Ecological separation of Anolis lizards in a Costa Rican rain forest

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Title:
Ecological separation of Anolis lizards in a Costa Rican rain forest
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Anolis lizards in a Costa Rican rain forest
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vii, 403 leaves : ill., map ; 28 cm.
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English
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Corn, Michael Jon, 1944-
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Subjects / Keywords:
Lizards -- Costa Rica   ( lcsh )
Lizards -- Ecology   ( lcsh )
Animal ecology -- Costa Rica   ( lcsh )
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non-fiction   ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1981.
Bibliography:
Includes bibliographical references (leaves 397-402).
Statement of Responsibility:
by Michael Jon Corn.
General Note:
Typescript.
General Note:
Vita.

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University of Florida
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All applicable rights reserved by the source institution and holding location.
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oclc - 07904954
ocm07904954
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lcc - QL666.L25 C67 1981a
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AA00011130:00001

Full Text












ECOLOGICAL SEPARATION OF ANOLIS LIZARDS
IN A COSTA RICAN RAIN FOREST






BY

MICHAEL JON CORN


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA


1981














ACKNOWLEDGMENTS

I would like to thank the people of Rio Frio, Costa

Rica, for their friendship and hospitality, especially the

McGinnis family and their employees at the Hotel Cabana.

I also appreciate the cooperation of Mr. Ed Pattimore of the

Standard Fruit Company and the hospitality of his family.

Mr. Jack D. DeMent, vice president of Castle & Cooke Foods,

provided weather data for their Costa Rican sites.

Support for the field research was provided in part by

a National Science Foundation Traineeship, by collection

funds from the Florida State Museum, and by the Organization

for Tropical Studies (O.T.S.). I am grateful to these organi-

zations, but particularly to the Costa Rican office of O.T.S.

Sr. Jorge Campabadal and his staff were particularly helpful.

My introduction to tropical reptiles and their fasci-

nating ecological relationships was in an O.T.S. ecology

course in the summer of 1967. Drs. Tom Emmel and Roy McDiar-

mid provided an especially sound beginning to my studies.

In addition I also benefitted from many discussion with

two other O.T.S. faculty members, Dr. Jay Savage and Norm

Scott.

I would like to thank my many friends and colleagues

at the College of Lake County: the Sabbatical Leave

Committee for allowing me to return to Gainesville to finish








the prey analysis and begin writing; my friends in the Se-

curity Office for providing me a quiet, secluded work area

and for some "friendly nagging"; the A-V department, espec-

ially Mr. Bill Kniest, who photographed the figures; Mr.

Dan Ziembo, for the excellent background drawing in figures

3-1 and 5-26; Mrs. Lynn Floor, for typing the final draft;

and my fellow biology faculty members, for their continued

support and encouragement.

I am greatly appreciative of Drs. Archie Carr, Thomas

C. Emmel, Brian McNab and Hugh Popenoe, my graduate commit-

tee. Through the many years since I started this project,

they have been supportive and cooperative beyond any reason-

able expectation. Drs. Carr and Emmel have been especially

helpful in straightening out, or sometimes bending,

administrative pathways.

Finally my most important acknowledgment is to my

wife, Mary, and my children, Michael Jr., and Melanie.

Mary has helped in every stage of this project from the very

beginning to the final typing. They have all put up with

a seemingly endless, time-consuming project called "Daddy's

Dissertation" instead of doing many other activities.

I am most grateful for their love and support,


iii














TABLE OF CONTENTS


ACKNOWLEDGMENTS. . . ii

ABSTRACT . .. vi

SECTION

I GENERAL INTRODUCTION. . 1

II DESCRIPTION OF THE STUDY AREA 3

III DESCRIPTION OF THE ANOLES .. 15

IV BODY SIZE . . 23

1 Introduction. . .. 23
2 Materials and Methods . ... 25
3 Sexual Dimorphism . .. 27
4 Interspecies Comparisons. 30
5 Body Weight to Length Relationships 36
6 Seasonal Variation in Size. . 38

V FOOD. . . 96

1 Introduction. . .. 96
2 Materials and Methods . .. 97
3 Indices of Resource Use . 99
4 Intraspecies Comparisons by Size. .104
5 Intraspecies Comparisons by Sex .. 115
6 Intraspecies Comparisons by Season 118
7 Interspecies Comparisons ... 122
8 PreyUtilization by All Anoles .. 129

VI REPRODUCTION. . ... 245

1 Introduction. . 245
2 Materials and Methods . .. 247
3 Results for Anolis humilis ... 250
4 Results for Anolis limifrons ... 254
5 Results for Other Anolis ... 257
6 Female Reproductive Cycles ... 263
7 Interspecies Differences in Egg
Production Rate. ... .. 268
8 Male Reproductive Cycles ... 269
9 Fat Cycles and Reproduction .. 275










VII POPULATION SIZE AND STRUCTURE .. 332

1 Introduction. . .. 332
2 Materials and Methods . .. 333
3 Density and Biomass of Anoles .. 335
4 Population Structure of Anoles. .. 341
5 Density, Biomass and Population Structure
of Other Lizards . ... 346
6 Density of Frogs. .... ..... 349

VIII SUMMARY AND CONCLUSIONS . 376

APPENDICES

1 Definitions of Morphological Counts and
Measurements . .... 384

2 Specimens Used. . .. 385

LITERATURE CITED . ... 397

BIOGRAPHICAL SKETCH. . ... 403










Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy


ECOLOGICAL SEPARATION OF ANOLIS LIZARDS
IN A COSTA RICAN RAIN FOREST

By

Michael Jon Corn

August 1981

Chairman: Archie Carr
Major Department: Zoology


Ecological separation in eight sympatric congeneric

lizard species in the genus Anolis was studied at Rio Frio,

a site in a lowland tropical rain forest on Costa Rica's

Caribbean slope. Four aspects of the ecology of the Rio

Frio anoles are considered: (1) body size, (2) food, (3)

reproduction, and (4) population size and structure.

Size and prey patterns have been thoroughly described

for island anoles by other authors, but the Rio Frio anole

community contains more species in a single habitat than do

the island communities. In many respects, these lizards

show size-prey patterns similar to island anoles, e.g.

ecological separation is achieved primarily through spatial

differences, both vertical and horizontal, and size differ-

ence. But in contrast, differences in mainland anoles are

not as clear-cut, large male size is found in only two

species, body size differences are reduced, and prey size










differences are generally less than would be found within

or between island species.

Anoles that live without congeners on islands tend

toward certain size patterns. At Rio Frio, it was found

that anoles which are restricted to peripheral habitats,

e.g. streamside or exposed canopy, are little influenced by

congeners, and show the same size, sexual dimorphism and

prey size patterns as do solitary island anoles.

Anoles collected in May, just after the dry season,

weigh less at any length than those collected at other

times. That this is a result of dry season food stress,

due to reduced populations of small arthropods, is further

corroborated by the finding that lizards collected in May

frequently had not eaten.

Reproduction in Rio Frio Anolis is continuous throughout

the year. There is some variation in some species, apparently

in response to environmental change, but in none is there a

complete cessation of reproduction. Young are present in

all seasons.

Density of low bush and ground amphibians and reptiles

at Rio Frio was estimated from fenced quadrats. For all

species collected, density is 3752-5378 individuals/ha; for

Anolis alone, the estimate is 688-1252 anoles/ha. Anoles

had a combined estimated biomass of 422-1026 g/ha.


vii














I. GENERAL INTRODUCTION

This study concerns the ecology of the lizards of the

genus Anolis at a site in the lowland rain forest on Costa

Rica's Caribbean slope. Anoles are generally small, arboreal,

insectivorous lizards that make up a large, important part of

the fauna of the American tropics. In some areas, usually near

the geographic or climatic limits of the genus Anolis, only a

single species may be found, but in the rain forests many

species co-exist. Rio Frio, the site of this study, was

chosen because at least eight species of anoles live in close

contact, apparently dividing their resources--space, food,

time, etc.--in a way that minimizes, without altogether

avoiding, the evolutionary friction of competition.

Rio Frio is the Standard Fruit Company's name for a very

large banana plantation. When I first visited there, most of

the area was rain forest, slightly disturbed and with some

second growth, but basically mature forest. The banana

company was in the process of cutting the forest and replacing

it with bananas. Most of the lizards collected for this

study were taken as the trees were felled; a few were collected

in still undisturbed areas. When I last visited Rio Frfo in

May 1970, about two-thirds of the area had been cleared of

forest, and banana production was in full operation. At the

time of this writing, early 1979, the forest and most of its

animal inhabitants are gone.

1





2



So this study is about the ecological separation of the

anoles in a rain forest that used to be. Its conclusions

do apply, I believe, to the increasingly diminishing patches

of Costa Rican rain forest that remain.

Four aspects of the ecology of the Rio Frio anoles are

considered here: (1) body size, (2) food, (3) reproduction

and (4) population size and structure. Because of the dis-

tinctness of the sections, each will have its own, more

specific introduction.













