Citation
The thermal biology of the Turks and Caicos Islands rock iguana Cyclura carinata

Material Information

Title:
The thermal biology of the Turks and Caicos Islands rock iguana Cyclura carinata
Creator:
Auth, David L
Publication Date:
Language:
English
Physical Description:
xvi, 367 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Body temperature ( jstor )
Cloud cover ( jstor )
Cooling ( jstor )
Ecology ( jstor )
Female animals ( jstor )
Heating ( jstor )
Iguanas ( jstor )
Lizards ( jstor )
Neurons ( jstor )
Oxygen ( jstor )
Iguanas ( lcsh )
City of Gainesville ( local )
Genre:
bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis--University of Florida.
Bibliography:
Includes bibliographical references (leaves 347-366).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by David Leslie Auth.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Resource Identifier:
023376857 ( ALEPH )
AAL3883 ( NOTIS )
06750362 ( OCLC )
AA00004907_00001 ( sobekcm )

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Full Text


THE THERMAL BIOLOGY CF THE TURKS AND
CAICOS ISLANDS ROCK IGUANA CYCLURA CARINATA
By
DAVID LESLIE AUTH
ISSERTATION PRESENTED TO THE GRADUATE COUNCI
THE UNIVERSITY OF FLORIDA
PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1980


ACKNOWLEDGEMENTS
I am grateful to numerous members of the faculty of the University
of Florida: to Walter Auffenberg, Lewis Berner, Robert DeWitt, Thomas
Emmel, and Howard Wallace, for their faith in my competence as a zoolo
gist; to Archie Carr, John Kaufmann, Brian McNab, and Hugh Popenoe, for
their forebearance, patience, and time as members of the Ph.D. committee
to John Anderson, John Brookbank, William E. S. Carr, Thomas Hollinger,
David Johnston, James Nation, Michael O'Rand, and Ronald Wolff, for the
pleasure of being a laboratory teaching assistant for their courses; to
Gerald Olsen, for introducing me to the anesthetic halothane; to Robert
DeWitt, John Swel, Brian McNab, and Horst Schwassmann, for use of equip
ment; and to James Heath, whose work and brief personal contact started
my interest in lizard thermoregulation.
I thank the New York Zoological Society for the financial support
necessary to accomplish the research. The University of Florida has
provided extensive support throughout my graduate career, in the form of
teaching assistantships in the Zoology Department ar.d the College of
veterinary Medicine. Educational assistance through the G.I. Bill and
the Naval Reserve proved essential.
John Iverson introduced me to the habitats and habits of Cy atura
aavinata and continued to aid me in many ways during the study. Diderot
Gicca, Dagmar Werner, and Thomas Wiewandt willingly shared their knowl
edge of other species of Cyatuva.
ii


C. W. (Liam) Maguire and Bill and Ginny Cowles of the Meridian
Club, Pine Cay, permitted me to stay for practically nothing in the lap
of luxury while working in the field. They were most hospitable hosts.
Gaston Decker, Chuck and Cathy Hesse, and Francoise de Rouvray, also of
Pine Cay, helped me in many ways,
Alan Bolten, Jon Baskin, and William Link were superior companions
and considerate office mates. Alexis Arends, Susan Barnard, Hugh Ellis,
Michael May, Ken Prestwich, and Perran Ross gave needed technical
assistance. Karen Bjorndal, David Deitz, Dale Jackson, Howard Kochman,
Anne and Peter Meylan, and Michael Marshall all provided moral support.
I owe the greatest debt to my mother, Mildred G. Auth, my father,
Eugene F. Auth, and my brothers, Ronald and Dennis.
iii


TABLE OF CONTENTS
ACKNOWLEDGEMENTS ii
LIST OF TABLES vii
LIST OF FIGURES ix
ABSTRACT xiv
INTRODUCTION
1
METHODS AND MATERIALS 7
ECOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CYCLUEA CARINATA ... 24
Climate of Water Cay 24
Diel Body Temperature Cycle of a Heliothermic Lizard 31
Regulating Phase Body Temperature in the Field 34
Temporary Hyperthermia and Hypothermia 44
The Relationship of Pertinent Environmental Variables
to Regulating Phase Body Temperatures 55
Season
Habitat Complexity .
Cloud Cover
Shaded Air Temperature
Substrate Temperature.
55
62
67
73
75
Patterns of Heat Transfer between Penned Iguanas and
their Environment: The Three-Transmitter Study 78
Statistical Analyses of Regulating Phase Body Temperature. . 89
Body Temperature Maxima and Minima ..... 39
Skewness of Body Temperature Frequency Distribution. . 107
Standard Deviation of Mean Body Temperature 112
Rates of Lizard Heating and Cooling in the Field 119
Morning Heating and Regulating Phase 119
Lizard Cooling in Burrows 125
ETHOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CYCLURA CARINATA. . 132
Lizard Activity 132
Burrow Movement 132
Control of Lizard Emergence and Submergence 132
iv


Morning Heating Phase 146
Regulating Phase 149
Foray Movement and Body Temperature 154
Feeding, Transmitter Passage Time, and Defecation. . 166
Drinking 168
Nasal Salt Gland Secretion 170
Time-Motion Study of Penned Lizards 171
Behavioral Thermoregulation 174
Description of Thermoregulatory Movements and Postures 174
Shuttling Behavior 184
Sun Orientation 187
Thigmothermia 200
Low Body Temperature Behavior 213
High Body Temperature Behavior 216
Head-Body Temperature Differences 224
The Integument: Metachromatism and Reflectivity 227
The Integument: Sensory Spots 231
PHYSIOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CY CUJEA CAMINATA. 236
Heating and Cooling Rates 236
Aerobic Metabolism 243
Heart Rate 253
Recovery from Maximum Exercise 259
DISCUSSION 270
Body Temperature Relations 270
Comparison of Cyclura oarinata's Mean Body
Temperature with Other Diurnal Lizards 270
Seasonal Variation in Body Temperatures of
Active Lizards 276
Stability of the Mean Preferred Body Temperature .... 277
Late Afternoon Maximum Body Temperature 279
Thermal Safety Margins 281
Panting Threshold 282
Body Temperature and Reproductive State 284
Ethological Considerations 285
Lizard Activity 285
Eurythermy vs. Stenothermv 288
Comparative Thermoregulatory Behavior of
Heliothermic Lizards 296
Complexity of Social Behavior in Cyclura aavinata
as a Function of Latitude 305
Nocturnal Lizard Aggregations 306
v


Physiological Considerations 307
Aerobic vs. Anaerobic Metabolism 307
Neural Basis for Shuttling Behavior in Lizards 312
SUMMARY OF RESULTS AND CONCLUSIONS 326
LITERATURE CITED 347
BIOGRAPHICAL SKETCH 367
vi


LIST OF TABLES
Table
1 Daily variation in relative humidity 30
2 Summary of regulating phase body temperatures of
lizards in the field 35
3 Hyperthermia in Cyolura carinata in the field 47
4 A seasonal comparison of some environmental variables
at Site II on Water Cay 56
5 Seasonal changes in regulating phase body temperature
in the field and in the laboratory thermal gradient .... 57
6 Vegetational cover at Sites I and II on Water Cay 66
7 Body temperatures of Cyclura car-Lnata confined to
cages with two structurally different habitats 68
8 Skewness of frequency distributions of minimum and
maximum body temperatures 91
9 Means and standard deviations of minimum and maximum
body temperatures of individual lizards and correlation
of maximum vs. minimum and minimum vs. maximum pairs . 92
10 Linear regression analysis of daily minimum vs.
maximum Tfc's and mean of daily Tfo minima vs. mean of
daily Tjy maxima for individual lizards 101
11 Skewness and kurtosis of Tfo frequency distributions .... 118
12 Rates of lizard heating and cooling in the field during
the morning heating and regulating phases 120
13 Lizard cooling in burrows in the field 124
14 Lizard body temperature at emergence in the field 131
15 Emergence and submergence times, temperatures, and
light intensities for free-ranging lizards in the
field 146
16 Possible acclimatization of lizard activity 165
vii


17 Effect of body temperature on the feeding rate and meal
size of penned Cyctura oavina.ta 167
18 Time-motion study of Cyclura aari-nata in the Gainesville
pen 172
19 Skin patch heating 198
20 Estimated thermal conductance during heating and cooling
of lizards in still air 240
21 Correlations of lizard behaviors with body temperature
during heating and cooling in the cage and on the jig . 245
22 Recovery oxygen for Cyclura carinata after seven minutes
of maximum exercise 265
23 Comparison of actual mean recovery time after maximum
exercise with recovery times at three hypothetical rates
of oxygen utilization 267
24 Field body temperatures of diurnally active lizards .... 271
viii


LIST OF FIGURES
Figure
1 Black body temperatures in the field 25
2 Air temperature in the shade at Gainesville, Florida
and Grand Turk, British West Indies 26
3 Mean maximum wind speed at Site II in July 28
4 Monthly precipitation on Grand Turk, British West Indies. 29
5 Body temperature record of Lizard 2 on February 10, 1976. 32
6 Summary of field body temperatures during the regulating
phase 45
7 Lizard hyperthermia and hypothermia during the regulating
phase 46
8 Patterns of regulating phase body temperature and distance
traveled in Cyctura occrznata in a laboratory thermal
gradient 51
9 Hypothermia in a free-ranging lizard 54
10 Seasonal change in Cyclura regulating phase body
temperature in the Caicos and Gainesville, Florida 59
11 Relationship of mean body temperatures in the field and
the thermal gradient for active lizards from several
families 61
12 Influence of habitat on body temperature 65
13 Effect of cloud cover on mean regulating phase body
temperature 69
14 Effect of overcast duration on lizard body temperature
change 72
15 Correlation of lizard body temperature during the
regulating phase with shaded air temperature 74
16 Thermoregulatory flexibility 77
ix


17 Three transmitter record of a large male Cyolura
oarinata in the Gainesville pen on August 21, 1975 80
18 Partitioning of heat transfer in active penned
Cyolura oarinata 84
19 Rate of core body temperature change during the four heat
transfer conditions 86
20 Correlations of the sum of daily heat transfer conditions
with lizard activity and mean body temperature 88
21 Time distributions of four combinations of maximum and
minimum body temperature 97
22 Correlations of maximum vs. minimum Tfo and the mean of
Tfo maxima vs. minima for individual lizards 100
23 Hourly change in minimum and maximum body temperatures. . 104
24 Timing shift of daily minimum and maximum Tfo1 s 106
25 Skewness of daily Tjy frequency distributions 110
26 Comparison of the level and precision of thermoregulation
of Cyolura oarinata and Dipsosaurus dorsalis in continuous
thermal gradients Ill
27 Correlation of the standard deviation of mean daily Ty
with mean daily Ty 113
28 Effect of body weight and mean Tfo on the standard
deviation of mean Tfo 116
29 Hourly change in the mean and standard deviation of lizard
body temperature and shaded air and sand temperatures . 118
30 Correlation of mean rates of heating and cooling in the
field with lizard body weight 124
31 Correlations of parameters describing regulating phase
body temperatures 126
32 Lizard movements in a burrow 134
33 Seasonal change in lizard emergence and submergence times 136
34 Correlations of times of emergence and submergence with
body weight in penned Gainesville lizards 140
35 Gradual reentrainment of lizard emergence time after
release 145
x


36 Lizard activity during the morning heating phase 148
37 Daily and seasonal variation in activity 151
38 Variable inhibitory effectiveness of cloud cover on
lizard activity 155
39 Influence of cloud cover on the length of lizard forays
and the mean rate of travel during forays 159
40 Correlations of foray length and rate of travel with mean
body temperature 161
41 Body temperatures during forays 163
42 Defecation body temperatures and timing 169
43 Correlation of lizard mobility with confinement area. . 173
44 Thermoregulatory motions and postures of Cyolura oarinata 175
45 Lizard shuttling in the summer field cages 185
46 Midday depression of shade seeking body temperature .... 188
47 Lizard orientation to a heat lamp during heating and
cooling in the photothigmotron 189
48 Positive and negative orientation during lizard heating
and cooling 192
49 Daily and seasonal variation in sun orientation in the
Gainesville pen 195
50 Left vs. right side sun orientation in the Gainesville
pen 196
51 Daily change in the pattern of travel during forays .... 199
52 Participation of the lizard's head, neck, and legs in the
reduction of substrate contact 203
53 Relative importance of lateral compression of the trunk
vs. head, neck, and trunk elevation 205
54 Conductive and convective heating and cooling of a single
Cyolura oar-inata 206
55 Conductive heating and cooling of a single Cyolura
carinata, with and without simultaneous radiative
heating with a heat lamp 209
xi


56 Thigmothermic responses of a single Cyclura oarznata to
periods of increasing, stable, and decreasing substrate
temperature 212
57 Behavioral responses of Cyolura oarznata at low body-
temperatures 214
58 Daily adjustment of the panting setpoint to changing
heat input 220
59 Correlation of total panting time with head and gut
thermal lag times 222
60 Head-body temperature differences during heating and
cooling of a 1790 g male Cyolura oarznata in the
Gainesville rooftop arena 226
61 Integumental and substrate reflectivities 228
62 Density of sensory spots on different parts of the lizard
body surface 234
63 Heating and cooling rates of a 1435 g male Cyolura
oarznata in a constant temperature chamber 239
64 Breathing rates of lizards during heating and cooling in
a constant temperature chamber 247
65 Correlation of resting and maximum oxygen consumption
with body temperature 250
66 Analyses of aerobic metabolism 252
67 Heart rate as a function of body temperature 254
68 Heart rate analyses 256
69 Relationship of oxygen consumption to heart rate at
different body temperatures 257
70 Temporal pattern of body movements while struggling to
escape 260
71 Effect of body temperature on the rate of recovery from
seven minutes of maximum activity 263
72 Early and late phase recovery from maximum exercise in
Cyclura oarznata at different body temperatures 269
73 Summary of body temperatures of selected behaviors and
physiological maxima of Cyclura oarznata 293
xii


