 




0.07 


0.02 


0.75 


A. sagrei 9 


7 




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. mestrei — tend 
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 ah, 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 



THE ANOLES OF SOROA 



Table 4a. Perch heights for 1 1 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. glven the composite nature of 

these values, no standard deviations are reported. 



Perch Diameter (mm) 






Males 






Females 




Anolis Species 


N 


Mean 


SD 


Range 


N 


Mean 


SD 


Range 


Lizards sampled in natural areas 


















A. sagrei 


46 


0.9 


0.4 


0.2-1.9 


5 


0.9 


0.6 


0.3-1.8 


A. allogus 


139 


1.0 


0.6 


0.0-3.0 


16 


0.5 


0.4 


0.1-1.5 


A. mestrei 


93 


1.4 


1.1 


0.0-7.0 


21 


0.8 


0.9 


0.0^1.0 


A. homolechis 


84 


1.5 


2.4 


0.0 17.0 


23 


0.6 


0.5 


0.1-2.5 


A. alutaceus 


35 


1.5 


1.1 


0.2-5.0 


10 


2.2 


1.9 


0.3-6.0 


A. vermiculatus 


17 


1.6 


1.3 


0.0-4.0 


10 


0.5 


0.7 


0.0-1.5 


A. barbatus 


3 


2.8 


— 


0.5^1.0 


3 


5.2 


— 


0.3-19 


A. loysianus 


11 


3.1 


1.5 


1.1-5.0 


6 


1.6 


1.1 


0.3-3.0 


A. luteogularis (o* + 9) 


28 


5.9 


5.1 


0.3-20.0 










A. angusticeps 


3 


6.3 


3.8 


2.0-9.0 


9 


2.2 


1 


1.0-4.0 


A. porcatus 


8 


12.0 


9.7 


3.0 30.0 


7 


7.0 


4.7 


1.0-14.0 


Lizards sampled around human constructions 


















A. homolechis 


158 


1.0 


0.8 


0.1-9.0 


55 


0.6 


0.4 


0.0-1.5 


A. sagrei 


88 


1.0 


0.6 


0.0-1.8 


23 


0.5 


0.6 


0.0-1.8 


A. porcatus 


13 


2.0 


1.1 


1.0-5.0 


6 


2.4 


1.4 


1.5-5.0 



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 



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 1 1 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 



BREVIORA 



No. 520 



Table 4b. Diameters of perches (rock and ground perches excluded) used by 1 1 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. 









Perch Diameter (mm) 










Males 






Females 




Anolis Species 


N 


Mean 


SD 


Range 


N 


Mean 


SD 


Range 


Lizards sampled in natural areas 


















A. sagrei 


37 


6.2 


4.5 


0.2-20 


4 


6.3 


7.1 


0.2-16 


A. allogus 


139 


11.4 


13.9 


0.5-80 


13 


10.0 


12.7 


0.4-39 


A. mestrei 


40 


1.0 


12.9 


1.0-50 


5 


11.0 


7.6 


2.5-22 


A. homolechis 


64 


12.1 


13.2 


0.6-70 


26 


10.9 


13.4 


0.7-60 


A. alutaceus 


28 


1.1 


1.4 


0.2-7.0 


7 


2.1 


2.8 


0.3-8.0 


A. vermiculatus 


12 


14.6 


17.5 


1.5-67 


3 


10.0 


13.0 


2.0-25 


A. barbatus 


27 


1.7 


— 


0.5^1.0 


27 


2.0 


— 


0.3-19 


A. loysianus 


11 


18.7 


14.7 


2.0-50 


6 


27.5 


20.7 


8-60 


A. luteogularis (cr + 9) 


32 


12.5 


12.7 


1.0-50 










A. angusticeps 


4 


10.6 


9.9 


2.5-25 


9 


14.1 


12.0 


0.8-30 


A. porcatus 


7 


11.4 


9.7 


1.0-20 


7 


14.6 


10.6 


5.0-33 


Lizards sampled around human constructions 


















A. homolechis 


149 


15.8 


9.4 


1.3-60 


52 


13.1 


6.8 


1.3-30 


A. sagrei 


76 


6.6 


4.8 


0.5-25 


11 


5.0 


4.2 


0.5-15 


A. porcatus 


9 


11.9 


11.3 


1.5^0 


4 


6.6 


5.6 


0.5-14.0 



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 



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 
midafteraoon. 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 



THE ANOLES OF SOROA 



Table 5. Frequency at which 1 1 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 


52 


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 


41 


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 1 1 Anolis species at Soroa. Species are listed in descending order ok male 

mean body temperatures. 





ecies 








Body Temperature 


(°Q 












Males 








Females 




Anolis Sp 


N 


Mean 


SD 


Range 


N 


Mean 


SD 


Range 


Lizards sampled in 


natural areas 


















A. sagrei 




9 


32.1 


1.8 


29.6-34.9 


5 


32.9 


1.8 


31.6-33.9 


A. vermiculatus 




7 


29.6 


1.7 


26.7-32.0 


9 


29.8 


1.8 


27.0-32.1 


A. luteogularis 




8 


29.4 


1.1 


28.0-30.7 


— 








A. loysianus 




3 


29.3 


1.6 


27.7-30.8 


2 


28.6 


0.3 


28.4-28.8 


A. angusticeps 




— 








3 


29.2 


0.4 


28.7-29.4 


A. homolechis 




51 


29.2 


2.4 


24.2-33.5 


34 


28.7 


2.5 


22.3-33.4 


A. alutaceus 




4 


28.7 


1.3 


27.6-30.6 


3 


29.1 


1.0 


28.0-29.9 


A. mestrei 




53 


27.5 


1.9 


22.8-30.8 


23 


27.6 


2.0 


22.1-29.9 


A. barbatus 




3 


27.3 


0.2 


27.1-27.5 


— 








A. allogus 




82 


26.9 


2.2 


21.3-31.1 


18 


26.9 


1.9 


24.2-29.9 


Lizards sampled around human 


















constructions 




















A. porcatus 




3 


32.7 


2.19 


30.8-35.1 


7 


31.5 


1.4 


29.8-33.3 


A. sagrei 




87 


30.4 


3.36 


24.5-36.8 


16 


28.2 


2.1 


24.4-30.9 


A. homolechis 




133 


28.5 


2.02 


24.1-32.5 


54 


27.6 


2.1 


24.6-32.7 



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 



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 1 1 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 



II 



Table 7. Relative activity times for 1 1 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 




55 






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 


0.11 


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 1 1 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 


2 


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 



No. 520 



Anolis allogus 



Anolis alutaceus 




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. 




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 ANOLES OF SOROA 



13 



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 



(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 bar bat us 




'€2SPZsi&2 




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 



No. 520 



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 



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 



THE ANOLES OF SOROA 



15 



Anolis luteogularis 




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



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 



Anolis mestrei 






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ife'^yg. 4tewM 


- 


: nfm 





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 



No. 520 



Anolis porcatus 



A no lis sagrei 




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. 



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 



THE ANOLES OF SOROA 



17 



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 vevmiculatus 




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 



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 



BREVIORA 



No. 520 



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 etal, 2002). 



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 1 1 
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 Bermu- 
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 



THE ANOLES OF SOROA 



L9 



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 
of 29.2°C for A. allogus, 31.8°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. loysianus 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 




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 



No. 520 



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 
Tecnica de Los Cambios Globales y la 
Evolution 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. 

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