II. DESCRIPTION OF THE STUDY AREA

Rio Frio is located at 10 20' N, 830 53' W in Heredia

Province, Costa Rica (Fig. 2-1). It is approximately 18 km

SE of Puerto Viejo and 16 km NW of Guapiles, and has an

elevation of approximately 100 M. All collections were made

on land belonging to the Standard Fruit Company, now a large

banana plantation. Most of my specimens were collected in the

area between the Rio Sucio and the Rio Chirripo. Population

quadrat collections were made on the NW side of the Rio Sucio.

Rio Frio is on a relatively flat alluvial plain below the

Cordillera Central. The area is shown as Premontane Wet

Forest, basal belt transition, on the Holdridge Life Zone map

of Tosi (1969), but rainfall data for 1961-65 (Table 2-1,

Figure 2-2) show more than 4000 mm of precipitation. This

would indicate a Holdridge Life Zone classification of Tropical

Wet Forest. The vegetation appeared to fit the description

(Holdridge et al., 1971) of Tropical Wet Forest more closely

than Premontane Wet Forest (Table 2-2). For a more detailed

description of a very similar forest, see the description of

Finca La Selva (Holdridge et al., 1971) which is located

about 10 km from Rio Frio.

While collections were being made at Rio Frio, the forest

was being replaced by bananas in the following sequence:

1. Selected trees were harvested for lumber. This

involved relatively few trees and caused no











appreciable difference in the structure of the

forest.

2. Some time after this lumbering (as long as many

months in some areas), bananas were planted in the

forest.

3. One to a few days after the bananas were planted,

all trees were felled and left to rot.

4. Bananas grew up among the rotting trees, and second

growth was cut back.

Most of my specimens were collected as the trees were

felled in step 3 above. Population quadrat collections were

made in an area that had been lumbered (step 1 above) many

months previously, but had not been otherwise disturbed. No

attempt was made to census or collect anoles in areas after

the forest was replaced by bananas.

It is generally accepted that wet tropical forests are

wet the year around, at least from a temperate point of view.

La Selva/Los Diamantes receives as much rain during the three

driest months, January through March, as Illinois does in an

entire year. Nevertheless, Rio Frio does have seasonality of

rainfall (Fig. 2-2). The wet season extends from May through

December, and the dry season from January through April. The

wet season has a characteristic rainfall depression, called

locally the veranillo, in late August and early September.

Even during this period, the forest usually receives regular

rainfall, and the forest floor is continuously damp. During

the dry season, however, there may be several days or even a






5




week or more with very little or no rain. Occasionally, the

dry season is interrupted by a period of rain, such as occurred

during February 1970. At Rio Frio, 495 mm of rain fell during

the 13 days from 5 February through 17 February, with only one

of these days receiving no rain. In contrast, the other 15

days of February received only 24 mm of rain. Because rainfall

data for Rio Frio were not kept prior to December 1969, the

average data from Puerto Viejo (18 km NW) and Los Diamantes

(15 km SE) are used.

Day length at the latitude of Rio Frio (Fig. 2-2) ranges

from 12 hours, 42 minutes at the end of June to 11 hours, 32

minutes in mid-December, a total variation of only 1 hour,

10 minutes (List. 1966).

Seasonal temperature variation is shown in Figure 2-3

and Table 2-3. Again, Rio Frio data are available only after

December, 1969. For a typical annual pattern, the Puerto

Viejo/Los Diamantes average is shown.

Anoles were collected during four periods (Fig. 2-2):

(1) September and early October, a dry period (veranillo)

during the wet season; (2) November and early December, the

very wet, last part of the wet season; (3) February and early

March, the driest part of the dry season; and (4) early May,

the last of the dry season or beginning of the wet season.

These four periods will be designated hereinafter simply as

September, November, February and May respectively. A few

anoles of some of the less abundant species were collected

during the preceding August and are used in some sections.










The fauna of lowland tropical forests is distinguished

for its; diversity, and the forest at Rio Frio was no exception.

Many other animals interact with the anoles as predators,

prey, or competitors. Predators include snakes, birds, small

mammals and occasionally larger lizards. Prey items include

most of the huge number of insect species, spiders, snails,

smaller lizards or frogs, and almost any other moving inverte-

brate of appropriate size. Chief competitors, for food at

least, are probably birds and frogs, and possibly spiders;

and not other lizards. Most other lizards at Rio Frio are

either nocturnal or herbivorous, or are found primarily in

second growth (Table 2-4). Thus, the most prominent lizard

competitors of anoles are other anoles.


















TABLE 2-1. Average Monthly Rainfall

PUERTO VIEJO
PUERTO LOS AND LA SELVA
MONTH VIEJO DIAMANTES AVERAGE

July 522.0 517.1 520

August 417.0 326.3 372

September 270.0 388.6 329

October 385.7 505.0 445

November 418.4 557.5 488

December 477.2 544.2 511

January 324.4 196.3 260

February 174.6 134.3 154

March 198.3 231.5 215

April 259.7 202.0 231

May 399.8 375.1 387

June 425.3 407.4 416

Total 4272.4 4385.3 4328

NOTE: Average monthly rainfall (mm) for the years 1961-65 at
Puerto Viejo and Los Diamantes, the two weather stations
closest to Rio Frio.









TABLE 2-2.
Comparison of Tropical Premontane Wet Forest
and Tropical Wet Forest Vegetation

Tropical Wet Forest

General. Tall, multistratal evergreen forest. A few canopy
species are briefly deciduous, usually when flowering, but this
does not affect the evergreen aspect of the forest as a whole.
Number of tree species very large.



Upper canopy. Trees 45-55 m tall, with occasional larger
emergents; crowns round to umbrella-shaped, usually not in
lateral contact with each other. Clear holes up to 30 m long
and 125-200 cm in diameter. Smooth, thin, light-colored bark
and high buttresses very common, but trees lacking buttresses
or with dark, rough, flaking or lissured bark also occur.
Leaves elliptical, usually lacking lobes or teeth, often glossy,
mostly less than 10 cm long.

Lower canopy. Trees 30-40 m tall, filling spaces between upper
canopy trees. Crowns round, trunks mostly slender, lacking
large buttresses. Bark dark or light, mostly smooth.

Understory. Trees 10-25 m tall, dense, commonly with narrow
conical crowns and slender stems, often leaning, twisted or
crooked, usually with smooth dark bark; flowers and fruits
sometimes produced directly from the trunk and lower branches
(cauliflory). Leaves or leaflets relatively large (up to 20
cm long), elliptical, long, pointed tips ("drip tips"). Stilt-
rooted palms often abundant.

Shrub layer. Dwarf palms, often with undivided leaves, usually
abundant, mostly 1.5-2.5 m tall. Miniature trees with unbranched
main stems present but usually less common. Giant herbs with
banana-like leaves often prevalent, especially in clearings
and disturbed places.

Ground layer. Often bare, or with a few ferns, Selaginellas,
or tree seedlings.

Epiphytes. Including orchids, bromeliads, and large-leafed
herbaceous climbers aroidss, Cyclanthaccae) common but often
not conspicuous; moss layer on tree trunks very thin or lacking.
Large bush ropes uncommon; epiphytic shrubs and strangling
trees mostly rare.

Tropical Wet Forest is distinguished from Premontane Wet Forest
by (1) larger buttresses of canopy trees; (2) larger trunk
volumes, especially in canopy trees; (3) bark generally smoother;
(4) palms much more common in understory and shrub strata;
(5) tree ferns less common; (6) more tree species per 0.1 ha;
(7) physiognomic differences between Tropical Wet and Premontane
Wet communities are subtle, but there are consistent differences
in floristic composition useful to the specialist; (8) more
luxuriant appearance than forests at higher elevations, which
feel cooler, darker, and more gloomy. (These subjective im-
pressions, of course, may not be shared by some observers.)











TABLE 2-2 extended

Tropical Premontane Wet Forest

General. Tall to intermediate semi-evergreen forest with two
or three tree strata; emergent, canopy, and subcanopy not al-
ways readily distinguished. In exceptionally dry years or on
edaphically dry sites, most canopy species may drop their leaves
for a few weeks. A few species are always dry-season decidu-
ous. The subcanopy, small trees, and shrub layers are evergreen.

Canopy. Trees mostly 30-40 m tall (with occasional emergents
up to 50 or 55 m), mostly with round to spreading crowns and
slender to stout trunks with clear lengths of 25 m or less.
Buttresses are common but much smaller than in Tropical Moist
and Wet forests, except in warm transitional areas. Bark mostly
brown or gray, moderately thick, flaking or lissured, but smooth
and light-colored in a few species. Leaves are mostly simple,
elliptic, glossy, 5-10 cm long, with entire or minutely toothed
margins, often somewhat crowded at the tips of the branches.
Number of species large.