74 Effect of increasing body temperature on frequency
distribution skew and standard deviation of the mean
for minimum and maximum body temperatures
75 Hypothetical neural basis for changes observed during
increasing mean body temperature in Cyolura carinata. .
. 319
. 324
xiii


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
THE THERMAL BIOLOGY OF THE TURKS AND
CAICOS ISLANDS ROCK IGUANA CYCLURA CARINATA
By
David Leslie Auth
June 1980
Chairman: Walter Auffenberg
Major Department: Zoology
The ecological, ethological, and physiological thermal biology of
Cyclura carinata, a large, tropical, herbivorous iguanine lizard, was
investigated on Water Cay in the Caicos Islands and in Gainesville,
Florida. The study lasted from June, 1974, until September, 1976, and
included four field trips totaling twenty weeks.
Far-field transmitters were placed in the coelom or gut of free-
ranging adult iguanas (675-1864 g) for long-term monitoring of core
body temperature (T^) in the field. Mean regulating phase T^ ranged
from 38.0 to 39.7 C (February-October), quite high compared with most
other iguanids. Stenothermia was pronounced, with a mean, regulating
phase range of only 3.3 C. Maximum voluntary T^, 43.8 C, was only
2.4 C below the CTMax. The CTMin, 12.8 C, almost equalled the lowest
recorded environmental temperature.
Iguanas were initially hyperthermic in both the field and a
thermal gradient, the former the possible result of bacterial infection
or a transmitter meal and the latter of thermoregulatory learning.
xiv


Individual differences in regulating phase T^ were partially due
to variable vegetational cover and, perhaps, egg development in females.
Mean operant T^'s did not correlate perfectly with seasonal change in
the ambient thermal environment. Mean regulating phase field T^
exceeded mean preferred TThe latter did not change with Gainesville
seasonal acclimatization, whereas winter variance of mean T^ was less
than summer variance in both field and thermal gradient.
Mean regulating phase T, was inversely correlated with cloud cover
u
time, the correlation improving with seasonal cooling. T^ was nearly
independent of shaded air temperature.
Heat transfer through the dorsal and ventral body was examined
using three transmitter placements on penned iguanas. Heat was usually
gained dorsally and lost ventrally; iguanas were thigmothermic only 20.7
percent of their active period, usually in combination with heliothermia.
Individual frequency distributions of regulating phase maximum and
minimum T-, s were commonly skewed positively and negatively, respectively,
whereas inclusive individual distributions were usually negatively
skewed. Standard deviations were slightly but not significantly greater
for mean minimum than mean maximum T-^' s. Change in T^ between any two
consecutive maxima was positively correlated with change in two
consecutive minima, with the first minimum immediately before or after
the first maximum. Individual mean maximum and minimum T^'s were
directly and linearly correlated, the regression line extrapolating
to approximately the maximum voluntary tolerance. These results are
discussed in terms of dual-limit thermoregulation.
Iguanas were diurnally active all year and rarely escaped into
burrows due to overheating. Mean regulating phase heating was 1.15


times faster than cooling and 3.0 times slower than heating during
morning basking. Iguanas also heated faster than they cooled in a
constant temperature chamber, both when restrained and when permitted
to move about. Maximum T^ commonly occurred during late afternoon
basking. This and other results indicate that surface rather than
core body temperature is regulated.
Both length and rate of travel were independent of mean T^ between
35.0 and 40.5 C. Arboreal climbing, burrow excavation, defecation,
drinking, feeding, nasal salt gland secretion, and panting were also
examined in relation to mean T-, .
b
Thermoregulatory behaviors and postures were described. A
photothigmotron was constructed to quantify iguana light shadow and
substrate contact in a controlled thermal environment. Shuttling and
sun orientation were examined. A time-motion study revealed the low
activity rate of penned iguanas and activity partitioning between
foraging, social, and thermoregulatory behaviors. Thermoregulatory
behavior was quite similar to other heliothermic iguanids.
Resting and maximum oxygen consumption, and heart rate and
oxygen consumption during recovery from maximum exercise were determined
at different 's. Iguanas fatigued rapidly during maximum exercise,
yet rarely developed large oxygen debts in the field. Heart rate
increment, initial rate of recovery, and oxygen scope were greater
in the preferred T^ range.
xvi


INTRODUCTION
Although much is known regarding the thermal biology of small
lizard species (see Heatwole 1976 for a review), large lizard species,
because of their rarity in numbers and usual distribution in isolated
areas, have received lesser attention. This is unfortunate, for large
mass confers certain advantages and disadvantages due to low heating
and cooling rates. This study investigates the thermal biology of a
species of rock iguana in the genus Cyalura Harlan, one of nine genera
in the subfamily Iguaninae (Avery and Tanner 1971). Although the genus
includes some of the largest lizards in the world, Cyalura aarinata is
one of the smaller members (maximum weight of 1,864 g for males,
1,135 g for females).
Preliminary observations by Auffenberg (personal communication) in
1973 indicated that Cyalura caminata probably had a fairly high diurnal
operant or eccritic body temperature (2^). This, coupled with large size and
a herbivorous diet, uncommon in lizards, suggested that its thermal biology
could in some ways be unique. A great deal of thermally related data had
already been collected for the desert iguana, Dipsosaurus dorsalis, a. small
herbivorous lizard of the North American Southwest with a very high
preferred (Porter et at. 1973). Thermally related comparisons of
Cyalura aarinata with this species as well as with the larger iguanines,
Amblyrhynahus cristatus (White 1973), Iguana iguana (McGinnis and Brown
1966), and Sauromalus obesus (Johnson 1965) seemed worthwhile.
Continuous monitoring of the T^ of free-ranging lizards in the
field using radio-telemetry forms the basis for the present study. This
1


2
approach has only rarely been followed for large lizards (Stebbins and
Barwick 1968, McNab and Auffenberg 1976).
The natural history of Cyalura aarinata was examined simultaneously
from four directions by Dr. Auffenberg (feeding strategy), John Iverson
(autecology and ethology) and myself, under a grant from the New York
Zoological Society. The field site was located primarily on Pine and
Water Cays, on the northwest side of the Caicos Island Bank. Since the
goal of this research was to gain as much information as possible which
could be useful in guaranteeing the species' future survival, I chose to
attack the problem of thermal biology broadly, rather than emphasizing
any one aspect such as thermoregulation. Ecological, ethological, and
physiological approaches were taken, working only at the organismic
level and using free-ranging field, controlled field, and laboratory
studies. This synthetic approach has been utilized rather successfully
by Grenot (1976) for Uromatix aaanthinurus, another fairly large
herbivorous lizard.
The field ecological approach to Cyalura aarinata's thermal
biology depends on a thorough understanding of the thermal environment
in which the lizard lives. This means that cloud cover, wind speed,
black body temperature, shaded air temperature, and substrate tempera
ture have to be monitored daily and seasonally as well as the lizard's
body temperature and its thermally related behavior. Then, environ
mentally related questions can be answered. How does the lizard
respond to the rather constant easterly winds? Are these xeric habitat
animals able to stay out of their burrows even during the hottest part
of the summer day or are they forced underground, as is the smaller


3
Dipsosaurus dorsalis (Porter et al. 1973)? Is T^ independent of shaded
air temperature, as expected in a heliotherm? The populational dis
tribution on the islands should be examined in terms of habitat types
available and any variation in mean T\ with vegetational density
determined. The iguana distribution (before the beginning of the study)
was minimally affected by introduced mammalian competitors, since the
human population was small and the only other mammal was the introduced
Norway rat.
Continuous T^ records of free-ranging lizards provide a large
amount of analyzable data, as shown by Stebbins and Barwick (1968) from
their monitoring of a single lace monitor lizard. Maximum and minimum
T-0's during the regulating phase, times of emergence and submergence,
and heating and cooling rates can easily be extracted from the record.
Then, many interesting questions can be addressed. How do the maxima
and minima vary during the day on clear days? Does body temperature
variability decrease as body weight increases? Does this variability
decrease as mean T^ increases? How is mean related, if at all, to
reproductive state, sex, and size?
Heath (1964) and DeWitt (1967) were two of the first investigators
to report a significant temperature differential between the head and
body of lizards during heating and cooling, calling into question the
significance of measuring just cloacal temperature and bringing up the
question of what body temperature is regulated. Since then, many
workers have simultaneously measured more than one body temperature in
lizards (Johnson 1972, Webb, Johnson, and Firth 1972, Crawford 1972,
Spray and Belkin 1972, 1973, Crawford and Barber 1974, Parmenter and


4
Heatwole 1975, Pough and McFarland 1976, McNab and Auffenberg 1976,
Crawford et at. 1977). For a large lizard like Cyolura earznata,
multiple body temperature measurements are advisable as part of the
study. Head-gut and dorsal black body-gut-ventral body surface tempera
tures were measured to determine internal temperature differences and
dorsal and ventral heat transfers with the environment, respectively.
Lizards had to be confined in outdoor pens to maintain continuous visual
contact, a serious problem with free-ranging lizards.
Two aspects of the ethological thermal biology of Cyolura
oavznata are investigated. The first addresses the question of how
daily variation in the thermal environment and T^ influence lizard
activity. Lizard movement in burrows, emergence and submergence timing,
movement during the morning basking and regulating phases, feeding rate,
total food consumed, passage time of transmitters through the gut,
defecation timing, and secretion timing of the nasal salt gland are
considered. Lizards vary considerably in the amount of time spent
in actual motion. "Widely foraging" predators, such as varanids and
teiids, are generally more active than "sit and wait" insect feeding
iguanids (Pianka 1973). Herbivorous lizards should fall somewhere in
between, if they feed on a variety of plants, since they must go to
where the plants are located and then actively pick items up. The
partitioning of the lizard's active time between feeding, social, and
thermoregulatory movements should vary consistently between each of
these three dietetic categories. Ideally, a continuous time-motion
study should be conducted in the field, as accomplished for Egermza
aunn-ngham-i by Wilson and Lee (1974). However, this proved impossible


5
from a fixed blind, due to vegetational density and the considerable
distance traveled by adult iguanas. Penned lizards were observed
instead.
The second ethological approach is to discover how Cyclura
carinata regulates its body temperature and then to compare this
behavior with that of other lizards, reported on extensively in the
literature. This has traditionally been the emphasis of lizard thermal
biology studies, since the time of Cowles and Bogert (1944). The roles
of thermoregulatory postures were comprehensively catalogued in a
qualitative manner, similar to Brattstrom's work on Amphzbolurus
barbatus (1971) and Greenberg's on Sceloporus cyanogenys (1977).. Some
of the behaviors were quantified under controlled field and laboratory
conditions. A laboratory device was built to quantify the degree of
sun orientation and thigmochermia, using a photographic technique
similar to Heath's for Phynosoma (1965). I did not attempt to find out
to what extent body posture controlled T-, under a defined set of
thermal conditions, as Muth (1977b) and Porter and James (1979) have
recently done.
Lizards usually heat faster than they cool (Bartholomew and
Tucker 1963, 1964, Morgareidge and White 1969, Weathers 1970, Weathers
and Morgareidge 1971, Weathers 1971, Baker et at. 1972, Ellis and Ross
1977). This is attributed to increased peripheral blood flow during
heating. The ratio of heating to cooling rate increases as body mass
increases in the American alligator, to the extent that while hatchlings
heat and cool at about the same rate, a 1000 kg adult would theoretically
heat nearly three times faster than it cools (Smith 1979). The testing


6
of Cyclura carinata is justified on comparative grounds and because of
its large size, especially since the data obtained complement the
heating and cooling rate results for the species in the field.
Many rate processes become somewhat temperature independent in
reptilian preferred T^ ranges (see Dawson 1975 for a review). Selected
physiological parametersresting and maximum oxygen consumption, rest
ing and maximum heart rate, and oxygen consumption during recovery from
maximum exercisewere measured in Cyclura cari'riata to see how they
varied with The study of recovery from maximum exercise was
especially interesting, since the amounts and kinds of voluntarily
tolerated, strenuous activity the species engaged in were observed in
the field. Bennett's methodology (1972) for his comparison of oxygen
consumption, oxygen debt, and heart rate in Varanus gould't'i and
Sauromalus kispidus was closely followed, permitting comparison
with results obtained for Cyclura.