Small tree stratum. Composed of a dense layer 10-20 m tall of
sapling canopy trees mixed with small, slender-trunked trees
having relatively deep crowns and smooth, often dark bark. Stilt
roots and long, strapshaped leaves are common. Tree ferns are
occasional; stilt-rooted palms occur in warm transitional areas.



Shrub layer. A dense stratum of single-stemmed "miniature"
trees and young canopy trees 2-3 m tall. Small palms are
generally rare or lacking, but are abundant in warm transition-
al areas.


Ground layer. Generally bare except for ferns, which may be
abundant (especially along trails), and tree seedlings.

Epiphytes. Orchids and bromeliads mostly not conspicuous;
herbaceous vines are common, mostly climbing the larger trees.
A thick layer of moss covers the tree trunks.



Premontane Wet Forest is distinguished from Tropical Wet Forest
by (1) much lower frequency of palms in small tree and shrub
strata; (2) higher frequency of tree ferns; (3) smaller but-
tresses in canopy trees, high buttresses rare; (4) bark gener-
ally somewhat rougher; (5) trunk and crown volume of canopy
trees somewhat smaller; (6) fewer tree species per 0.1 ha.


SOURCE: Holdridge et al., 1971.



















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TABLE 2-4. Other Lizards at Rio Frio


Anguidae
Celestus sp.
Diploglossus monotropis

Gekkonidae
Lepidoblepharis xanthostigma
Sphaerodactylus spp.
Thecodactylus rapicauda


Iguanidae
(Anolis 8 species)
Basiliscus plumifrons
Basiliscus vittatus

Corytophanes cristatus
Iguana iguana
Polychrus gutturosus


Scincidae
Leiolopisma cherries
Mabuya unimarginata


Teidae
Ameiva spp.


nocturnal (?), arboreal
nocturnal (?), terrestrial


nocturnal, terrestrial
nocturnal, arboreal
nocturnal, arboreal




diurnal, arboreal, riparian
diurnal, terrestrial,
riparian
diurnal, low arboreal
diurnal, arboreal, riparian
diurnal, high arboreal



diurnal, terrestrial
diurnal, terrestrial,
clearings


diurnal, terrestrial,
clearings


Xantusidae
Lepidophyma flavimaculatum


nocturnal, terrestrial










- 110


100


850



100 Km 84









83




FIGURE 2-1. Map of Costa Rica. Shown are the location of
Rio Frio (circle), a Standard Fruit Company
banana plantation, and the two nearest towns
that are on most maps: Puerto Viejo (diamond)
and Guapiles (triangle).











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III. DESCRIPTION OF THE ANOLES

Part of the rationale for studying ecological inter-

actions among sympatric congeners is that, being very close

in an evolutionary sense, they should show intense and, there-

fore, important competition. In this study, ecological

relationships of eight species of the large iguanid genus

Anolis are discussed. These eight species, although diverse

in their ecological niches, are very close phyletically.

The anoles are a group of over 200 iguanid species, the

distribution of which centers in the American tropics and

extends into the subtropics. All eight species of anoles

present at Rio Frio belong to what Etheridge (1960) and others

have termed the Beta Section, as species derived from Central

American ancestry are designated (in contrast to the Alpha or

South American derived species). Three species groups, desig-

nated as series by Etheridge (1960), are represented: petersi

series (A. biporcatus, A. capitol and A. pentaprion), fuscoauratus

series (A. carpenter, A. limifrons and A. lionotus) and

chrysolepis series (A. humilis and A lemurinus). Etheridge

(1960) suggested that the petersi series of large lizards is

a relatively primitive group, and that the fuscoauratus series

and chrysolepis series are its more advanced descendants.

The size, shape, color and gross habitat distribution are

ecologically important attributes of anoles. Some of these

will be discussed more thoroughly in later sections, but in











order to help the reader visualize these characteristics in

the eight species discussed here, a brief summary is given

below.

Anolis humilis is the smallest and most numerous anole

at Rio Frio. Maximum snout-vent length (SVL) is only 40 mm

in my samples, although Fitch (1975) found them up to 43 mm,

and maximum weight is 1.7 g. A. humilis is a stout-bodied

anole of more or less uniform brown dorsal color; females

occasionally have a light stripe or a rhomboid pattern on the

back. Males have a relatively large dewlap, red bordered

with yellow, while females lack a dewlap, but may have a

small patch of red on the throat. A. humilis is usually found

close to the ground; Fitch (1975) found 92% on the ground or

below 20 cm. They rarely go above 50 cm on a tree trunk, even

when chased.

Anolis lemurinus, the second member of the chrysolepis

series, is considerably larger than A. humilis. My largest

specimen has a 60 mm SVL, and the heaviest is 5.06 g. Taylor

(1956) describes a specimen with a 63.5 mm SVL. Also a stout-

bodied anole, A. lemurinus is dark gray in color usually with

a blotched dorsal pattern; females may have a light band or a

series of diamonds on the dorsal midline. Males have a rela-

tively small dewlap, deep red in color, lighter at the edge.

The female dewlap is very small, but colored like that of the

male. A. lemurinus is found on the trunks of large trees,

usually on the lower part of the trunk, but when disturbed

will flee quite high.










Anolis limifrons is a slender, gray anole that is second

only to A. humilis in abundance at Rio Frio. Slightly longer

than A. humilis, A. limifrons has a maximum SVL of 42 mm in

the Rio Frio collection; Fitch (1975) lists 45 mm as the

maximum. Maximum weight in the Rio Frio sample was 1.5 g;

Fitch (1975) gives 1.7 g. Males are more or less uniform light

gray, lighter on the venter having a small white dewlap with

an orange spot in its center. Females are colored like the

males, or with a light dorsal band or diamond pattern. Females

lack a dewlap. A. limifrons is a shrub-level anole usually

found on vertical trunks of shrubs, saplings or larger trees,

almost always between 0.5 and 2 m above the ground. Fitch (1975)

found 87.3% of this species perching above ground level, with

a mean perch height of 0.78 m. More than any other Rio Frio

anole, A. limifrons is found commonly in second growth and

disturbed areas, but is still more abundant in less-disturbed

forest.

Anolis carpenter is closely related to A. limifrons, and

is virtually the same length. The longest Rio Frio specimen

is 43.5 mm SVL; Fitch (1975) records it up to 45 mm SVL. It

is slightly more slender than A. limifrons, with the heaviest

A. carpenter weighing only 1.2 g. A. carpenter, which was

not described until 1971 (Echelle, Echelle and Fitch), is a

very cryptic, lichen-green anole with a light half-ring below

the eye and a white venter. Males have a large bright orange

dewlap; females lack a dewlap but may have a trace of orange










pigment on the throat. The habitat of A. carpenter is

problematic. Fitch and his co-workers (Echelle, Echelle and

Fitch, 1971; Fitch, 1975) found their first ten specimens on

lichen-covered rocks at ground level in the Tropical Moist

Forest, premontane transition Life Zone. They then obtained

14 more lizards in 10 months at Finca La Selva in the Premon-

tane Wet Forest, basal belt transition Life Zone. Of these

.14, only 5 were from undisturbed rain forest; the rest were

from cacao or palm groves. Five of these lizards were found

at ground level, three on tree bases and two in leaf litter.

Fitch (1975) states that all were "on or near lichen-mantled

tree trunk or log which provided concealing background," and

that three of fourteen recorded "dropped to the ground almost

immediately after being flushed." When I first visited Rio

Frio in August 1969, I found A. carpenter to be very common

among the just-felled trees. In fact, A. carpenter was so

common that I collected several, decided it was a common

species that I didn't recognize, and released them all. Only

after a later conversation with N. J. Scott, did I realize

that the species was probably then undescribed. A. carpenter

was also quite common among the cut trees in September and

October, but less common in November, December, February and

March. None were seen in May. Only one specimen, a 26 mm SVL

o, was collected outside the area of newly felled trees. This

individual was collected from a small sapling after it had

been flushed out from a liana about 2.5 m above the ground.










I would suggest that the apparent rareness of A. carpenter

reflects its preference for lichen-covered perches, probably

at moderate heights. The few specimens found on the ground

(with the exception of the type series) are there because of

a "drop-and-freeze" escape behavior. While not as common as

A. limifrons or A. humilis, it is like several other rain

forest reptiles (for example, see Corn, 1974), only encountered

when disturbed from its high arboreal habitat.

A. lionotus is the largest anole of the Rio Frio

fuscoauratus series, and the most restricted in habitat. It

is never found more than a few meters from the edge of a river

or stream. These lizards perch on the bare bank, rocks,

debris and occasionally on vegetation. When disturbed, they

hide under debris, or readily enter the water, where they

swim and hide under submerged material. The largest Rio Frio

A. lionotus is 71 mm SVL, weighing 6.6 g. Fitch (1975) found

much larger specimens, up to 85 mm SVL and 13.7 g. The color

of both sexes is chocolate-brown with cream-colored lateral

stripes and a lighter venter. Males have a large, pale orange

dewlap, which females lack.

Anolis capitol is a rare, large anole. The largest Rio

Frio specimen was 87 mm SVL, weighing 13.0 g. Both Fitch

(1975) and Andrews (1971a) found them as long as 95 mm SVL.

The color of A. capitol is a mottled olive-green to brown.

Females occasionally have a distinct brown mid-dorsal stripe.

The male dewlap is very small and greenish-white; an even

smaller, similarly colored dewlap occurs on females.











A. capitol is found on the ground or on tree trunk bases or

buttresses, usually lower than 1 m. Escape behavior involves

freezing, or, occasionally, running across the ground or onto

a tree trunk, never higher than two meters or so. A. capitol

was probably much more common at Rio Frio than my collecting

suggests, but its escape behavior made it very difficult to

find in the newly felled trees.

Anolis biporcatus is the largest Rio Frio anole, reaching

93.5 mm SVL, and weighing up to 20.0 g. Taylor (1956) states

that the maximum SVL is 102 mm. Dorsal color can be changed

from bright green through medium brown in both sexes. The

male dewlap is large, with a bright blue center and an orange-

red outer portion. The dewlap of the female is smaller.

A. biporcatus is a tree anole, found from lower sections of

trunks to high in the crown.

Anolis pentaprion is a moderately large, short-legged

anole. My longest specimen is 66.5 mm SVL, and the heaviest

4.3 g; Taylor (1956) and Fitch (1975) examined a specimen of

75 mm SVL. Dorsal coloration is light gray sometimes mottled

with dark patches. The dewlap is large and deep purplish-red

in color. The dewlap of the female is only slightly smaller

than that of the male. The tail is relatively short and

somewhat prehensile. A. pentaprion is the only truly open-

habitat heliotherm among the eight Rio Frio anoles. Most

specimens were found on exposed tree trunks in open country

or high in the crowns of forest trees. All of mine were taken










from just-felled trees. In forests A. pentaprion probably

spends most of its time high in the leaves and twigs, where

it can bask, or at least take advantage of the higher tempera-

tures of the exposed tree crown. Campbell (1971) obtained a

mean of 330 C for 12 records of A. pentaprion in a thermal

gradient. The short legs and prehensile tail are undoubtedly

adaptations for movement on small twigs and branches.

From the preceding descriptions, it can be seen that the

eight anole species at Rio Frio are well distributed throughout

the habitat, with some important separation, both vertically

and horizontally (Fig. 3-1). Anolis capito, A. humilis and

A. lionotus form a group of highly terrestrial species, with

A. humilis and A. capito restricted to forest floor and tree

bases, and A. lionotus almost completely separated from all

other anoles in its riparian habitat. A. biporcatus and

A. lemurinus are usually found on tree trunks of moderate to

large size, and only rarely elsewhere. A. limifrons occurs

on a rather broad spectrum of perches, from ground to moderate-

ly high trunks, but is primarily found in the shrub layer.

A. pentaprion and A. carpenter occupy the highest habitats,

with A- pentaprion preferring the sunny crown and A. carpenter

restricted to dark, lichen-covered trunks, branches and lianas.
























- -


wr


i- '7P"-V














IV. BODY SIZE

1. Introduction

The size of animals has always been of theoretical

interest to zoologists, and Hutchinson's (1959) classic

observations on size ratios of closely related species

pointed the way to an interesting and fruitful area of

investigation for ecologists. How do the sizes of sympatric,

closely related species increase or decrease their ability

to co-exist? From Hutchinson's (1959) own ratio suggestion,

several authors have proposed various models, and many others

have compared sizes within a variety of taxa (See Schoener,

1974, for an extensive review.). The sizes of anoles in

simple and more complex West Indian faunas have been the

subject of numerous studies (Schoener 1967, 1968, 1969a,

1969b, 1969c, 1970; Schoener and Gorman, 1978; Williams,

1972; and others). Only Fitch (1976) and Duellman (1978)

have discussed size in sympatric mainland species.

Two particularly interesting generalizations have come

from these studies. From the work of Schoener, have come

what Williams (1972)called Schoener Rules:

An anole occupying an island without congeneric
competition tends to a range of sizes with
maximal head lengths of between 20 and 25 mm
and maximal snout-vent lengths of between 65
and 96 mm. (p. 52)











If two anole species occur on an island,
one will be smaller and the other larger,
the ratio of the two sizes ranging from
1.5 to 2. (pp. 57-58)

The greater the diversity of island anole
faunas, the greater the disparity between
the largest and smallest species. (p. 48)

These are rules meant to apply only to anoles in the West

Indian islands, but they also appear to have some appli-

cation to the more complex faunas of the mainland.

A second tentative generalization comes from Williams'

(1972) fascinating inferential model of the historical co-

adaptation of the complex anole fauna of Puerto Rico. Build-

ing upon a foundation of comparative osteology, karyotypes,

electrophoretic patterns and ecological information, Williams

has attempted to reconstruct the probable course of evolution

of the Puerto Rican anoles. From this and his wide knowledge

of other anole faunas, Williams (1972) concluded that Puerto

Rican anoles (and supposedly other anoles in complex island

faunas)

are first able to utilize size differences as
a major means of syntopic co-existence. Beyond
the stage of the third species, however, size
ceases to have the same importance, and spatial
shift and climatic shift become
essential elements in the adaptations that
permit the addition of species to the fauna.
(pp. 87-88)

In light of these ideas, it would seem reasonable

that any discussion of the ecology of anoles should first











focus on size distribution among the species under consider-

ation. That is the purpose of this section.

2. Materials and Methods

Times and places in which collections were made are

described above (section II). All anoles collected at Rio

Frio are considered in this section. Anoles were taken during

the morning and early afternoon, usually 9:00 A.M. to 2-3:00

P.M., and held in plastic bags until processing. As soon as

possible after collection, usually within six hours, each

lizard was weighed (alive) to the nearest 10 mg on an Ohaus

balance. Each was then killed by an injection of sodium nem-

butal, and the snout-vent length and tail length were measured

to the nearest 0.5 mm with a plastic ruler. Head and hind leg

measurements (with a vernier caliper) and lamellae counts

(See Appendix 1) were made after the lizard had been preserved

in 10% formalin. Sex was determined by dissection (See

Appendix 2).

An outstanding problem of any morphological comparison

of different taxa is to determine which specimens should be

compared. I considered four possibilities:

1. All specimens of each species. The weakness of this

is that collecting bias might increase or decrease

the mean size data.

2. Sexually mature adults. Assessment of egg production

makes this method plausible for females, although

seasonal cessation of reproduction is a problem in










some species. For males, however, seasonal variation

in testis size and sperm production in relatively

small subadultt" size males (Fitch, 1956) makes

designation of a minimum adult male size extremely

arbitrary.

3. The ten (or other arbitrary number) largest of each

species. This would be useful if maximum size were

desired, but sample size affects this choice too

greatly for its use here. For example, the ten

largest individuals of A. capito comprise 43% of my

collection of that species, while the ten largest

A. limifrons make up only 2.8% of my collection of

that species.

4. The largest third (or some other fraction) of each

species. This essentially eliminates the effect of

sample size, yet, if the fraction is appropriately

small, a reasonable sample of "largest adults" can

be compared. For a more thorough discussion of this

method, see Schoener (1969c).

Because an unbiased approximation of adult size is desirable

for size comparisons, I have chosen the fourth alternative,

carried out in the following manner. All specimens of each

category considered (species, sex or collecting period) were

ordered from longest to shortest. The length of the individual

that was one third of the number of specimens from the longest

was used as the lower limit for that sample. All specimens










of that length or greater are used in size comparisons for

that category, regardless of whether it is length or weight

being compared. When used hereinafter, the term "longest

third of" will refer to such a sample of the indicated

category.

3. Sexual Dimorphism

Inasmuch as sexual dimorphism in body size is often strong

in anoles (Rand, 1964, 1967a; Schoener, 1967, 1968, 1969c;

Schoener and Gorman, 1968; Fitch, 1976), it was deemed appro-

priate to determine the patterns) and degree of intraspecific

sexual dimorphism before making size comparisons among the

species. Table 4-1 lists means and standard errors for snout-

vent length (SVL), head length (HL) and body weight (BW) for

all specimens of all eight species of Rio Frio anoles. Table

4-2 indicates the ratios and significance of differences

between those means. All eight species show a significant

(P<.05) degree of sexual dimorphism in all three parameters

except in the case of A. biporcatus for HL.

Two patterns of dimorphism are seen, one in which the

females are larger than the males (6 species), and the second

in which the males are larger than the females (2 species).