METHODS AND MATERIALS
The field study site was located in the Caicos Islands, between
21 and 22 North latitude and 71 30' and 72 30' West longitude,
approximately 150 km north of Hispaniola and 890 km southeast of Miami,
Florida. Pine Cay was the base of operations, a privately owned island
with excellent lodging, potable water, an air strip, and a large popu
lation of Cyalura cari,nata (as of August 1973). Field work was conducted
primarily on Water Cay, connected by a traversable sand bridge to Pine
Cay. Iguana density was adequate here at the beginning of the study, no
human habitation was present, and bulldozed paths provided easy access to
most areas at the northeast end of the island. The two islands are
located on the northern perimeter of the triangularly shaped Caicos
Bank, a shoal with a northern base of 125 km and a southerly directed
altitude of 75 km. Water Cay has an area of approximately 225 ha, with
a leeward northwestern sandy beach, a windward eastern rocky limestone
coast, and a southern mangrove swamp. The height and density of the
vegetation increases with island age and soil depth and quality from the
northwestern to the southeastern side of the island. The highest point
is approximately five meters above sea level.
I spent a total of 140 days in the field during four trips:
June 1-July 28, 1974; September 30-November 4, 1975; February 4-23, 1976;
and August 17-September 11, 1976.
Two field sites were chosen on the. eastern end of Water Cay. Site I
was in level, sandy, Semiopen Scrub, 600 m from the NE side and 275 m from
7


8
the NW side of the island. Site II was in Dense Scrub, with Rocky
Coppice bordering the Pine-Water Cay inlet, 200 m from the SE corner of
the island (see Auffenberg, in prep., for descriptions of vegetation
types). Work was confined to approximately one hectare at each site.
Site I bordered a bulldozed road and Site II was crisscrossed by an
interconnected road network.
A moderately large hotel complex was completed on Pine Cay in the
summer of 1973. By the time of my first trip, iguana populations had
already declined on both Pine and Water Cays due to predation by man,
dogs, and feral cats. The subsequent collapse of the population on Pine
Cay has been documented by Iverson (1977). Cyclura density at Site II
was estimated at 10 adults per ha in July, 1974. Density was somewhat
less at Site I. Iguanas had practically disappeared from Site I by
September, 1975, necessitating a concentration of effort at Site II.
Iguanas had been extirpated from Site II by August, 1976, resulting in
a shift of field work further southwest along Water Cay and to Fort
George Cay, just north of Pine Cay. Due to the small number of hatchling
and juvenile iguanas, the study had to be restricted to adults.
Eleven male (X = 993 g, 435-1715) and four female (X = 748 g,
528-1135) iguanas were brought to Gainesville, where they were kept in
two outdoor galvanized sheet metal pens. One pen was circular, enclos-
? 2
ing an area of 16 m and the other was rectangular, enclosing 18 m .
Eight plywood boxes, each measuring 40 cm wide by 40 cm long by 10 cm
high, were perched on concrete blocks and provided shade, shelter, and
basking platforms. Substrates were beach sand and grass in the circular
and rectangular pens, respectively. From October 1 to March 30, the


9
pens were partially or completely covered with 4 ml polyurethane plastic
to increase temperatures. An electric forced air heater kept temperatures
above freezing on exceptionally cold nights. The captives were fed
bananas, green beans, white grapes, and fish-flavored catfood.
Several environmental variables were monitored in the field.
Measurements were taken hourly at a single location near the blind at
Sites I and II, where an open, sun exposed area bordered an area of
fairly dense vegetation. Changing temperatures between readings were
estimated to occur linearly. Substrate temperatures were taken on dry
sand with a Number 421 banjo probe attached to a Model 46 Tele-Thermometer
(Yellow Springs Instrument Co.). Air temperatures were taken at a height
of 2-3 cm with a Number 405 air probe. A black body for estimating
daily variations in solar heat input was constructed by placing a
mercury thermometer in a groove carved in a balsa wood block. The block
was then covered with a thin brass plate (0.4 mm) and coated with flat
black paint. A record was kept of the times the sun was obscured by
clouds. Light intensity was measured with a Type 213 light meter
(General Electric) directed from a height of 20 cm at a piece of white
painted cardboard placed on the ground. Instantaneous wind speed was
measured with a Dwyer wind meter. Average wind speed was determined
over a one minute period with an Air Meter Model W131 (Weather Measure
Corp.). Relative humidity was measured with a sling psychrometer.
Core body temperatures of free-ranging iguanas were measured with
Model L far-field transmitters (Mini-Mitter Co.). The transmitter was
a crystal-controlled blocking oscillator operating in the citizen's
band range (27 MHz). Osgood and Weigl (1972) describe the circuit diagram


10
and construction. The signal, a series of beeps increasing in frequency
with temperature, was received with a Model HA-420 three-channel walkie-
talkie (Lafayette) with an attached beat frequency oscillator. Trans
mitter range was approximately 200-300 feet, depending on the topography.
Accuracy was +0.5 C. The transmitter was powered by two hearing aid
batteries (Mallory MS 76), which had a working life of one to two months.
The transmitter was embedded in epoxy resin and enclosed in a plastic
capsule. The capsule was then coated with a mixture of 50 percent
paraffin and 50 percent beeswax, followed by dental acrylic (NuWeld,
I.D. Caulk Co.). The package weighed about 15 g and was smaller (1.7 by
3.7 cm) than a Cuolura egg. A single mercury thermometer (Scientific
Products), checked for accuracy against a NBS certified bomb calorimeter
thermometer (Parr Instrument Co.), was used to calibrate all transmitters
used in this study. Most transmitters were recovered after the monitor
ing period. Drift never exceeded 0.5 C.
During the summer field trip of 1974, a transmitter was implanted
in each of eight adult iguanas. The operation was performed during the
night of the day of capture. The lizard was fastened to a wooden jig
similar to that used by Bartholomew and Tucker (1963). The individual
was then placed in a large freezer cabinet until body temperature had
fallen to 2-5 C. T^ was monitored intestinally with a Tele-Thermometer
Probe Number 701 inserted approximately 10 cm into the distal colon
and taped to the tail. The individual was kept on ice during the
operation. The transmitter was inserted through a dorsoventral
incision made on the right lateral abdominal wall about 1 cm anterior
to the. rear leg. The incision was closed with nylon fishing line.


11
Individuals were released the day following the operation and monitored
immediately. During subsequent field trips, transmitters were placed
in the gut immediately after capture. A gauze pad soaked with halothane
(Ayerst Laboratories) was positioned at the bottom of a 250 ml beaker.
See McDonald (1976) for a discussion of this inhalation anesthetic. The
beaker was then slipped over the lizard's head. A rubber apron taped
to the mouth of the beaker extended over the forelegs, reducing the rate
of loss of gas anesthetic. Lizards were usually apneustic when the mask
was initially placed over their heads, the duration of apnea being
inversely correlated with body temperature. The drug took effect within
a minute after breathing commenced. Complete anesthesia lasted between
five and ten minutes, and recovery was seemingly complete within an hour.
All individuals were weighed (Ohaus Dial Spring Scale), measured (snout-
vent and total length), examined for scars and ectoparasites, and marked
on both sides of the trunk with a bright orange Arabic numeral (Aero-
gloss paint, Pactra Industries).
The body temperatures of eleven different, free-ranging adult
iguanas were monitored during the study (10 males, 1 female). Two males
were monitored on two separate trips. Recording periods extended from
three to twenty days. Temperature readings were taken every five to
fifteen minutes from before the individual emerged up to two hours
after it submerged for the day.
I observed and monitored iguanas from a blind during the first
field trip. Lizards were attracted to the vicinity with a sardine bait
and noosed from inside the blind. Less timid individuals were noosed
without bait and blind.


12
Iguanas were less timid during che second and third trips, per
mitting more thorough observation of specific individuals by continuous
tracking (6 adult males tracked 1-6 days, X = 4 days). Following initial
transmitter placement, iguanas were monitored without further visual
contact until the start of the second or third day. Since iguanas
usually retreated into their burrows when disturbed during the initial
basking period, visual contact was always delayed for a time after
emergence. I crawled along behind the lizard at a distance of approxi
mately five meters during its feeding forays. Visual contact was
intermittently lost when the individual moved off into dense scrub. The
lizard was usually quickly resighted by following the same direction of
travel. Signal strength, as perceived by the human ear, reached a
maximum approximately three meters from the transmitter. Bushes along
the route of travel were flagged with orange plastic tape marked with the
dace and time. A crude map of the route was recorded in a notebook,
along with observations of the individual's behavior. The locations of
the flags were later marked on a detailed map of the site. Average
rates of travel and distances traveled could then be calculated at
leisure.
Near-field transmitters, built according to Mackay (1970), were
used during the study. They were cylindrically shaped, measuring approxi
mately 1 by 3 cm and weighing 4 to 6 g. The signal, a series of clicks
increasing in frequency with temperature, was received at a maximum
distance of one meter with an AM transistor radio (50-160 KHz). The
package was sealed in the same manner as the far-field transmitters.
Iguana movement in the burrow and T^ were monitored
simultaneously during the second and third trips by placing both a


13
near-field and a far-field transmitter in the gut of a single large male.
After the lizard submerged for the night, its position was located by
moving the AM radio to where the signal was strongest. The radio was
again placed directly over the lizard early the next morning prior to
emergence. The signal was monitored at the blind six to nine meters away
from the burrow by listening through an earplug connected to the. radio.
When the lizard moved, indicated by fading or loss of the signal, I
quietly moved the radio to the lizard's new position and returned to the
blind. This procedure was repeated until the lizard emerged. To see
how far it would submerge, the lizard was deliberately scared into its
burrow at different times of day. The location of the burrow terminus
was determined by digging at the end of the monitoring period.
In June of 1974 body temperatures of four iguanas were monitored
two at a time in two field cages at Site I. The cages were constructed
of quarter inch hardware cloth in the shape of a cylinder 1 m high
by 2 mlong. Two, 15 cm diameter, 1 m long cylinders were attached to
one end of each cage and covered with sand to simulate burrows. Shellac
coated copper wire was wrapped around the entire cage to act as an
antenna for the near-field transmitters. Strands were approximately
10 cm apart. Each two leads were attached to a circuit breaker connected
to two similar leads from wire wrapped around the ferrite antenna of the
AM radio. Thus, monitoring the T^ of one lizard could be immediately
followed by monitoring the second by switching circuits. Individuals
were kept in a complex habitat during the first four to seven days.
One-half of the cage was covered with thatch palm fronds and branches
were arranged beneath the fronds inside the cage, a 11owing the iguana


14
to get nearly one meter off the ground while still remaining in the
shade. The burrows provided the only shade during the next two to three
days of monitoring, in what was called the simple habitat. The body
temperatures of two caged and two free-ranging iguanas could be monitored
simultaneously from the blind.
Preferred 2V's were determined in two thermal gradients constructed
of plywood, each 2.2 m long, 0.4 m wide, and 0.6 m high, with tops
covered with quarter inch hardware cloth. Two 250 w heat lamps were
suspended at one end of each gradient; their height was adjusted so that
the black body temperature immediately beneath them was 60 C, slightly
below the lowest monthly maximum recorded in the field (63 C in February).
The gradients were placed in a controlled environmental room (Environ-
Room, Lab-Line Instruments, Inc.) set at an air temperature of 25 C (+ 1.0
C accuracy, air velocity next to the lizard less than 0.25 m/second).
The air temperature in the cool end of the gradients gradually increased
to a maximum of 28-30 C after four hours of the eight hour photoperiod.
The photoperiod was centered at 1200 E3T. A 40 w fluorescent bulb
provided less than ten footcandles of illumination. Preferred 's
were determined for eight adult iguanas, a group of four acclimatized
to Gainesville temperatures both in July-August and December-January.
Summer and winter lizards were monitored individually, for four to
eight consecutive days, respectively. Monitoring commenced the day
following transmitter placement. Lizards received neither food nor
water while in the gradient.
Low body temperature behavior was examined in six adult Cyclura
acclimatized to Gainesville temperatures in May. Starting at 1200-1600
(all times reported in this study are EST), the individual was cooled in


15
the environmental room set at an air temperature of 5-8 C. Body tempera
ture was monitored colonically with the Tele-Thermometer. The losses of
seven behavioral responses were noted as T^ decreased: 1. crawling,
2. turning over when placed on the back (critical thermal minimum),
3. maintaining a horizontal head orientation when the body was rotated
about its long axis, 4. maintaining a constant field of view by rotating
the head or eyes when the body was rotated laterally, 5. moving the legs
when pinched while resting on the back, 6. bloating, and 7. closing and
opening the eyes. Each lizard was cooled an additional 3 C, then exposed
to room temperature outside the environmental room. The lizard's body
temperature was again recorded as each of the seven behavioral responses
was regained.
To examine the timing and degree of heat transfer through the dorsal
and ventral surfaces of the iguana's body, three-transmitter experiments
were run in a circular Gainesville pen in late August and early
September. Two far-field transmitters were fastened with electrician's
tape to the outside of a piece of television antenna cable. When the
cable was wrapped around the lizard's waist just anterior to the rear
legs, the transmitters were situated ventro-laterally on either side of
the trunk. The cable ends were held together dorsally with a rivet.
The temperature-sensitive thermistor bead, embedded in epoxy resin, was
located at the end of a short probe wire extending from one end of the
transmitter. One probe tip was taped to the middle of the ventral side
of the cable, to monitor the body-substrate interface temperature
immediately under the lizard. The other, painted flat black to monitor
dorsal heat input, was suspended free in the air about 1 cm anterior
to the cable and 5 mm above the middle of the lizard's lower back.