The patterns and relative positions (Fig. 4-1) are fairly

consistent for SVL, HL and BW. Using measurements from both

living and preserved specimens, Fitch (1976) examined sexual

dimorphism in 54 groups of anoles from Mexico to South America,

but primarily middle America. He found SVL ratios as low as

0.80 (Anolis vittigerus of Panama) and as high as 1.36











(A. cuprinus of xeric Mexico), but only in 34 of 54 was the

dimorphism significant. In his samples of A. biporcatus,

A. carpenter and A. lemurinus, males and females were not

significantly different in SVL. Differences in our findings

may be due in part to inadequate sampling of large males

(lemurinus, lionotus, and pentaprion) by me and/or Fitch's

combining of samples from various localities, seasons or

years. SVL ratios in the six species studied by Duellman

(1978) at Santa Cecilia, Ecuador,ranged from 0.89 (A. trachy-

derma) to 1.10 (A. punctatus).

The pattern in which the males are larger than the females

is the rule in West Indian anoles, especially in solitary

species (Schoener, 1969a). This is apparently true also for

mainland Alpha anoles. Both species which have larger males

than females reported by Duellman (1978) are in the Alpha

section of Anolis. All seven of the Alpha anoles measured

by Fitch (1976) have larger males (significant in 6 of 7).

In Beta anoles, both patterns are common, but seem to

sort out on a climatic (faunal size?) basis. Seventeen popu-

lations studied by Fitch had significantly larger males. He

considered 12 of the 17 (including A. lionotus and A. pentaprion)

to be primarily inhabitants of "severely seasonal" climates.

These are areas which also have reduced anole faunas, compared

to lowland rain forests. Only 3 of 21 Beta groups attributed

to "lowland rain forests" by Fitch have significantly larger

males. Of the 11 populations having significantly larger

females, eight are "lowland rain forest" forms, and the other










three inhabit montanee or cloud forest" areas, both of which

are "relatively seasonall" No species which have the females

larger than the males inhabits severely seasonal areas. The

Rio Frio anoles fit this pattern precisely.

The development of sexual differences in solitary anoles,

in which males are always larger than females,is seen to be

a result of lack of congeneric competition (Schoener, 1967).

The two Rio Frio species with larger males are the two with

the least congeneric pressure. A. lionotus is the only anole

which normally occurs in the riparian microhabitat, i.e. it is

essentially a solitary anole, and it very closely fits the

solitary anole size pattern. Another point, which holds for

A. lionotus (but not for A. pentaprion) is Schoener's (1969b)

observation that relative head size is less in solitary anoles.

Table 4-3 indicates that A. lionotus, the "most nearly solitary"

anole at Rio Frio, has the shortest head relative to SVL.

A. pentaprion inhabits the outer branches and twigs of the

forest crowns. No other anole, with the possible exception

of the much smaller A. carpenter, overlaps with it in this

exposed, climatically severe microhabitat.

The pattern in which the females are larger than the

male is seen by Fitch (1976) to be an asset to species in

seasonal habitats, allowing them to maximize egg/young pro-

duction. While the six Rio Frio species with this pattern

are all primarily inhabitants of seasonal microhabitats, these

are also species which generally occur with the greater numbers











of congeneric competitors and can, therefore, least support

the food niche expansion that sexual dimorphism requires. Any

increase in the number of anole species in a fauna will result

in a general decrease in the availability of prey. Most par-

ticularly this will reduce the density of the least common

large items, which are the prey of the large males in dimorphic

species (Schoener, 1967, 1968; Schoener and Gorman, 1968).

Schoener's (1969a, 1969b) models and examples (1969b, 1970)

suggest that such an increase in food competition might result

in a reduction of body size of all species, as smaller lizards

are better able to exploit the more numerous small prey items.

It would seem a reasonable explanation then, that the general

anole pattern in which males are larger than females is

abandoned in rain forest Betas because of the greater compe-

tition in multi-anole fauna. Male size is more reduced than

is female size giving the size advantage to the reproductively

more important, egg producing females. The following section

(V) will further explore sexual differences in food.

4. Interspecies Comparisons

When an anole species exists without congeneric competitors,

Schoener (1967, 1968, 1969c) suggests that an optimum size

exists. When more than one species is present, sizes must

diverge from the solitary anole size to alleviate pressure on

particular prey sizes. Table 4-1 indicates the wide range of

sizes of Rio Frio Anolis (Fig. 4-2, 4-3). Note the range,

from a minimum mean SVL of 33 mm for A. humilis males to a











maximum of 91 mm for A. biporcatus females, an almost three-

fold difference. As would be expected, the difference in

weight is even greater, from 0.65 g (A. carpenter males) to

17.1 g (A. biporcatus females), a difference of over 26 times.

The expectation for a complex assemblage of species is

that each species will differ from the next by some constant

factor. Hutchinson (1959) found a range of 1.1 to 1.4 with an

average difference of 1.3 times, in a variety of taxa. Table

4-4 shows that this is not the case for Rio Frio anoles. Indeed,

if eight species could exist with the smallest 33 mm in SVL

and the length of each larger species increased by 1.3 times,

the largest in the sequence would be just over 207 mm SVL,

much larger than any mainland anole. To complicate this

possibility further, Schoener (1970) has shown that the

separation ratio should increase between larger species, so

the largest in an eight species sequence would be much larger

than any known anole. Since there is apparently not a constant

or increasing ratio between all eight species at Rio Frio,

is it possible to subdivide the assemblage into some logical

grouping that will show the expected pattern?

One possible sorting of species might be along phyletic

lines. The species groups (or series) of Etheridge (1960)

apparently represent well established evolutionary lines.

Table 4-5 shows that, within each of the three series represented

at Rio Frio, the separation ratios are not exactly in keeping

with Hutchinson's (1959) suggestion. The two members of the











chrysolepis series are too widely separated. In the fusco-

auratus series two species are too close, and the third is

too large. Finally, in the petersi anoles, the females of the

two smaller species are separated by too large a ratio, though

the males fall in the low end of Hutchinson's bracket, and

the larger two are just barely different enough to pass the

lower limit. In fact, only 1 out of 10 SVL comparisons and

5 of 10 for HL fall within the 1.1 to 1.4 range, even though

the average separation ratios are near 1.3 (1.32, SVL; 1.30 HL).

Though a slight expansion of the ratio range would encompass

most, if not all the ratios in Table 4-5, perhaps something

other than a phyletic grouping might yield a more logical,

consistent set of size relationships.

In an evolutionary sense, there seems to be no reason why

congeners should differ in size simply because of phyletic

relationship. Indeed, Hutchinson (1959) proposed that the

divergence of syntopic species pairs occurs because of their

ecological similarity. Therefore, ecological resemblances

(microhabitat, perch site or structural niche) provide a

better basis for grouping the species. Most of the anoles

which I collected or observed at Rio Frio were in the area of

newly felled trees. With this type of data, it is not possible

to establish numerical boundaries on the structural habitat

of each species as is frequently done (Rand, 1964; Schoener,

1967, 1968; Schoener and Gorman, 1968; Schoener and Schoener,

1971a, 1971b; Andrews, 1971a, 1971b). However, for the












discussion here, general descriptions of the habitats

(such as those given in Section III) will be quite use-

ful. As Williams (1972) explains with regard to habitat

similarity in two Puerto Rican anoles:

They both may be crudely described as crown
animals of the shaded forest. It is
possible to quibble a bit about this but
the same modal situation describes both.
A giant anole such as cuvieri, though it
is, in fact, seen some of the time at every
level (personal observation), is most
often seen high in the crown. A dwarf
anole such as occultus is seen on branches
and twigs of small diameter and
therefore not infrequently on bushes and
vines, but certainly its modal structural
niche includes the crown. (p. 74)

The eight Rio Frio species are best sorted into four

groups:

1. Ground dwellers. This set includes A. humilis

and A. capitol, both of which inhabit tree

bases, logs and the intervening ground

surface.

2. Riparian species. Only A. lionotus fits here.

It is thus horizontally removed from the

other seven species.

3. Trunk dwellers. Included here are A.

limifrons, primarily an inhabitant of

lower trunks, smaller trees and low shrubs;

A. lemurinus, found on larger, higher trunks;

and A. biporcatus, on large trunks up to, and

possibly into, the base of the crown.











4. Crown inhabitants. Although A. biporcatus might be

included here, I am using this to mean only the

small branches and twigs, the habitat of A. pentaprion

and probably A. carpenter.

Table 4-6 gives the length ratio of the eight species grouped

by perch site.

The extension of the Schoener Rules (Williams, 1972)

and other findings of Schoener (1970) to Rio Frio anoles leads

to the expectation that: (1) when an anole is without congeners

in it's particular habitat, it will tend toward a particular,

sexually dimorphic size pattern (55-75 mm SVL for males,

40-60 mm SVL for females); (2) when two anoles share a micro-

habitat, they will differ by a factor of 1.5 to 1, with one

larger and one smaller than the solitary anole size; and

(3) when three anoles occupy the same structural level, the

size ratio will be greater between larger species than between

smaller. Schoener (1970) found an average ratio of 1.5 between

smallest and 2.2 between largest species in 10 syntopic trios.

The Rio Frio anoles seem to fit these expectations.

The only anole that rarely shares it's habitat with other

anole species is A. lionotus, and it clearly fits the expected

pattern, except that there is not quite as much sexual di-

morphism as expected. One factor that might push the female

size upward, i.e. limit the dimorphic spread, is the presence

of the smaller, more numerous A. humilis on the adjacent, non-

riparian forest floor.










Two anole habitats, the ground and the crown, are also

relatively close to expectation. The two anoles with the

lowest perch sites, A. humilis and A. capitol, are on either

end of the solitary anole size range, but they are separated

by more than the 1.5 to 2.0 suggested by rule two. The ex-

ceptionally small size of A. humilis is perhaps explained by

its partial microhabitat overlap with A. limifrons. This

small, abundant species might compete in such a way as to

push the size of A. humilis down. In fact, both A. humilis

and A. limifrons may be reduced in size by any overlap (Schoener,

1969a, 1969b). The crown anoles, A. carpenter and A. pen-

taprion, are separated by the required size range in males, but

not in females this being due to their having opposite

patterns of sexual dimorphism. Though A. carpenter is below

the bottom end of the solitary anole size range, A. pentaprion

is not larger than the solitary size which rule two specifies

it should be. This may be due to the small sample size.

However, in equally small samples, Fitch (1976) found a mean

of 74.2 mm SVL for male and 60.0 mm for female A. pentaprion.

These measurements would make A. pentaprion males 2.03 times

and females 1.45 times the length of A. carpenter and just

longer than the size range of solitary anoles. It may be that

the size of A. pentaprion is held down by the influence of the

larger A. biporcatus in the crown-- or even by the presence

of Polychrus gutturosus, a larger, although primarily vege-

tarian iguanid which was found in crowns at Rio Frio.











The only place where three species of anoles regularly

perch is the area broadly termed "trunk." Here the pattern

of increasing separation in larger species (Schoener, 1970) is

seen, although the magnitude is somewhat less than Schoener's

average for 10 islands. In both sexes, the difference in

length between A. biporcatus and A. lemurinus is greater than

the difference between A. lemurinus and A. limifrons.

In general then, the sizes of Rio Frio anoles are well

explained by Schoener's island patterns if habitat types are

considered separately. That small anole faunas on West Indian

islands are isolated by sea water, while the Rio Frio anoles

are segregated only by the much more nebulous boundaries of

perch site preference, should explain the slight variation.

5. Body Weight to Length Relationships

Schoener (1969a) predicted that weight could be related

to length by a power function {such as BW=a(SVL)b} and that

the power, b, ought to be close to 3, in groups such as Anolis

lizards. He was able to confirm this only with data for the

Puerto Rican A. gundlachi from Turner et al. (1965). He also

expected that b might prove adaptive where there is an advan-

tage in increased length without increased weight, as in a

twig inhabiting anole. Data from Rio Frio Anolis fit these

expectations quite well (Table 4-7, Figs. 4-4 to 4-11). The

exponent b ranges from 2.46 to 3.27, with a mean of 2.91.

Because of the weight increase associated with the repro-

ductive activity of larger females, the regression line for











females bends more than the curve for males. For females b

averaged 2.98, while the mean for males was only 2.83. In all

species except A. lemurinus, males have a lower b than females,

reflecting the relative slimness of large males. This dif-

ference was significant only for A. limifrons and A. biporcatus

(Table 4-8). Male A. limifrons often perch higher than females

(Fitch, 1975), so the "slimness" of males may be an additional

aid in moving along small stems and vines, or over leaves.

No information is available on sexual differences in perch

heights of A. biporcatus.

Though comparable reports are not available for other

Anolis, at least two other tropical iguanids seem to show the

same sexual difference. In two Costa Rican populations of

Basiliscus basiliscus, Van Devender (1978) found males to have

a significantly lower b than females. Recalculation of

weight-length regression equations for Ctenosaura similis

given in a different form by Fitch and Henderson (1977) also

show a higher b for females than males.

Comparisons of b between pairs of species show few

differences (Table 4-8). The exceptionally slim A. carpenter

have a lower b than males of all other species. This difference

is significant for comparisons with all species except A. hu-

milis and A. pentaprion. This may reflect a real adaptation

to the twig-vine habitat, or it may be an artifact of the

rather small, restricted sample of A. carpenter since 14 of

21 weighed were within 5 mm in SVL and 12 were within 2 mm











(Fig. 4-6). Female A. biporcatus have a higher b than all

other females, but this was significant only for comparisons

with A. carpenter and A. lemurinus. The large b may reflect

the fact that A. biporcatus perches only on the broad trunks

and large limbs of trees; but more likely it simply reflects

the fact that A. biporcatus females are by far the largest

anoles at Rio Frio, and are almost always gravid.

6. Seasonal Variation in Size

There are two ways in which seasonality may cause differ-

ences in the sizes of lizards. A decrease in mean size may be

caused by changes in growth rate or by different reproductive

rates, which will alter the fraction of small lizards in the

population. A change in the weight-length regression power,

attributable to a size dependent change in food consumption,

could be the cause of a seasonal difference. Even though

little seasonality is apparent at Rio Frio (see Section II),

anoles may respond to minor variations. In this section

that possibility will be considered.

Table 4-9 shows SVL, HL and BW of four species of Rio

Frio Anolis for each of the four collection periods. Samples

of the other species were too small for adequate seasonal

comparison. Table 4-10 shows significance of seasonal dif-

ferences in SVL and BW (see also Figs. 4-12, 13, 14, 15).

A. humilis shows very little seasonal change. Females

are largest (both SVL and BW) in the September sample, the

climax of the rainy season, but this is significant only when











compared to February, the driest season. There are no sig-

nificant changes in the males of A. humilis.

Size of A. limifrons is affected by a decrease in the

number of large females and an increase in the number of small

males in the November sample, and by a decrease in the number

of juveniles and small of both sexes in the September and

May samples. The size then has an apparent "dip" at the end

of the rainy season. A similar pattern was seen by Fitch (1973a)

in A. limifrons from Turrialba, Costa Rica, which has a rain-

fall pattern similar to that of Rio Frio. In both sexes, May

specimens were the longest, but were not as heavy as the

shorter September lizards. This would seem an indication that

they are not feeding as well at the end of the dry season as

they did during the wet months. Even though the sample sizes

are very small, A. lemurinus and A. lionotus appear to be

similar to A. limifrons in seasonal size pattern. Both have

a late rainy season decrease in mean size, with an increase in

length through May.

There is one immediately obvious difference in the BW-SVL

regression curves (Figs. 4-16, 17, 18, 19)--that for May is

well below all others. This is further evidence that, at any

SVL, A. humilis or A. limifrons will weigh less in May than

in any other period. The November curve is the highest in

three of the four indicating that anoles are heaviest at the

end of the rainy season.

The same pattern is approximately true for A. lemurinus

and A. lionotus. Even though sample sizes are small for some










periods and no A. lionotus were weighed in September, the

regression curves of the rainy season samples are highest, and

the May curves are lowest (Figs. 4-20, 21), as is true for

A. limifrons and A. humilis. The May curves are altered some-

what by the absence of juvenile specimens of either A. lemurinus

or A. lionotus.

Differences in b, the exponent of weight-length relation-

ships are not so easily explained (Tables 4-11, 12). First,

for A. humilis and A. limifrons in all periods, b is larger

for females than for males as found in the-preceding section

for all species. This is significant only for the two rainy

season samples of A. limifrons. But there is some variation

within the sexes that is more puzzling. In three of the four

species, the May samples have the highest b, although in some

comparisons this difference is not significant. The regression

exponent is low in most February samples and low in September

samples of A. humilis.

The regression exponent, b, is essentially the factor deter-

mining that- the big get bigger, i.e. the:.upward curvature of the

regression line. As such, it appears that even though most

groups are lightest in May, the longer individuals are com-

paratively heavier than the shorter ones, an indication that

a late dry season food decline has a more pronounced effect on

smaller lizards. This may be caused by a variety of factors,

probably including actual physical struggle for food items,

size proportional territory (and, therefore, more potential

prey for larger individuals), and higher perches and a larger










scanning radius for larger anoles. The only exception is

A. lemurinus--ttI sample of May females has a heavier BW as

well as a longer SVL. However this is from a sample of two,

so the differences are not significant.

The only exception to the May peak of b is the case of

A. lionotus. In A. lionotus, the complete absence of small

individuals causes the regression line to rotate clockwise.

If the curve were drawn below the SVL of the smallest May

A. lionotus collected (56.5 mm), it would predict improbably

heavy juveniles (dotted line on Fig. 4-21). If normal-sized

juveniles had been collected, they might well have bent the

regression curve to the "normal" pattern.





















Sizes of Longest Third of Rio Frio Anois
Sizes of Longest Third of Rio Frio Anolis


SPECIES

humilis



limifrons



carpenter



lemurinus



pentaprion



lionotus



capitol



biporcatus


SEX


C?



o-

c?