16
A near-field transmitter was placed in the lizard's gut. Three large
males (1195, 1240, and 1260 g), monitored one at a time for a total of
nine days, moved freely about the pen. Seven other iguanas were also
present. Readings from the three transmitters were taken every five to
ten minutes. To obtain gut temperature readings, the AM radio was taped
to the end of a wooden pole and extended from outside the pen to within
a meter of the lizard. Shaded air, sun-exposed sand, and black body
temperatures were also measured. The lizard's behavior was observed
and notes taken for later correlation with the transmitter records.
To determine whether feeding rate or meal size was significantly
less immediately after morning emergence than during the higher T-^'s of
the regulating phase, iguanas in the Gainesville pen were presented with
known weights of white grapes. Food was either placed on trays on the
ground, before individuals emerged for the day, or on trays affixed to
the tops of posts, after individuals had reached regulating phase body
temperatures. The elevated trays were large enough for only one
individual to sit on, preventing feeding by several animals on a single
known weight of grapes. Also, by placing grapes on three post tops
simultaneously, one feeding lizard was rarely displaced by another.
Animals were fed on clear days after at least two days with no food.
Sun orientation directions of lizards were measured with a hand
held Silva compass. Records were obtained in the field, the Gainesville
pen, and a 2 m circular arena placed on top of the zoology building. To
determine the importance of vision in sun orientation control, lizards
were blindfolded by placing ellip.tically shaped plastic disks over their
eyes and securing the disks with several layers of electrician's tape


17
wrapped around the head. A stretch sock, with an opening for the
nostrils, was then placed over the head and taped at the neck to prevent
the lizard from scratching off the tape.
A photothigmotron was built to quantify thermoregulatory postural
changes in Cyolura. A square plywood substrate box, measuring 65 by 65
by 5 cm, was lined with Plexiglas and sealed internally with fiberglass
cloth and resin. A 0.3 mm thick aluminum sheet formed the top of the
inner Plexiglas box and was affixed to it with Al-40 adhesive (Devcon).
A grid of 1 cm squares was scored on the exposed surface of the
aluminum. The substrate box rested on a 15 gallon water bath
(Precision Scientific Co.). The water bath pump was connected via
Tygon tubing (Norton Plastics and Synthetics Division) to two ports
drilled in the side of the substrate box. Water at the desired tempera
ture was pumped through the entry port in the substrate box and circulated
evenly via six inner exit ports pointed in different directions. A small
hole was made at one corner of the aluminum top to permit pressure
equalization when the pump was initially turned on. A short tube was
attached externally to the rim of the hole to prevent water overflow.
To confine the lizard to the aluminum substrate, a one meter high hard
ware cloth cage, reinforced with wooden frames at top and bottom, was
placed on the substrate box. The cage top was covered with a Plexiglas
sheet, on which was placed a 35 mm camera directed downward toward the
aluminum grid. A single 250 w heat lamp, attached to a tripod resting
on the floor, was directed slightly downward toward the side of the cage.
The observer's side of the cage was covered with vinyl plastic provided
with a small observation port. The photothigmotron was placed inside
the Environ-room.


18
Before each iguana was introduced into the photothigmotron,
several body areas were determined. The lizard was anesthetized with
halothane and placed on a Plexiglas sheet. Outlines of the outer edge
of the body and the inner area of substrate contact were marked on the
sheet and later transferred to a piece of paper. Three separate
measurements of each area were made planimetrically and the mean taken.
Contact area of the anesthetized lizard was assumed to equal maximum
contact of the free-ranging lizard. The day before the first experiment,
the lizard was placed on an insulative piece of plywood resting on top
of the aluminum substrate of the photothigmotron. One experiment was
performed each day, starting at midday. The photoperiod was ten hours.
Animals were kept in the cage for three to five experiments, during
which they received neither food nor water. An hour before heating or
cooling of the lizard began, the water bath pump was turned on to bring
substrate temperature to the desired level. The plywood was then
gently removed, shifting the individual onto the metal substrate. The
heat lamp was also turned on in some experiments. Photographs of the
dorsal aspect of the lizard were taken at intervals. Core body tempera
ture was measured with a near-field transmitter placed in the digestive
tract. The extent of leg and tail contact was noted, as well as how
much of the anterior trunk was elevated off the substrate. The black
and white photograph negatives were projected with a slide projector
onto a piece of white paper and the body outline traced. The dorsal
outline area was determined planimetrically. Head, anterior trunk, leg
and tail noncontact areas as well as a standard correction based on the
difference between the inner and outer outlines of the anesthetized


19
lizard were subtracted from the dorsal outline area to get the approxi
mate contact area. This value could then be converted to a percentage
of the maximum possible contact.
Head temperature was measured at two different locations, using
probed near-field transmitters. The transmitter was taped to a piece
of television antenna cable fastened with a rivet around the lizard's
neck. In a large male monitored in the photothigmotron, the thermistor
probe was inserted into the orbital venous blood sinus through a 1 mm
diameter hole drilled in the skull. The probe wire was glued to the
center of a circular cork disk, which was glued to the top of the head
with epoxy resin. Initial placement of the probe resulted in a temporary
eye bulging (also noted during cannula placement in Phvynosoma aormutvm
by Heath 1966). In a second large male, monitored in the arena atop the
zoology building, the probe tip was placed just inside a slit made in
the tympanic membrane. The probe wire was glued to a plastic disk, which
was glued to the outside of the tympanum. Lizards were allowed to
recover from implantation for three days before testing.
Skin reflectivity was determined for two Cyclura adults from Water
Cay, one in the light and one in the dark color phase. The light phase
lizard was killed by heating it above the lethal maximum body tempera
ture and the dark phase lizard died of starvation. Epidermal-dermal
squares, measuring approximately 25 mm on a side, were removed from the
lizards immediately after death and placed on water-moistened filter paper.
Patches were taken from the mid-dorsal, ventral pectoral, and ventral
abdominal trunk and the dorsal and ventral tail just distal to the anus.
Integumental reflectivities were immediately determined, using a Bausch


20
and Lcmb Spectronic 20 Spectrophotometer with reflectometer attachment.
Monochromatic reflectance was obtained at a 90 angle to the skin surface,
using a magnesium carbonate block as a standard. Readings were taken
between 340 and 960 nm at 20 nm intervals. Reflectivity over the same
range was also determined for a common substrate on Pine and Water Cays,
fused corralline sand covered with two species of blue-green algae.
To check for possible reflectance of near-infrared light (700 to
900 nm) from the integument of live Cyotura, two adults were photographed
during daily activity in Gainesville, using Kodak Infrared Ektachrome
film (Schwalm et at. 1977). When the camera lens is covered with a
yellow filter, reflected near infrared light appears red in the developed
film.
Skin patches were heated from different angles to determine whether
the rate of integumental heating decreased as the radiant heat shadow
caused by scale imbrication increased. The patches were removed from
various locations on a single adult iguana, tacked to separate pieces
of plywood, dried, and cooled to 20 C in a controlled environmental
room. A YSI Probe 421 was placed under the patch to monitor the time
required to heat the dried skin to 50 C with a 250 w heat lamp. Each
patch was heated 11 times, with the lamp at a constant distance of 17 cm,
oriented normally to the skin and at 15, 30, 45, 60, and 75 from normal,
both directly anterior and posterior to the imbricate edges of the
scale rows.
The histological structure of integumental "sensory spots"
(Miller and Kasahara 1967) were examined in tissue sections prepared for
light microscopy. Material had been fixed in formalin and transferred


21
to isopropyl alcohol for preservation at the Florida State Museum.
Tissue blocks were taken from the lizard's rostral, labial, parietal
eye, and dorsal tail regions, dehydrated in ethyl alcohol, embedded in
methacrylate plastic, and sectioned at 5 micrometers thickness vertical
to the skin surface. Sections were mounted serially and stained with
Mayer's hematoxylin and eosin-phloxine.
Six adult iguanas were heated and cooled in a controlled environ
mental room (Environette, Lab-Line Instruments, Inc.). Each individual
was habituated for one day to its small hardware cloth cage, placed in
one Environette set at an air temperature of 20 C. Polyurethane panels
enclosed three sides of the cage, reducing the air flow over the lizard's
body surface to practically zero. The cage floor was also composed of
polyurethane. On the second day a YSI rectal probe, coated with petroleum
jelly, was inserted into the lizard's large intestine to a depth of 7-10
cm and taped to the tail. The cage was then rapidly moved to another
similarly wind-baffled box in another Environette set at an air tempera
ture of 40 C. When within 1-2 C of ambient air temperature, the lizard
was rapidly transferred back to the 20 C Environette and cooled. During
the third and final day in the chamber, the individual was secured in a
supine posture to a wooden jig. The lizard's legs were free to move.
A rectal probe was again inserted into the intestine and a banjo probe
glued middorsally to measure body surface temperature. The individual
was then heated and cooled again. Animals were observed continuously
during heating and cooling through an observation port in the chamber
door. Breathing rates were recorded intermittently. The photoperiod
was ten hours. Heating began at 1300 and cooling was always over by


22
2100. Lizards were weighed at the start and the end of the test period
and given neither food nor water.
Heart rate and oxygen consumption at rest, during maximum activity,
and during recovery from exhaustive exercise were determined for five
adult iguanas (two females and three males; X = 1252 g, range = 834-1722).
Each lizard was tested on six consecutive days, starting at an air
temperature of 15 C and increasing at 5 C increments to a maximum of
40 C. Experiments were conducted after 2100 on individuals which had
received neither food nor water for two days prior to the first run.
After each run the chamber air temperature was elevated five degrees
for the next night's run. When not being tested, animals were permitted
to move freely about the 1.4 by 2.5 m Environ-Room, kept on a 10L:14D
cycle.
The lizard's exhaled air was trapped in a rectangular, black-
painted Plexiglas mask fitted over the head. The mask had a neoprene
apron glued to its open posterior end, which fitted over the lizard's
anterior trunk and forelegs. Room air was sucked into the open posterior
end of the apron, past the lizard's head, and out the front of the mask
via Tvgon tubing through columns of soda lime and Drierite, to remove
carbon dioxide and water vapor, respectively. Flow rates ranged from
280 to 2440 cc/min. The gas then passed through a flowmeter, a pump,
another flowmeter, and finally through a Beckman C-2 paramagnetic oxygen
analyzer, located outside the Environ-Room. Gas flows were measured
with rotameters calibrated with a Brooks Vol-U-Meter. The decrease in
percent oxygen resulting from the lizard's respiration was recorded
continuously, except during the exercise period, with a Honeywell
Electronic 15 stripchart recorder. Lizards were kept in total darkness


23
during all oxygen consumption measurements. Oxygen consumption was
calculated from oxygen content and flow rate data using the appropriate
equation given by Depocas and Hart (1957).
For each run, a rectal YSI probe was inserted into the lizard's
intestine. The animal was then fastened in a supine posture to a wooden
jig, with its head resting inside the mask, taped to the jig. Oxygen
consumption was monitored until a stable resting rate was achieved.
The lizard was then removed from the jig and mask and exercised by hand
manipulation for several minutes, sufficient to completely exhaust the
animal. The lizard was immediately refitted into the mask and refastened
to the jig. ECG needle leads were pushed beneath the skin of the shoulder
and the base of the tail. Heart rate was measured every 4-10 minutes of
the recovery period with a Physiograph Four A with high gain preamplifier
(E and M Instrument Co.), located outside the Environ-Room. When
oxygen consumption again fell to the resting rate, the lizard was
manipulated by hand for another two minutes to get a maximum heart rate
reading.


ECOLOGICAL ASPECTS OF THE THERMAL BIOLOGY
OF CYCLURA CARINATA
Climate of Water Cay
The Caicos Islands are in the northern tropical zone, slightly
below the Tropic of Cancer. They are also in the tropics under the
Koppen System (Koppen 1923), since mean monthly shaded air temperature
never falls below 18 C. Water Cay has a trade wind littoral climate,
with fairly uniform high temperature and distinct wet and dry seasons.
Figure 1 summarizes the black body temperature measurements taken
during three trips to Water Cay. Cyclura had to remain intermittently
in the shade on sunny days during a large portion of its activity period
or face lethal body temperatures due to radiant heat input. Adult
lizards provided with some well timed overcast periods could theoret
ically survive in an exposed location all day in at least October to
February. Yearly photoperiod varied by almost three hours, reflected
in the difference in width of the heating curves for February vs. July.
Figure 2 illustrates the annual air temperature cycle near the
two primary locations where Cyclura was observed. The range of mean
shaded air temperature was only 4.5 C on Grand Turk. Of course, both
daily and seasonal microclimatic extremes at ground level were greater.
Gainesville, located in the subtropics, was too cold part of the year
to support a Cyclura population. From the beginning of April until
the end of September, penned Gainesville lizards ate readily and gained
weight. Feeding was negligible during the other six months of the year.
24


25
on i -
7 8 9 10 II 12 13 14 15 16 17 18
HOUR INTERVAL
Figure 1. Black body temperatures in the field. February, June, and
October measurements are represented by solid, dashed, and dotted
lines, respectively. Maximum temperatures during the recording
periods are represented by open squares, mean temperatures for the
hourly intervals by open circles, and minimum temperatures by
solid circles. All weather conditions encountered were included
in the calculations. Horizontal lines are means and vertical lines
are ranges of regulating phase Tfo's of lizards at Site II for
February, July, and October, respectively.


AIR TEMPERATURE (C)
26
Figure 2. Air temperature in the shade at Gainesville, Florida (Agronomy
Farm Weather Station; 29 40' N, 82 20' W; Prine 1977; and
Grand Turk, British West Indies (Auxiliary Air Field; 21 26' N,
71 8' W). Vertical lines extend from the maximum to the
minimum recorded Ta each month (dashed line for the year 1976
at Gainesville, solid line for the years 1900-1968 at Grand Turk).
Horizontal lines are mean monthly temperatures on Grand Turk.
Circles and squares represent mean maximum and mean minimum air
temperatures, respectively. Air temperatures were measured in
a standard NOAA shelter (Stevenson screen) at a height of 1.5 m.