9







a"

9!


9i


NL- SVL : (mm)

53 33.22(+0.14)

49 35.76(-0.21)

72 37.15(+0.13)

72 39.59 (0.15)

11 36.50(d0.41)

7 41.50(-0.42)

24 47.79(0.31)

13 57.38(0.47)

4 61.50(11.86)

5 56.80(0.37)

14 65.11(d1.20)

12 58.58(d0.58)

5 75.80(1.10)

4 83.25(-1 .25)

12 88.75(+0.32)

8 90.63(d0.80)


HL (mm)

9.00(d0.06)

9.51(0.05)

9.66(+0.04)

9.94(+0.05)

9.26(0.17)

10.07 (0.15)

12.39(+0.11)

14.57(+0.19)

16.38(+0.50)

14.32(0.19)

15.51(+0.28)

13.86(0.14)

18.52(+0.10)

21.20(+0.34)

23.45(+0.11)

23.91(:0.19)


BW (g)

0.90(0.02)

1.23(0.03)

0.84(0.01)

1.09(0.02)

0.65(d0.03)

1.08(+0.03)

2.28(d0.06)

4.14(0.16)

4.00(+0.14)

3.19(0.04)

4.95(+0.26)

3.92(0.11)

9.16(+0.77)

12.47(0.23)

14.54(+0.19)

17.14(+0.77)


tApproximately 1/3 of all those in each category includes the
longest third plus any others of the same SVL as shortest specimen
in longest third. N (BW) may be less than NL (SVL and HL) if not
all of the longest tHird were weighed.

Given as: mean(standard error).


TABLE 4-1.




















4' K K -I
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-K -I -


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0

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4-4 0
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Mm










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-K -K -I -
-K -I -
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TABLE 4-3. Relative Head Length

SPECIES RELATIVE HLE

MALE FEMALE

lionotus 0.238 0.237

capitol 0.244 0.255

carpenter 0.254 0.243

lemurinus 0.259 0.254

limifrons 0.260 0.251

biporcatus 0.264 0.264

pentaprion 0.266 0.252

humilis 0.271 0.266

EMean HL mean SVL.














SVL Ratios Between Species

SVL (mm) RATIOt


TABLE 4-4.

SPECIES

1. Males:

humilis

carpenter

limifrons

lemurinus

pentaprion

lionotus

capitol

biporcatus

2. Females:


humilis 35.76

limifrons 39.59

carpenter 41.50

pentaprion 56.80

lemurinus 57.38

lionotus 58.83

capito 83.25

biporcatus 90.63

tLonger SVL shorter SVL.

t test, comparing each with next
as in Table 4-2.


33.22

36.50

37.15

47.79

61.50

65.11

75.80

88.75


SIG4


n.s.

***

***

n.s.

***


1.10:1

1.02:1

1.29:1

1.29:1

1.06:1

1.16:1

1.17:1






1.11:1

1.05:1

1.37:1

1.01:1

1.03:1

1.42:1

1.09:1


longer; significance


**

***

***

n.s.

n.s.

***

***
















TABLE

SERIES


4-5. Length

SPECIES

1. Males:


Ratios by

SVL (mm)


Taxonomic Series

RATIO HL (mm)


chrysolepis


fuscoauratus


petersi


chrysolepis



fuscoauratus





peters


humilis

lemurinus

carpenter

limifrons

lionotus

pentaprion

capitol

biporcatus

2. Females:

humilis

lemurinus

limifrons

carpenter

lionotus

pentaprion

capitol

biporcatus


RATIO


33.22

47.79

36.50

37.15

65.11

61.50

75.80

88.75


35.76

57.38

39.59

41.50

58.83

56.80

83.25

90.63


1.44



1.02

1.75



1.23

1.17


1.60



1.05

1.42



1.47

1.09


9.00

12.39

9.26

9.66

15.51

16.38

18.52

23.45


9.51

14.57

9.73

10.07

13.86

14.32

21.20

23.91


1.38



1.04

1.61



1.13

1.27


1.53



1.03

1.38



1.48

1.13















TABLE 4-6. Length Ratios by Perch Site


PERCH SITE



streamside

ground and
streamside


ground


trunk


crown


streamside

ground and
streamside


ground


trunk


crown


SPECIES

1. Males


lionotus

humilis
lionotus
capitol

humilis
capitol


limifrons
lemurinus
biporcatus

carpenter
pentaprion
biporcatus

2. Females

lionotus

humilis
lionotus
capitol

humilis
capitol

limifrons
lemurinus
biporcatus

carpenter
pentaprion
biporcatus


SVL (mm)


65.11

33.22
65.11
75.80

33.22
75.80

37.15
47.79
88.75

36.50
61.50
88.75


58.83

35.76
58.83
83.25

35.76
83.25

39.59
57.38
90.63

41.50
56.80
90.63


RATIO HL (mm)


1.96
1.16


2.28


1.29
1.86


1.68
1.44


1.65
1.42


2.33


1.45
1.58


1.37
1.60


15.51

9.00
15.51
18.52

9.00
18.52

9.66
12.39
23.45

9.26
16.38
23.45


13.86

9.51
13.86
21.20

9.51
21.20

9.73
14.57
23.91

10.07
14.32
23.91


RATIO


1.72
1.19


2.06


1.28
1.89


1.77
1.43


1.46
1.53


2.23


1.50
1.64


1.42
1.67















Weight to Length Relationships


SPECIES

A. humilis



A. limifrons



A. carpenter



A. lemurinus


SEX

3+juv.

p+juv.

c~+juv.

j+juv.


c?


dc


A. pentaprion



A. lionotus 0



A. capitol a



A. biporcatus



NOTE: Regressions are in
or BW=aSVLb.


a

4.66x10-5

3.16x10-5

2.84xl0-5

1.78xl0-5

9.34x-5

3.08xl0-5

3.10x10-5

4.43xl0-5

2.46xl0-5

1.76x10-5

2.93xl0-5

2.07x10-5

4.12x10-5

1.79x10-5

2.61x10-5

0.68xl0-5

the form of


b r

2.82 0.981

2.95 0.985

2.85 0.975

2.99 0.981

2.46 0.981

2.81 0.983

2.90 0.988

2.82 0.992

2.93 0.991

3.00 0.977

2.89 0.995

2.99 0.995

2.84 0.993

3.04 0.996

2.95 0.994

3.27 0.986

In(BW)=ln(a)+bln(SVL)


For A. humilis and A. limifrons small unsexed specimens
were used with both sexes.


TABLE 4-7.















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r --I0

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or deo V C o n C 0 CN

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snyouoTTj
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snufTJntua&





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r-4o oD l o

HOW LA 0


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000




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mP r
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TABLE 4-9.
Sizes of Anoles by Collection Period


PERIOD

A. humilis C
SEP
NOV
FEB
MAY

A. humilis 0
SEP
NOV
FEB
MAY


A. limifrons e
SEP 16
NOV 18
FEB 12
MAY 21

A. limifronsEP
SEP 16
NOV 9
FEB 21
MAY 17

A. lemurinus d
SEP 8
NOV 7
FEB 4
MAY 2

A. lemurinus 9?
SEP 3
NOV 3
FEB 3
MAY 2

A. lionotus (
SEP 1
NOV 4
FEB 8
MAY 2

A. lionotus 0
SEP 0
NOV 5
FEB 8
MAY 1


NLa SVL (mm)


33.09(10.30)
33.17(o.31)
33.39(t0.47)
33.54(10.22)


36.42(+0.48)
35.92(+0.53)
35.07(0.35)
35.78(0.26)


37.00(10.30)
36.33(+0.19)
37.21 (0.13)
38.00(+0.27)


40.13(+0.21)
37.61(+0.35)
39.26(+0.30)
40.82(+0.18)


47.25(0.76)
44.86(12.17)
47.00(+0.58)
48.25(0.25)


59.17(+0.60)
49.17(+4.18)
53.67 (2.17)
58.75(+1.25)


57.0
55.13(3.85)
65.00(+1.53)
70.75(2.25)



51.60(+2.06)
58.69(+0.56)
62.5


HL (mm)


9.14 (0.15)
9.11(00.10)
9.12(10.09)
8.79(10.11)


9.58(+0.07)
9.58(+0.13)
9.44(+0.08)
9.44(10.12)


9.73(0.08)
9.55(0.08)
9.73(0.07)
9.69(0.07)


10.15(+0.09)
9.71(10.09)
9.89(0.11)
10.08(0.06)


12.34 (0.19)
11.84(10.44)
12.18(+0.17)



15.10(+0.40)
13.23(+0.98)
14.10(0.49)
14.55 (0.75)


14.3
13.00(+0.90)
15.58(0.34)
16.90(+0.40)



12.15(+0.05)
13.99(+0.16)
14.3


Nw BW (g)


0.87(+0.03)
0.99(+0.03)
0.96 (0.04)
0.82(+0.03)


1.34(+0.07)
1.29(+0.05)
1.19(0.04)
1.15(+0.06)


0.89(+0.04)
0.83(+0.02)
0.89(0.03)
0.84(0.02)


1.27(+0.04)
1.08(+0.03)
1.05(+0.03)
1.11(0.03)


2.30(+0.11)
2.13(+0.25)
2.22(+0.06)
1.95(+0.05)


4.73(+0.33)
2.71(0.54)
3.35(+0.50)
4.02(+0.48)



3.34(+0.63)
4.94(+0.39)
5.85(+0.44)



2.78(+0.38)
4.03(+0.12)
4.26


%Arrangement as in Table 4-1.

N=2; head length not measured on all A. lionotus











TABLE 4-10.
Significance of Seasonal Differences Size
in Rio Frio Anolis


A. humilis

mean SVL





mean BW


SEP


SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY


n. s.
n.s.
n.s.


*
n.s.
n.s.


NOV

n.s.

n.s.
n.s.

n.s.

n.s.
***


FEB


n.s.

n.s.

n.s.

n.s.
n.s.

*


MAY

n.s.
n.s.
n.s.



n.s.
n.s.
n.s.


A. limifrons


mean SVL





mean BW


SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY


*
***
***


n.s.
n.s.


n.

n.s.
n.s.
n.s.


n.s.
n.s.


***
n.s.

n.s.


**
n.s.
n.s.


A. lemurinus


mean SVL





mean BW


SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY


n.s.
n.s.
n.s.



n.s.
n.s.
*


n.s.

n.s.
n.s.


n. s.
n.s.


n.s.
n.s.

n.s.

n.s.
n.s.

*


n.s.
n.s.
n.s.



n.s.
n.s.
n.s.


A. lionotus


mean SVL




mean BW


NOV
FEB
MAY

NOV
FEB
MAY


*
*



n.s.
*


n.s.


n.s.


NOTE 1: t-test; significance as in Table 4-2.


NOTE 2: Female comparisons are on the upper right, males on
the lower left


















TABLE 4-11.
Seasonal Weight to Length Relationships


SPECIES

A. humilis


A. limifrons


A. lemurinus




A. lionotus


SEX PERIOD


'+j


both




both


SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY

SEP
NOV
FEB
MAY

NOV
FEB
MAY


a

8.07x10-5
-5
2.54xl0
3.79x10-5
1.95x10-5

5.56xl0-5
2.23x10-5
3.19xl0-5
1.14x10-5

4.11x10-5
4.30x10-5
4.33x10-5
0.93xl0-5

1.77xl0-5
1.56xl0-5
3.58xl0-5
0.78xl0-5

1.96x10-5
2.87xl0-5
5.04x10-5
0.39x10-5

2.02xl0-5
2.36x10-5
20.38x10-5


b r


2.66
3.03
2.89
3.04

2.79
3.07
2.96
3.21

2.76
2.75
2.75
3.14

3.02
3.06
2.80
3.20

3.04
2.94
2.78
3.42

2.99
2.95
2.41


0.979
0.985
0.996
0.988

0.985
0.989
0.993
0.993

0.990
0.989
0.985
0.985

0.995
0.991
0.986
0.986

0.988
0.995
0.990
0.958

0.995
0.994
0.968


NOTE: Regressions as in Table 4-7.















TABLE 4-12.
F Test for Seasonal Difference

lis SEP NOV FEB

SEP 5.38* 1.84

NOV 8.28** 1.09


FEB

MAY


A. limifrons

SEP

NOV

FEB

MAY

A. lemurinus

SEP

NOV

FEB

MAY


2.76

6.73*


1.10

0.01


0.15


0.00

0.01

9.01**





0.77

3.12

1.45


0.00

12.89**







2.24

4.11*


1.78


3.50

4.65*


13.38**









6.14*


A. lionotus

NOV

FEB 0.13

MAY 6.11* 2.18

NOTE: Significance and arrangement as in Table 4-8.


A. humi


in b

MAY

9.99**

1.61

7.84**


ovso

0.74

0.15

0.49

2.40



5.46*

8.80**

0.29

0.37


2.19

1.22

15.39**











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ID






0




0








o


r1


0





>
0 *
-,-I4































FIGURE 4-2. Snout-Vent Lengths of Longest Third of Rio
Frio Anolis. Mean SVL indicated by a hori-
zontal line, range by a vertical line, 95%
confidence interval by stippled bar (males,
on right) or cross-hatched bar (females, on
left), and sample size by number above each
figure.











5 8


90

+ 4
sto








70 .





S60 5


55
o 24
o)
50 o


45 7

L4 72 11 49
40 I
S53

35 *

-I 3 0
30 I
4 y.
































FIGURE 4-3. Body Weight of Longest Third of Rio Frio Anolis
Mean BW indicated by a horizontal line, range
by a vertical line, 95% confidence interval by
stippled bar (males, on right) or cross-hatched
bar (females, on left), and sample size by num-
ber above each figure.



























4


121


+


9






0

3 .
5






O ^

(u


20

19

18

17

16

15

14


13

12

11

E-4 10

:z
u9
9


18





0


+4

E
*""
e-


6
.- 9
--I
4
*u
Q)
4

US

Iq:


47




E-
















CO





aQ 0 r'
0 a)

>1
-^





-H U) a)
o>
) 0 -r



* H !
o Q
r.








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0) -


*-C *r


0 C















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to fa
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a )UH a)




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.rl (U
















i > 44
()a) r-4 44)
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61









f 00






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IO ^ cs o oo o or Co o




BW (g)

























0

tr

4.4


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I

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rd
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a)



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C:


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0
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4-1































65



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\ N


























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.
o


















































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BW (9)

























t
*

tr
-H
4-I

r-.

CO
u)













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11




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0
44


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'Fr




























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Ln









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n

BW (g)


























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r-l
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c)
























0
r:
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r&e






































































































BW (g)


* X \


Ln








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L)










in
U,


* i


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O
0
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01



Q)





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O



















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rZ4
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4-








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3,

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r:








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*<
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* *


BW (g)


0







r-






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C*L
Ln






* LA

Er


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U)


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En


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0








3
tr











































C;
f-
0







4-
t0


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'-I






H
r4
0}




















0



































0











Ln
.o






0










o












m


BW (g)


0* *


























-4
o,-I
t-4












rl
Ul


O



















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0(
V-I








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3
u
















Uv
a.




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C-I






01
><-








CQl



















*ox\


o W N
(1 r-


Ln
LfA


BW (g)































FIGURE 4-12.


Seasonal Variation in Size of Anolis humilis.
For the longest third of each sample, mean
body weight (BW) and mean snout-vent length
(SVL) are given as horizontal lines.
Stippled bars indicate 95% confidence inter-
vals; vertical lines show the range; and
sample sizes are given by numerals above the
range.








1.8


1.6


1.4


1.2.


1a. o


0.8


0.6





41


39


37


35


33


31


8

12


.


SEP NOV FEB MAY
FEMALES


14 8
13








SEP NOV FEB MAY
MALES










9
15 13









SEP NOV FEB MAY
MALES


SEP NOV FEB MAY
FEMALES


12 15



9
































FIGURE 4-13. Seasonal Variation in Size of Anolis limifrons.
Symbols are as in figure 4-12.







1.6


1.4 a


1.2


ml1.0 -


0.8 -


0.6 "


SEP NOV FEB MAY
MALES


16 2



18 12



+4+


SEP NOV FEB MAY
MALES


SEP NOV FEB MAY
FEMALES


16 17
21



9

f [


SEP NOV FEB MAY
FEMALES


20
7
11 '
6 21
S174


42


40 *


38 u


36 a


34 .

































FIGURE 4-14. Seasonal Variation in Size of Anolis lemurinus.
Symbols are as in Figure 4-12.

















2

I-

0




65

60 -

55

50

45

40

35*


2

+ 2
3 3+
4-








SEP NOV FEB MAY
FEMALES


3-
-f-


SEP NOV FEB MAY
MALES


2
34-

4-


SEP NOV FEB MAY
FEMALES


SEP NOV FEB MAY
MALES


ff-- ::=


8 7 2

Ut4

































FIGURE 4-15. Seasonal Variation in Size of Anolis lionotus.
Symbols are as in figure 4-12.


















I-b


2


NOV FEB MAY
?ALES


+


MAY


NOV FEB


FEMALES


ii


4


NOV FEB MAY
MALES


8

5+


NOV FEB MAY
FEMALES


-4
Im


75 -


70


65


6-60


55


50

45


--em-


--v-


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0
S








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0 <- (
















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rl rl -4 0 0 0 0
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BW (9)


87













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tP



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r-H

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or


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rz:











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0










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0









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+3
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H













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(UDa







(0*
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Er













i 0


Sin







.0







\ 0
S l*















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ti







U)

LA


BW (g)