27
Special precautions had to be taken to prevent the lizards from freez
ing to death between November and February.
An easterly wind blew almost continually over the islands. The
mean surface wind speed on Grand Turk was 24.6 km per hour (15.3 mph).
Figure 3 illustrates the mean maximum July wind speeds on a road pro
tected on the windward side by a limestone ridge and dense scrub at
Site II. Wind speeds were greatest during the middle of the day, a
typical pattern (Geiger 1957). Wind speed was greater at 50 cm than
at 3 cm above the ground, also a typical pattern. Thus, lizards basking
or feeding above the ground were usually cooled at a greater rate by
forced convection than lizards on the ground. Wind speed was consider
ably higher facing the inlet between Water and Pine Cays just a few
meters away. Since lizard burrows were common near the top of the
ridge, some interesting wind related behavior was expected there,
perhaps similar to the seeking out of minimal wind exposure at low
Tfo and maximum exposure at high Tj, as seen in Amblyrhynehus ovistatus
of the Galapagos Islands (Bartholomew 1966, Bartholomew et al. 1976).
Rainfall was distinctly seasonal in the Caicos (Figure 4). Mean
annual rainfall on Pine and Water Cays was probably slightly greater
than on Grand Turk (597 mm or 23.5 inches). Four seasons characterize
the western islands of the Caicos; a cool dry winter from January to
March, a warm dry spring from March to May, a hot, increasingly wet
summer from June to October, and a warm, wet autumn from October to
December.
Water Cay was fairly humid (Table 1). Daily relative humidity
decreased as air temperature increased, minimizing near midday. Of


MEAN MAXIMUM WIND SPEED (M/SEC)
28
Figure 3. Mean maximum wind speed at Site II in July. Ten consecutive
wind speed maxima were measured each hour for seven days at
3 cm (open rectangles) and 50 cm (solid rectangles) above the
ground. Horizontal lines are means, vertical lines are ranges,
and rectangles are plus and minus two SE of the mean.


MONTHLY RAINFALL (MM)
29
190
Figure 4. Monthly precipitation on Grand Turk, British West Indies. The
line represents the mean monthly rainfall for the years 1900-1968.
Total rainfall per month for several months on Pine Cay, Caicos
Islands, are represented by dots.


30
Table 1. Daily variation in relative humidity. Measurements
taken during hours of precipitation were not included.
Data were collected at Site I (Jun) and Site II (Feb,
Jul, Oct) on Water Cay.
MEAN HOURLY
RELATIVE
HUMIDITY
HOUR INTERVAL
FEB
JUN
JUT.
OCT
N=4
N=12
N=15
N=7
DAYS
0700-0759
-
75
74
86
0800-0859
78
74
71
77
0900-0959
67
69
64
69
1000-1059
66
62
64
64
1100-1159
66
59
61
62
1200-1259
60
59
64
62
1300-1359
61
64
63
59
1400-1459
62
61
64
64
1500-1559
66
61
65
66
1600-1659
70
67
71
70
1700-1759
73
72
71
78
MINIMUM DAILY READING
56
46
52
62
MAXIMUM DAILY READING
79
80
80
87
RANGE
23
34
28
25
MEAN MONTHLY RH
66.
0
64.6
66.9
68.
0
2SE
2.
1
1.4
1.5
2.
4


31
course, daily absolute humidity did not vary as much as relative
humidity. Mean monthly relative humidity did not differ significantly
between February, June, July, and October.
Piel Body Temperature Cycle of a Heliothermic Lizard
The body temperature cycle was divided into four phases by Wilson
and Lee (1974). Some slight alterations were made for the following
scheme (Figure 5).
1. Morning Heating Phase The lizard emerged from its burrow and
heated rapidly in direct sunlight. Cyelura usually started its first
feeding foray below the first maximum Tfo, upon reaching the "normal
activity range" (Cowles and Bogert 1944). Wilson and Lee's "basking
phase" ends here. This term is confusing since basking did not cease
when the first feeding foray began. Also, Cyclura was so wary during
the June-July trip that it was very difficult to observe the start of
the first foray unless I was sitting in a well positioned blind. There
fore, the first phase has been called the morning heating phase, ending
at the first maximum TThis endpoint is obvious on a continuous T^
record.
2. Regulating Phase 2^ fluctuated between a series of maxima and
minima, staying within a fairly narrow range.
3. Burrow Cooling Phase It started at the lizards evening
submergence and ended during the night, when T^ had fallen to the 24
hour minimum. The decline in T, followed a Newtonian cooling curve,
with an ever decreasing rate of T^ decline as the stable deep burrow
temperature was approached.


BODY TEMPERATURE
TIME
Figure 5. Body temperature record of Lizard 2 on February 1, 1976. The Arabic numerals at the
top of the graph refer to the four phases of the cycle. The broken line at the
bottom indicates the periods when the sun was obscured by clouds. Capital letters
refer to data points discussed in the text.
Co
N>


33
4. Stable Tfo Phase The lowest T^ in the 24 hour cycle occurred
just before emergence in two lizards larger than Cyoliara carlnata, the
Lace Monitor, Vararais varius (Stebbins and Barwick 1968) and the Komodo
Dragon, Varanus komodoensis (McNab and Auffenberg 1976). However, T^
stabilized considerably before emergence in Cyclura, necessitating a
fourth descriptive phase. The term "inactive phase" used by Wilson and
Lee would be confusing since Cyclura was also inactive in the burrow
cooling phase.
A continuous body temperature record provides a substantial amount
of information about a lizard. It was sometimes difficult to determine
the exact time of emergence from the record, since deep burrow tempera
ture was slightly greater than surface temperature early in the morning.
The lizard in Figure 5 may have lost heat on the surface just after
emergence (A) or near the burrow mouth prior to emergence (B). Slight
differences in emergence T^ could sometimes be used to determine the
burrow the lizard spent the night in, without having to locate the
burrow the preceding night by following the increasing signal strength
with the receiver. The time of submergence (F) is easily discerned in
the February record. However, determination of submergence time from
the summer records was commonly more difficult, since often fell
prior to submergence and at a rate not easily distinguished from the
initial decline after submergence. Body temperature shifted abruptly
upward as the overcast cleared temporarily at (D), a change maintained
until submergence. Shifts of this sort were fairly common in the records.
Several further terms concerning lizard need to be defined.
All refer to measurements on a single individual taken during its


34
regulating phase. Unless otherwise specified, each T^ reading used in
a calculation occurred during a period of sunshine or overcast (10
minutes or less) which did not cause a decline in 2^.
1. Mean Daily Ty Once the continuous record for a single day
was plotted, the first and last maxima plus points at fifteen minute
intervals, on the hour, were averaged.
2. Daily Minimum, Maximum Ty The lowest, highest T^ recorded
for the day (C and E, respectively, in Figure 5).
3. Mean Daily Minimum, Maximum Ty An average of the daily
minimum, maximum Tfor a specified number of days.
4. Mean of Daily Ty Minima, Maxima An average of all minima,
maxima for one day.
5. Mean of Ty Minima, Maxima An average of all minima, maxima
for a specified number of days.
Regulating Phase Body Temperature in the Field
Cycluva oarinata is a stenothermal lizard during the regulating
phase (Table 2). The mean of the regulating phase T\ ranges for all
the monitoring days during clear weather was only 3.3 C (SD = 1.3,
range = 0.7-6.4, N = 106 days). The frequency distribution of these
Tjj ranges was positively skewed toward greater stenothermality, with
a median of 2.9 C and a modal class of 2.0-2.4 C.
Adults rarely went into their burrows at Sites I and II because
of extremely high surface temperatures, although this may have been a
common behavior in more sparsely vegetated areas. Free-ranging lizards
prematurely submerged for the day on only 19 occasions during 129
individual monitoring days: 45 percent due to rain, 40 percent due to


Table 2.
Summary of regulating phase body temperatures of lizards in the field. Data are
arranged by season, field site, sex, Increasing body weight, and individually by
decreasing daily mean Tfo, each line equaling one day's monitoring. Mean Tfo's were
calculated from the entire record, irrespective of weather conditions or lizard
location. The letters (B) and (SU) signify that the lizard was in a burrow or the
sun was obscured by clouds (sun under) when the minimum Twas recorded, respectively.
Lizards 2 and 3 were monitored in February and October.
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
DAILY
MAXIMUM
Tb
RANGE
LENGTH OF
REGULATING
PHASE (MIN)
% TIME
SUN
BEHIND
CLOUDS
% TIME
LIZARD IN
BURROW
1(F)
F
1135
II
2-19-76
37.7
0.82
36.2
39.6
3.4
293
4
0
17
37.5
1.85
33.4(B)
39.8
6.4
280
15
36
18
36.8
1.38
34.0(B)
38.8
4.8
240
30
19
2(F)
M
1665
II
2-05-76
40.8
0.46
40.2
41.5
1.3
142
10
0
6
38.6
0.72
36.0
39.2
3.2
288
14
0
7
38.4
0.68
37.1
40.0
2.9
264
8
0
12
38.3
0.52
37.0
39.2
2.2
315
0
0
11
38.3
0.27
37.5
38.6
1.1
187
28
0
9
38.0
1.40
34.7(SU)
40.0
5.3
332
8
0
8
37.7
0.94
35.4
39.0
3.6
333
0
0
10
37.6
0.79
35.9
38.8
2.9
244
41
0
3(F)
M
1775
II
2-12-76
40.0
0.60
38.1
41.0
2.9
427
0
0
13
38.5
0.53
37.1
39.4
2.3
363
40
0
11
38.2
0.50
37.0
39.2
2.2
288
20
0
14
38.2
0.63
36.6
39.5
2.9
426
8
0
15
38.1
0.61
37.0
39.2
2.2
327
19
0
19
37.8
0.48
36.4
38.6
2.2
330
3
0
17
37.4
0.79
35.6
38.5
2.9
505
38
0
18
37.4
0.69
35.8
38.7
2.9
360
43
0
20
36.0
1.81
32.6(SU)
38.6
6.0
313
63
0
CJ
l_n


Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
4(C)
F
645
I
6-10-74
40.8
2.03
37.6
9
40.4
1.14
38.2
8
40.3
1.63
37.4
11
39.9
3.00
35.8
7
39.1
0.93
37.1
5(C)
F
723
I
6-24-74
41.9
0.81
40.4
25
41.5
0.42
40.7
22
40.6
0.95
38.5
26
40.6
0.58
39.3
27
40.0
0.76
38.2
28
39.3
0.96
37.2
29
39.3
0.68
38.0
7-01
35.3
3.79
28.6(B)
6(C)
F
815
I
6-17-74*
39.0
0.48
37.8
9
38.8
1.29
37.2
8
38.3
0.40
37.4
18
38.3
4.18
33.9(SU)
10
38.3
1.10
36.4
7
37.6
1.29
35.7
14
37.0
1.37
34.8
11
36.5
1.36
34.9
7(C)
M
1290
I
6-24-74
40.7
0.63
39.4
22
40.3
0.63
38.0
29
38.7
0.96
36.3
25
38.1
1.24
35.3
26
38.1
1.01
35.7
DAILY LENGTH OF
MAXIMUM REGULATING
Tb RANGE PHASE (MIN)
% TIME
SUN % TIME
BEHIND LIZARD IN
CLOUDS BURROW
43.8
6.2
419
30
41.8
3.6
260
23
42.6
5.2
288
7
42.0
6.2
312
24
41.0
3.9
360
0
43.7
3.3
325
0
42.5
1.8
501
0
41.5
3.0
272
29
41.7
2.4
481
3
41.4
3.2
507
0
41.4
4.2
525
10
40.2
2.2
459
0
39.8
11.2
481
72
40.0
2.2
315
50
40.8
3.6
291
29
39.4
2.0
268
11
41.3
7.4
242
42
39.8
3.4
430
29
39.8
4.1
407
2
38.6
3.8
326
31
37.8
2.9
315
24
41.9
2.5
325
0
41.8
3.8
332
24
40.8
4.5
465
0
41.3
6.0
465
0
39.7
4.0
488
3
NOT
APPLICABLE
to


Table 2. Continued.
LIZARD
(FREE OR
CAGED) SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
7(Continued)
27
38.0
1.16
36.0
28
37.8
1.11
35.3
7-01
35.0
3.22
29.3(R)
8(F) F
675
I
6-26-74
41.9
0.44
40.7
25
40.9
0.87
39.3
14
40.6
0.97
37.7(SU)
16
39.8
0.66
38.5
24
39.8
0.46
38.4
22
39.6
0.60
37.3
17
39.6
1.38
36.7(SU)
23
39.2
0.71
37.5
18
38.4
1.93
30.3(SU)
12
35.2
3.04
30.4(B)
15
33.8
4.45
29.7(B)
9(F) M
1450
I
6-29-74
42.4
0.37
41.6
28
42.4
0.46
41.6
17
42.3
0.39
41.5
27
42.2
0.66
40.9
24
41.7
0.42
40.8
16
41.6
0.70
40.5
23
41.0
0.48
39.9
25
40.1
0.57
39.4
7-02
40.0
1.05
38.1
6-26
39.9
1.08
38.0
7-01
39.2
0.51
38.1
4
37.2
3.82
29.8(B)
6-15
35.0
4.31
28.9(SU)
% TIME
DAILY
MAXIMUM
Tb
RANGE
LENGTH OF
REGULATING
PHASE (MIN)
SUN
BEHIND
CLOUDS
% TIME
LIZARD IN
BURROW
40.1
4.1
507
0
NOT
39.9
4.6
417
12
APPLICABLE
39.7
10.4
507
69
II
42.9
2.2
420
0
0
42.1
2.8
493
0
0
42.1
4.4
321
21
0
42.3
3.8
457
0
0
40.6
2.2
360
7
0
40.3
3.0
378
22
0
42.1
5.4
393
51
0
40.2
2.7
339
10
0
40.5
10.2
300
34
0
40.9
10.5
429
0
43
42.1
12.4
330
45
170
43.3
1.7
490
0
0
43.5
1.9
525
0
0
43.0
1.5
390
54
0
43.2
2.3
486
0
0
42.6
1.8
378
7
0
43.3
2.8
457
0
0
41.6
1.7
351
10
0
41.2
1.8
450
0
0
42.5
4.4
510
14
0
41.4
3.4
480
3
0
39.9
1.8
267
47
0
40.9
11.1
455
26
17
41.0
12.1
358
35
0
co
--vj


Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(C)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
10(F)
M
787
II
7-18-74
40.8
0.85
38.8
13
39.9
1.79
36.9
15
39.5
0.58
38.2
12
38.6
1.07
37.1
16
38.5
1.07
35.2(B)
20
38.4
1.43
32.1(B)
22
38.4
1.39
35.9
21
38.1
1.21
35.3
19
37.4
2.39
31.9(B)
23
37.1
1.30
34.9(SU)
11
36.9
1.60
33.3(B)
17
34.4
3.30
30.1(SU)
11(F)
M
1598
II
7-09-74
41.4
0.87
39.0(SU)
10
41.3
0.33
40.3
11
40.8
0.36
40.1
12
40.1
0.59
38.8
18
38.9
0.66
37.4
13
38.8
0.78
37.1
6
38.5
0.25
38.2
22
38.2
1.11
36.6
15
38.1
0.66
33.7(SU)
19
38.0
0.86
36.5
20
38.0
0.64
36.9
21
37.6
0.88
35.7
23
37.1
1.49
33.8(SU)
25
36.7
1.27
35.1
7
36.2
1.11
34.1(SU)
17
34.2
3.31
29.5(SU)
16
28.0
0.10
28.0(B)
DAILY LENGTH OF
MAXIMUM REGULATING
Tfc RANGE PHASE (MIN)
% TIME
SUN / TIME
BEHIND LIZARD IN
CLOUDS BURROW
42.5
3.7
408
10
0
42.9
6.0
500
0
0
40.6
2.4
375
6
0
40.8
3.7
486
6
0
40.6
5.4
384
2
9
40.2
8.1
450
19
8
40.7
4.8
487
12
0
40.2
4.9
435
5
0
40.6
8.7
431
0
20
39.5
4.6
318
16
0
39.9
6.6
460
0
31
38.9
8.8
405
62
0
42.9
3.9
396
35
0
42.1
1.8
420
30
0
41.5
1.4
481
0
0
41.1
2.3
509
6
0
40.1
2.7
480
8
0
40.4
3.3
339
0
0
38.9
0.7
178
20
0
40.3
3.7
574
14
0
39.6
5.9
550
27
0
39.6
3.1
489
42
0
39.3
2.4
470
24
0
39.8
4.1
517
22
0
39.6
5.8
575
26
0
39.7
4.6
285
5
0
38.4
4.3
234
100
0
39.8
10.3
381
65
0
28.3
0.3
0
2
100
U)
09


Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
12(F)
M
1420
I
8-28-76
39.7
0.28
39.4
30
39.5
0.57
37.9
29
36.3
2.31
32.2(B)
13(F)
M
950
III
9-03-76
39.3
0.69
38.2
5
38.7
0.59
37.5
4
37.8
1.14
35.2
O
L
36.8
2.63
31.8(SU)
14(F)
M
1864
II
10-25-75
39.9
0.84
38.2
21
39.7
0.36
38.5
23
39.3
0.53
38.2
15(F)
M
1784
II
10-29-75
39.2
0.85
37.5
28
39.2
0.82
37.3
27
38.3
0.81
37.6
3(F)
M
1672
II
10-06-75
40.1
0.65
37.9
5
38.6
3.30
31.0(B)
7
38.2
1.44
35.0
8
38.1
1.45
34.6(B)
16
37.8
1.09
35.9
21
37.7
0.95
36.2
4
37.6
0.89
35.1
20
37.5
1.29
35.2
18
37.5
0.82
35.6
19
37.3
0.76
34.5
15
34.8
2.82
30.6(B)
DAILY LENGTH OF
MAXIMUM REGULATING
Tjy RANGE PHASE (MIN)
% TIME
SUN % TIME
BEHIND LIZARD IN
CLOUDS BURROW
40.2
0.8
114
0
0
40.2
2.3
327
27
0
39.9
7.7
387
38
25
40.4
2.2
195
35
0
39.6
2.1
315
3
0
39.4
4.2
213
62
0
39.3
7.5
417
45
0
40.8
2.6
186
24
0
40.7
2.2
300
18
0
40.2
2.0
357
6
0
41.0
3.5
281
0
0
40.7
3.4
313
31
0
40.4
2.9
330
30
0
41.5
3.6
445
5
0
42.3
11.3
487
10
20
40.6
5.6
345
30
0
40.4
5.8
380
0
12
39.4
3.5
159
19
0
39.7
3.5
255
13
0
39.4
4.2
312
73
0
39.7
4.5
270
22
0
39.0
3.4
310
59
0
38.3
3.8
354
5
0
39.2
8.6
285
21
49
u>


Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEICHT
(G)
SITE
DATE
KEAN
?b
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
DAILY
MAXIMUM
Tb
RANGE
LENGTH OF
REGULATING
PHASE (MIN)
% TIME
SUN
BEHIND
CLOUDS
% TIME
LIZARD IN
BURROW
2(F)
M
1507
II
10-16-75
40.1
0.83
39.3
41.4
2.1
90
33
0
19
39.5
0.37
38.5
40.1
1.6
308
6
0
18
38.6
0.76
37.2
40.2
3.0
360
51
0
Laid eggs in the cage during this day.
p-
o


41
human disturbances, and 15 percent due to extreme cloudiness. Rates of
cooling during burrow excavation and shade sitting were not consistently
different. Thus, the cause of the cooling could not be determined by
examination of the T^ record alone.
Figure 6 summarizes the regulating phase data for all the field
lizards. The rise in mean T^ over the 3.5 C range was steady. Minimum
T^'s fell into two groups having no correlation with increasing mean
2%/s, one group above and one below 39 C mean T* This probably is of
no significance. The correlation of maximum with mean T- was better.
The lowest maximum T^ recorded was 1.6 C above the highest minimum. Thus,
all ranges of individuals overlapped somewhat, no matter what the
sex, habitat type, or season of measurement.
The sample sizes were too small to determine conclusively whether
or not mean T7 differed significantly with sex. Lizard 9, a dominant
male, had a significantly higher T^ than Lizard 8 (p <_ 0.05), an adult
female monitored simultaneously. However, two of three caged females
in the simple habitat, Lizards 4 and 5, had significantly higher body
temperatures than the caged male, Lizard 7. Mean 's were not
significantly different in two free-ranging males and a female monitored
at Site II in February.
Gravid females may have lower regulating phase 2^'s than spent
females and males. The gravid June female (Lizard 6) had the lowest
mean of all those monitored. The means were significantly higher
(p _< 0.05) for the other two caged females (Lizards 4 and 5) and the
free-ranging female (Lizard 8) at Site I, all of which had probably
recently laid eggs, since they actively defended their burrows against
both encroaching males and females. However, Lizard 6's T^
was not


Figure 6. Summary of field body temperatures during the regulating phase. The lizards were
monitored from 3 to 16 days. Mean regulating phase T was calculated from values
taken at 15 minute intervals from the monitoring record. The number of days used
to calculate each mean was variable and usually equal to the total monitoring time
(Table 2). Periods of hyperthermia were included in calculation of the means,
whereas periods of declining Tfo due to overcast or burrow submergence were not.
Vertical lines are ranges, horizontal lines are means, and rectangles are 99 per
cent confidence limits of the mean. Sex is indicated by an M or F. The four
caged lizards in the complex habitat at Site I in June are represented by
rectangles filled with vertical lines; the same four lizards in the simple habitat
by dots; three free-ranging lizards in February at Site II by horizontal lines;
two lizards in June at Site I by diagonal lines sloping downward to the right; two
lizards in July at Site II by open circles; a single lizard in August at Site I by
open squares; a single lizard in September on Fort George Island by an open rectangle;
four lizards in October at Site II by diagonal lines sloping downward to the left.


44
43
42
4
40
39
38
37
36
35
34
33
wj/^0 mu,?.
M
JL
7 2 13 10 7 II 3 I! ¡2 5 14
INDIVIDUAL IDENTIFICATION NUMBER
2 4 8 4
4^
LO


44
significantly lower than the caged male, Lizard 7. On the single day
Lizard 6's Tu was recorded after egg laying, the maximum was 0.5 C
higher than the highest maximum of the seven monitoring days previous
to laying. The highest mean T^ recorded was on the day she laid her
eggs, but the mean was not significantly higher than the next two
highest means recorded prior to laying. The differences probably
would have been significant had not the day of laying been overcast
50 percent of the time.
Temporary Hyperthermia and Hypothermia
All eleven iguanas monitored in the field passed through a period
of hyperthermia soon after release (Figure 7 and Table 3). T* then
declined to a more stable, presumably normal range. The T^ decline
was either rather rapid as for Lizard 3 or more gradual as for Lizards
8 and 11. The extent of the hyperthermic response, as measured by tne
sum of the ranges of the maximum daily T^ and the mean daily 2^'s
between the hyperthermic peak day and the return to normalcy, was
usually slightly greater in coelomically than enterically implanted
lizards. However, the response may vary with season as well as with
transmitter location. The summer animals were probably suffering
from a bacterial infection resulting from transmitter implantations
(see Vaughn et at. 1974, Kluger et at. 1975, and Bemheim and Kluger
1976a, b for related laboratory studies). The second Th rise observed
in Lizards 8 and 9 may have been due to reinfection of the wound or
a secondary response to a chronic infection. In the latter case it
may be that lizards could not tolerate a high T^ for more than a few
days even though infectious agents were still present (Cowles and


Figure 7. Lizard hyperthermia and hypothermia during the regulating phase. Lines
connect T£, maxima, omitting those during periods of heavy overcast (open
circles). Open squares are daily Tfo means. Lizard 11 was prevented
from emerging on Day 3 by a land crab (Cavdisoma guanahumi) ensconced
for a single day just inside the burrow mouth. The transmitter was in
the gut of Lizard 3 and in the coeloms of Lizards 8 and 11.




Table. 3. Hyperthermia in Cyalura oavinata in the field. The ease of graphically distin
guishing a Tfo peak and a recovery from hyperthermia was classified as high, medium,
or low. The maximum 7', was the highest body temperature recorded for a lizard.
The low maximum Tfo was the lowest maximum recorded on the day recovery was com
plete or before a second rise in Tfo occurred (Lizards 8 and 9).
LIZARD
2
3
4
5
6
7
8
9
10
11
3
MEAN S.D.
MONTH
FEB
FEB
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JULY
JULY
OCT
LOCATION OF
TRANSMITTER
GUT
GUT
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
GUT
CAGED OR FREE
RANGING LIZARD
F
F
C
C
C
C
F
F
F
F
F
DAYS LIZARD MONITORED
8
10
6
11
13
11
15
12
13
20
19
CLARITY OF TB TREND
MEDIUM
MEDIUM
HIGH
HIGH
LOW
HIGH
HIGH
HIGH
HIGH
HIGH
MEDIUM
MAXIMUM Tb
41.5
41.0
43.8
43.7
39.9
41.9
42.1
42.9
42.6
43.5
42.9
42.9
42.3
42.3 1.2
DAY OF MAXIMUM TB
1
2
5
4
1
4
3/15
2/6
3
4
3
HOURS AFTER RELEASE
TB MAXIMUM RECORDED
4
28
100
75
4
72
49/339
25/125
55
73
48
48 31
LOW DAILY MAXIMUM TB
37.6
37.4

38.7
37.5
36.3
39.4
38.7
35.9
37.8
37.6
37.7 1.1
DAY OF LOW DAILY
MAXIMUM Tb
5
8
9
6
8
11
4
13
10
6
TIME OF Tb MAX TIME
OF LOW TB MAX (HOURS)
91
145
117
117
91
216/73
48/50
235
140
72
116 60
RANGE OF DAILY
MAXIMUM Tb'S
3.9
3.6
5.0
2.4
5.6
2.7/3.5
3.9/4.8
7.0
5.1
4.7
4.4 1.3


Table 3. Continued.
LIZARD 23456
MEAN Tp ON DAY OF
MAXIMUM Tb
40.8
40.0
40.8
41.9
38.8
MEAN Tp ON DAY OF
LOW DAILY MAXIMUM Tp
38.0
37.4

39.3
36.5
RANGE OF MEAN
2.8
2.6

2.6
2.3
7
8
9
10
11
3
40.7
40.6
41.9
41.7
42.4
39.9
41.4
38.6
38.0
39.2
39.9
37.1
38.1
38.1
2.7
1.4/2.7
1.8/2.5
2.8
3.3
0.5
MEAN S.D.
40.7 1.2
38.2 1.1
2.3 0.8
o-
oo


49
and Burleson 1944, Licht 1965b, Licht and Basu 1967, Kluger et at.
1975). The body cavity internal to the wound sites and surrounding che
transmitter suppurated only slightly or net at all at the time trans
mitters were removed. The mean time between release and the maximum
hyperthermic response was 2.4 days in summer Cyclura and 1-2 days in
Dipsosaurus injected with live Aeromonas bacteria (Bernheim and Kluger
1976a). Recovery was complete in Dipsosaurus by the sixth day after
injection and, on the average, by the seventh day after the operation
in summer Cyclura. These similarities may signify that similar dis
ruptions were in force in the brain thermoregulatory centers of both
species. Hyperthermia in enterically implanted lizards may have been
a feeding response to a transmitter meal or, somehow, a reaction to
handling or the use of halothane. The lizard with the weakest response
was Number 6, the gravid caged female. Embryonic development in Cyclura
carinata begins within the female's body and continues after nesting
at burrow temperatures of 28-29 C (Iverson 1979). Regulating phase
2^'s in the range of nongravid lizards may be fatal to developing
embryos in Cyclura, as they are in Iguana iguana (Licht and Moberly
1965). Thus, the hyperthermic response to bacterial infection may
have been counterbalanced somewhat by a hypothermic response to
gravidity.
Hyperthermia also occurred in the laboratory thermal gradient
(Figure 8). The high daily mean T^ was significantly greater (p _< 0.05)
than the low in six of eight animals. The hyperthermic responses in
the field and gradient were fairly similar; that is, a latent period
of variable duration was followed by a maximum Tand a decline. High
mean T^ occurred in three species of Phrynosoma when first housed in


Figure 8. Patterns of regulating phase body temperature and
distance traveled in Cyclura caminata in a laboratory
thermal gradient. Data are presented for eight
lizards, each monitored for four consecutive days.
Lizard 3 was included in both samples. Horizontal
lines are Tfo means, vertical lines are ranges, and
open rectangles are plus and minus two standard
errors of the mean. The total distance each lizard
walked (open circles) was measured during the full
eight hour photophase.


DAY IN GRADIE
BODY TEMPERATURE
DAILY DISTANCE TRAVELED (M)


52
terraria (Heath 1965) and in Anotis aarolinensis when first introduced
into a thermal gradient (Licht 1968). The authors believed the hyper
thermia, which ceased after two days, was due to lizard inexperience in
new surroundings. Learning the new thermal environment resulted in a
decline in T^ to the preferred level. The same explanation probably
applies to Cyclura. Four of eight lizards did not move voluntarily
during the first day. Two of the lizards were located under the heat
lamps when the photophase began. It was thought best to push them
toward the cool end of the gradient when they began to pant. The steady
increase in walking distance per day corresponded in part to learning
how to maintain preferred Tfo by shuttling.
The hyperthermic response, as measured by the difference between
the maximum mean daily T^ minus the mean preferred Tis similar in the
iguanid lizards studied to date: 1) 2.0 C in Anotas cavol'ir.ens'is by the
third day in a thermal gradient (Licht 1968), 2) 2.3 C in field monitored
and 2.2 C in gradient monitored Cyol^a aarznata, 3) 3.7 C in Dipsosanrus
doTsdlis on the second day after injection of dead Aeromonas hydrophila
(Bernheim and Kluger 1976a), and 4) 2.9 C in Phrynosoma by the third
day in a thermal gradient (Heath 1965). One explanation for the
similarity in the responses is the constancy of the small temperature
gap between the preferred T, and the CTM in these lizards.
Voluntary hypothermia during the regulating phase was a rare event,
observed in only two free-ranging summer lizards (8 and 10) on the day of
their release. Lizard 8 never ventured further than a meter from its
burrow the first day, retreating six times to the burrow for lengthy
cooling periods (Figure 9). As T^ maxima increased, T^ minima increased
also, resulting in a series of ascending peaks ending with the highest


Figure 9. Hypothermia in a free-ranging lizard. Lizard 8 was in its burrow (solid
line) or lay partially exposed in the sandy depression in front of its
burrow mouth (dotted line) for a considerable part of its first day
after release.


BODY TEMPERATURE


55
peak late in the afternoon. The same pattern was observed in only 2 of
51 thermal gradient records. Hypothermia in newly released field
lizards may have been due to a temporarily heightened thermal sensitivity
due to cooling to 3-12 C during the implantation operation the previous
night.
The Relationship of Pertinent Environmental Variables
to Regulating Phase Body Temperatures
Season
Table 4 summarizes the environmental temperature data accumulated
for Site II. Mean black body temperature was not significantly different
in February and October. The mean range was only 3.1 C between February,
nearly the coldest month, and July, nearly the warmest. The mean
differences between months for the other three temperature measures
in the table were significant. The ranking was also consistent. The
February-July mean temperature difference increased from 4.2 C for sun
exposed sand and 4.7 C for shaded sand to 6.5 C for shaded air. Although
the monthly percentages of time that clouds obscured the sun did not
differ significantly, the thickness of the cloud cover and the percent
age of the sky in overcast were greater during the rainy season in
October than in February and July.
Mean monthly did not correlate directly with changing monthly
environmental temperature. Mean T, at Site II was significantly higher
in October than in February and July (p < 0.01, student's t test) and
mean s in February and July did not differ significantly from each
other (Table 5) The seasonal variation in mean Ti was probably only
slightly greater than 0.5 C (July minus February mean). The significant


56
Table 4. A seasonal comparison of some environmental variables
at Site II on Water Cay. The data were collected at
approximately hourly intervals from 0900 to 1700.
Values are for a variety of weather conditions in
February (15 days), July (16 days), and October (16
days).
N
X
X+2SE
X-2SE
HIGH
LOW
BLACK BODY TEMPERATURE
October
191
44.1
45.4
42.8
64.4
27.0
February
109
44.2
46.1
42.3
63.0
24.1
July
181
47.2
48.8
45.6
67.4
28.3
SUN EXPOSED SAND
TEMPERATURE
February
101
33.4
34.9
31.9
>52.0
23.7
October
190
35.8
36. 7
34.9
>52.0
25.1
July
177
37.6
38.5
36. 7
>52.0
26.8
SHADED SAND TEMPERATURE
February
109
26.3
26.9
25.7
36.3
21.4
October
192
29.1
29.5
28.7
39.6
25.1
July
182
31.0
31.3
30.7
35.5
26.5
SHADED AIR TEMPERATURE
(HEIGHT 6 CM)
February
109
26.3
26.7
25.9
32.6
22.1
October
193
30.5
30.9
30.1
38.1
24.2
July
181
32.8
33.1
32.5
37.2
27.8
PERCENT OF TIME SUN
OBSCURED BY CLOUDS
February
15
21.5
31.2
11.8
63
0
October
16
22.1
32.3
12.3
73
0
July
16
24.2
37.5
10.9
100
0
N
TOTAL (mm)
PRECIPITATION
February
15
5
July
16
36
October
16
83


57
Table 5. Seasonal changes in regulating phase body temperature in
the field and in the laboratory thermal gradient. Only
periods when the sun was not obscured by clouds were
included in the field calculations. Only third and fourth
day monitoring records were used in the calculation of mean
T-n's for gradient lizards. Values were taken from the T^
records at 15 minute intervals. All lizards were non-
hyperthermic.
MONTH
DAYS
FEMALES/
MALES
SITE
N
X Tb
X+2SE
X-2SE
SD
LOW
HIGH
FREE-1
RANGING LIZARDS
February
13
0/2
II
280
38.0
38.1
37.9
0.82
34.7
40.0
June
9
1/1
I
231
39.7
39.8
39.6
0.89
37.3
42.5
July
13
0/2
II
365
38.1
38.2
38.0
1.24
33.7
40, 7
October
13
9/3
II
268
38.5
38.7
38.3
1.25
34.5
41.0
THERMAL
GRADIENT LI
ZARDS
July-Aug
8
2/2
222
36.9
37.1
36.7
1.86
31.6
41.8
December-Jan
8
2/2
174
37.1
37.3
36.9
1.08
34.0
39.3


58
difference between June and July T^ means was probably due to the
difference in the amount of vegetation at Sites I and II. The variance
of mean T^ in February was significantly lower (p <_ 0.01) than in July
and October. The winter variance was lower primarily in the upper
portion of the T^ range (Figure 10). Lizards had been acclimatized to
the more thermally variable Gainesville climate before testing in the
thermal gradient (Table 5). Yet, summer and winter sample means did
not differ significantly on any of the first four days in the gradient.
As in the Caicos, the variance of the mean T^ for winter animals was
significantly less than for summer animals (p < 0.01) and the lower
winter variance was more evident at high T^ (Figure 10). Thus,
acclimatization to low environmental temperatures in Cyclura appears to
involve an increased avoidance of high and low regulating phase T^'s
without change in mean T
Mean T^ was significantly lower for Cyclura monitored in the
gradient than in the field (Table 5). The July-August gradient mean was
2.8 C lower than the June field mean and 1.2 C lower than the July field
mean. The difference was only 0.9 C for the winter field and gradient
samples. In Figure 11 mean field 's of active lizards have been
plotted against mean gradient 's for lizard species from several
different families. Mean gradient 2%/s are nearly always lower than
mean field T7's above a mean field Tof 37 C. Allowing T^ to rise
above the preferred gradient T^ is probably adaptive in diurnal helio
therms temporarily living in hot environments, such as Cyclura in
summer, since morning and afternoon activity periods are thus lengthened
(DeHitt 1967, Licht et al. 1966).


PERCENT OF SAMPLE
59
BODY TEMPERATURE INTERVAL
Figure 10. Seasonal change in Cyclura regulating phase body temperature in
the Caicos and Gainesville, Florida. The February and July field
distributions and the October field distributions are for two and
three nonhyperthermic free-ranging male lizards,respectively.
Each month's field sample consisted of a total of 13 monitoring
days. The two thermal gradient distributions consisted of third
and fourth day results for two males and two females.


60
There are at least three other reasons why lizards commonly have
higher field 's than gradient i^'s. First, Regal (1971) found that a
dominant Ktauberina riversiana provided with a subdominant lizard in the
same gradient box maintained a higher T^ than the same dominant lizard
monitored separately. Since gradient lizards are usually tested singly,
Tb's may be lower due to missing social interaction present in the field.
Secondly, field s measured by mercury thermometer rather chan
telemetry, as was the case for practically all the points in Figure 11,
may be biased to those lizards in full sun and plain view as opposed
to sequestered individuals in deeply shaded surface areas (McGinnis and
Dickson 1967). Thirdly, lizards are commonly not fed while in the
thermal gradient, although reptiles often have elevated active temperatures
after feeding: lizards (Cogger 1974), snakes (Regal 1966), and turtles
(Gatten 1974c).
The single anguid plotted in Figure 11, Gerrhcnotus multicarinatus,
which has such a low field temperature mean compared to its gradient
mean, is probably forced to be eurythermal due to its habitat choice.
It is a secretive, ground dwelling lizard living amongst shaded vegetation.
The lizard does occasionally bask, reaching a voluntary maximum of
approximately 35.7 C, considerably higher than the mean for active
lizards (Cunningham 1966). The field T-^' s for the five gekkonids in
the figure were measured while the animals were in their diurnal retreats.
Even though their s during nocturnal activity must necessarily conform
to air and substrate temperatures, gekkonids in their diurnal retreats
commonly seek out as high or higher than their preferred gradient
T-^'s. The high diurnal T-' s may be necessary for digestion of food
collected during the night (Bustard 1970, Piar.ka and Pianka 1976).


61
Figure 11. Relationship of mean body temperatures in the field and the
thermal gradient for active lizards from several families.
Solid triangles represent agamids, open triangle,anguid; open
squares; gekkonids; solid circles, iguanids; circles enclosing
dots, Cyctuva at Site I in June and Site II in July vs. the
summer gradient mean and at Site II in February vs. the winter
gradient mean; open circles, scincids. Data were taken from
several sources: Bradshaw and Main 1966, Brattstrom 1965, Cowles
and Bogert 1944, Cunningham 1966, DeWitt 1967, Fitch 1954,
Fitzpatrick et al. 1978, Grenot 1976, Heatwole 1976, Huey 1974,
Lee and Eadham 1963, Licht and Basu 1961, Licht et at. 1966,
McGinnis and Brown 1966, Norris 1953, Parker and Pianka 1974,
Pentecost 1974, Pianka 1971, Pianka and Parker 1972, Pianka and
Pianka 1976, Spellerberg 1972b, Vance 1973, Wilhoft 1958,
Wilhoft and Anderson 1960.


62
Egg laying was delayed in the Gainesville pen. A female captured
on April 4 laid in Gainesville on June 12, within the normal Caicos
nesting season of the first two weeks in June. However, another female,
which had been captive in Gainesville since July 30, 1974, laid in the
pen on July 25, 1975. Yet another female, captive since February 25,
1976, laid on July 25. The rate of egg development may have been slower
in Gainesville's cooler climate or shorter photoperiod (Mayhew 1961).
However, the delays may also have been partially due to a lack of nesting
burrows in the pen, resulting in egg retention. For example, a caged
female on Water Cay, captured on June 5, did not lay until June 17.
Habitat Complexity
A foraging Cyoluva aarinata, a semiarboreal lizard sometimes
traveling up to 1600 ft (488 m) a day, should receive a radiant heat
input from direct and filtered sunlight that is inversely proportional
to the density and height of the plants in its territory. If the habitat
niche of the species is sufficiently broad, the regulating phase Tjy of
individual lizards should increase significantly as vegetational cover
decreases. This has been demonstrated for the eurythermal lizard, Anolis
aoulatus (Ruibal and Philobosian 1970). Cyclura burrows were absent
from both beach and swales, infrequent in Mixed Woodland, perhaps due
to low food plant diversity, and infrequent in Mesic Coppice, perhaps
due to insufficient open areas for basking. Iguanas once lived in all
the other vegetation zones on Pine and Water Cays, reaching the highest
density in Rocky Coppice. This generalist habitat niche is not surpris
ing considering the lack of significant vertebrate competitors.


63
Monitoring sites were carefully chosen in order to find the maximum
mean T, difference due to differences in vegetational cover. The highest
regulating phase T^'s were expected in the sparsely vegetated Open Scrub
nearest the beach. However, man and his dogs had nearly extirpated the
lizards from this zone by the time my study commenced. Therefore, Site I
had to be located further inland in semi-Open Scrub bordering the Dense
Scrub. Lizard T,'s were expected to be lowest at Site II, a thickly
vegetated area covered with a mixture of Dense Scrub and Rocky Coppice,
bordering a rocky coast. The site was crisscrossed by a network of
bulldozed roads. On the average, bushes and trees were 63 percent taller
at Site II than at Site I (Table 6). Site II also had approximately 82
percent more shade, when roads were excluded from the transects.
As expected, mean T. was significantly higher at Site I, although
the difference was not very great (mean difference = 1.6 C, Table 5).
The small lizard sample sizes were compensated for somewhat by the long
monitoring times. Inland environmental temperature and wind velocity
differences between sites were negligible. Thus, the T^ difference
was probably due to the vegetational cover difference. The site
difference was significant in hyperthermic as well as nonhyperthermic
lizards (Figure 12). The mean T^ difference was slightly less for
maximally hyperthermic than for nonhyperthermic lizards. The high
Tj^ distributions were negatively skewed while the low 2^ distributions
were more normally distributed.
Caged lizards were monitored in an attempt to produce a significant
difference in mean T, by varying the amount and kind of shade available
in the lizard's habitat. It was hypothesized that, given only two
burrows as heat sinks, each with relatively constant air and substrate


Figure 12. Influence of habitat on body temperature. Each upper
frequency distribution is for three days of monitoring
of a maximally hyperthermic adult male Cyclura carinata
(Site I, solid line; Site II, dashed line). The lower
distributions are for the same two lizards cn the three
days of lowest mean 2^, unaffected by long periods of
overcast. Horizontal"lines are ranges, vertical lines
are mean Tfo's, and rectangles are plus and minus two
standard errors of the mean.


65
Tb INTERVAL


66
Table 6. Vegetational cover at Sites I and II on Water
Cay. The height of a bush or tree was
measured at approximately 7.5 m intervals along
the transects, which were 30 m long.
SITE I
SITE II
N (PLANTS)
HEIGHT OF VEGETATION
20
20
MEAN HEIGHT (M)
1.77
2.88
2 SE
0.39
0.53
RANGE
0.79-3.88
1.12-4.57
N (TRANSECTS)
PERCENT SHADE
5
5
MEAN PERCENT SHADE
43.3
79.0
2 SE
7.5
8.3
RANGE
33.1-51.7
68.9-90.4


67
temperatures below the lizard's minimum regulating phase Tmean
and the standard deviation of mean T^ would be greater than for the
same lizard given vegetational shade, two burrows, and branches,
which allowed greater convective cooling of lizards perched off the
ground. Overall, mean T^ was significantly higher in the complex
habitat (p < 0.01), but individually, this was true only for Lizard 5
(Table 7). The results may have been opposite to those expected
because of lizard hyperthermia. Lizards 5-7 had their hyperthermic
peaks while in the complex habitat, perhaps overriding the effect of
the vegetational shade and the branches. The standard deviations of
overall mean T- in the two habitats were about the same, again contrary
to expectation. The mean of the four lizards caged in the complex
habitat did not differ significantly from the two free-ranging lizards
also monitored at Site I in June (Table 5). Clearly, caged Cyoluva
required only a very simple habitat to thermoregulate as well as free-
ranging lizards.
Cloud Cover
Mean regulating phase T, decreased linearly as overcast time
increased (Figure 13). The correlation coefficient was significantly
less than zero (p <_ 0.0001 for r = -0.042, X = -0.042 Percent Cloud
Cover + 39.582). As expected, the correlation coefficient increased as
the weather got colder (June-July, r = -0.193, N.S.; Oct, r = -0.344,
N.S.; February, r = -0.541, p <_ 0.01). Mean T^ per day was never
depressed below 36.5 C by cloud cover times of less than 30 percent
of total time. Continuous overcast periods had to be fairly lengthy
to depress mean below the mean minimum T^ for clear weather, since


68
Table 7. Body temperature of Cyclura cavinata confined to cages
with two structurally different habitats. Lizards were
first monitored in the complex habitat for three to
four days, then in the simple habitat for two to five
days.
COMPLEX
HABITAT
SIMPLE HABITAT
LIZARD
DAYS
N
MEAN
rn
13
SD
2SE
DAYS
N
MEAN
tb
SD
2SE
4
3
60
39.9
1.17
0.30
2
48
40.4
1.40
0.40
5
4
101
41.1
0.85
0.17
3
95
39.6
0.87
0.18
6
3
64
38.1
1.10
0.28
5
97
37.7
1.54
0.31
7
4
104
38.8
3.54
0.69
3
90
38.2
1.15
0.24
Overall
14
329
39.7
1.63
0.18
13
330
38.8
1.60
0.18


MEAN BODY TEMPERATURE
43
0 10 20 30 40 50 60 70 80
PERCENT OVERCAST TIME
Figure 13. Effect, of cloud cover on mean regulating phase body temperature. Practically all field monitoring
is included (N = 124 days). Each symbol represents one day's monitoring; February (solid tri-
angles), June (solid circles), July (open circles), Angust-September (solid squares), and October
(open squares). The regression line was fitted by the method of least squares.


70
large adult Cyctuva cooled slowly and radiative heat input continued
during light overcast. The high mean T^ of 42.3 C during June, achieved
with an overcast time of 54 percent, was possible due to the thinness of
the cloud cover.
Lizard T-^ sometimes increased rather than decreased during over
cast periods, especially when the overcast lasted only a short time
(Figure 14). When the lizard had been heating prior to the overcast,
the post overcast T-h rise can be explained as a result of thermal lag.
However, T. also commonly rose initially during overcast after decreasing
before overcast, the maximum T^ during overcast being higher than the
previous maximum before overcast. Iguanas sought out hot substrates
avoided while the sun was shining and assumed a prostrate posture
(Figure 44, Illustrations 4 and 11). It appears that higher core Tj_
was tolerated under these conditions, since overheating, as indicated
by body surface temperature, was not possible. The noises of lizard
activity; that is, walking, chasing, arboreal feeding, and dropping to
the ground after feeding, nearly ceased during extended overcast periods.
Except for short walks, iguanas remained practically motionless until a
few minutes after an extended overcast cleared. The abruptness of the
activity cessation varied directly with the severity and duration of
the overcast. Short periods of overcast, frequently encountered,
resulted in little or no curtailment of activity. The T^ change during
overcast differed slightly with season. On the average, T-^ decreased
2.0 C per hour during February, 1.3 C during June-July, and 2.6 C
during October, the month of heaviest overcast. A large increase in
was most likely during summer overcast.


Figure 14. Effect of overcast duration on lizard body temperature change. Data
from February are indicated by open triangles, June and July by solid
circles, and October by open circles. Regression lines were calculated
by the method of least squares. All lizards were probably out of their
burrows during the full overcast period.


ro


73
Shaded Air Temperature
In Huey and Slatkin's cost-benefit model for thermoregulation
(1976), the value k represents a lizard's "thermoregulatory strategy,"
varying from perfect thermoregulation (k = 0) to passive nonregulation
(k = 1). The linear regression coefficient r for a graph of T^ vs.
shaded T' at the lizard's body surface is a good estimate of k. A lizard
living in a tropical forest would be expected to have a high r value,
since direct solar radiation near the forest floor is only patchily
available. On the other hand, regulating phase T- should be practically
independent of shaded T^ in a heliothermic lizard like CycZuva, living
in fairly open habitats. CyoZm'a, with its low r values, is a nearly
perfect thermoregulator (Figure 15; v = 0.151 in Graph 1, 0.013 in
Graph 2, 0.131 in Graph 3, and 0.154 in Graph 4),
Mean T^-T^ during the regulating phase at Site II was least in
July (X = 5.9 C, SD = 2.00, Range = 1.2-10.4 C, N = 68), greater in
October (X = 7.2 C, SD = 1.65, Range = 4.2-12.2 C, N = 67), and greatest
in February (X = 11.0 C, SD = 2.11, Range = 5.5-15.6 C, N = 58), since
shaded air temperature decreased faster seasonally than 2b,, again
expected in a heliotherm.
One problem with this analysis is that the air temperature was not
measured at the surface of the lizard, but rather at a site near the
blind. An attempt was made to select the lowest shaded air temperature
at the site. The assumption was made that this air temperature was also
available to the lizard. If this assumption is correct, the value of r
is still a measure of the degree of heliothermy. Of course, the
correlation of any instantaneous thermal input with core T^ declines as
body size increases in a lizard moving rapidly through a varied thermal
environment.


BODY TEMPERATURE
74
22 26 30 34 38 22 26 30 34 38
SHADED AIR TEMPERATURE
Figure 15. Correlation of lizard body temperature during the regulating
phase with shaded air temperature. Each graph represents the
data for one dominant male (Graph 1, Lizard 3; Graph 2, Lizard 9;
Graph 3, Lizard 11; Graph 4, Lizard 3). Tfo was independent of
shaded air temperature in all four lizards. Data were collected
on five to seven clear days in February (Site II), June (Site
I), July (Site II), and October (Site II), respectively.


75
Substrate Temperature
Heliothermic lizards are able to behaviorally maintain body tempera
ture within narrow limits, somewhat independently of ground surface
temperatures, which fluctuated with changes in daily and seasonal
thermal flux, cloud cover, and wind speed. McGinnis (1970) coined the
term "thermoregulatory flexibility" for this ability of heliothermic
lizards. Even during the hottest days, shaded substrates other than in
burrows were available at temperatures well below lizard T^ (Figure 16).
Iguanas rested quietly in the shade during maximum midday insolation
rather than retreating to a burrow, in a posture somewhere between
prostrate and upright (see Figure 44). T* was usually considerably
less than the sand surface temperature in the sun (T ) on clear
days, except during the early morning and late afternoon (upper and
middle graphs). Hyperthermic lizards in June decreased the ^ssn~-j-,
difference more than February nonhyperthermic lizards by spending con
siderable time in exposed areas. Iguanas remained above ground during
mildly heavy overcast on partly cloudy days, receiving maximum insol-
ative heat input with minimum convective heat loss by sitting in exposed
areas protected from the wind. T-, was commonly higher than T under
these conditions (lower graph). Nonetheless, T^ eventually decreased
during extended overcast. T-, decreased more slowly than T in the
late afternoon on clear days due to the longer heat retention time of
the lizard's body than the surface of the windswept sand. The negative
T -T-i difference in the late afternoon was not as great as shown in
ssn o
the middle graph, however. Large males, the last lizards to submerge,
sought out the last remaining sunspots in the habitat, unavailable to
me for temperature measurements without disturbing the lizards. Also,


Figure 16. Thermoregulatory flexibility. Tfo's of emergent lizards
have been subtracted from the sand surface temperature in
the shade (solid circles, Tssh Tj;) and from the surface
temperature of sand continually exposed to the sun (open
circles, Tssn Solid and dashed lines encompass the
Tgg-^ Tj, and Tgsn Tfo values, respectively. Solid and
dashed lines also indicate the hourly mean values (upper
two graphs only). Data for the upper, middle, and lower
graphs were collected on 14 clear days in February at
Site II, 8 clear days in June at Site I (lizards hyper
thermic), and 5 partly cloudy days in June at Site I,
respectively. Mean Tss}i, Tfo, and Tssn values are given
on the left side of each graph.


(CLEAR)
r^.
co
I L L. I i i I
CVl CD Tj- o
J L
I
J L.
J!L
J L.
-I L.
CO
I
OJ
CO
I
CO
o
<*
I
VJSSX QNV ql-Mssx
J L
CO
I
X L
J. X.
CD
i i
jJ
LU
5


78
iguanas kept their ventral body surfaces in contact with the substrate
during this period. The sand surface temperature important in con
ductive heat exchange, lying beneath the motionless lizard's body, did
not decrease as rapidly as T
^ J ssn
Patterns of Heat Transfer between Penned Iguanas and
their Environment: The Three-Transmitter Study
Heat transfer occurs everywhere on the lizard's body surface.
Dorsal heat exchanges are primarily radiative and convective while ventral
exchanges are also conductive. Heat loss due to evaporation of water from
cutaneous and respiratory surfaces and heat gain due to metabolism and
condensation of water vapor will be assumed to be approximately equal
in the following analyses and thus, ignored. Based on temperature
received from the dorsal surface, gut, and ventral surface transmitters,
four heat transfer conditions were possible: 1. The lizard simultaneously
received a net heat input from both dorsal and ventral body surfaces,
that is, both the dorsal and ventral transmitter temperatures were
higher than the gut temperature, 2. the lizard received heat dorsally
and lost it ventrally, 3. lost heat dorsally and gained it ventrally,
and 4. lost heat both dorsally and ventrally. All four heat transfer
conditions usually occurred sometime during an emergent lizard's day,
as illustrated in Figure 17 for a large male.
Three problems concerning the limitations of the transmitter
technique should be pointed out. Ideally, the time constant for
transmitter sensing of a rapid temperature change in the environment
should be infinitely small. However, this was not the case. For
example, the lizard in Figure 17 moved to the shade at 1109, resulting


Figure 17. Three transmitter records of a large male Cyc'luva aavinata in the Gainesville pen on
August 21, 1975. The lizard's black body temperature at five mm above and immediately
anterior to the rear legs, gut temperature, and ventral surface temperature, also just
anterior to the rear legs, are represented by the dot and dash line, solid line, and
dotted line, respectively. The lizard was in the shade much of the day (rectangles),
occasionally entering different plywood shelters (S2, S3, etc.). Transitions from one
heat transfer condition to another are demarcated by vertical lines and labeled by
number at the bottom of the graph. The heat transfer conditions are: 1) dorsal and
ventral surface temperatures higher than the gut (core) temperature, 2) dorsal surface
higher, ventral surface lower than the core, 3) dorsal surface lower, ventral higher
than the core, and 4) both surfaces lower than the core. The day was almost perfectly
clear.


0£l 0£,9[ 0£9I Og>l 0£,£l Qggl Q£JI Qg.OI 0£6
08


81
in a rapid change from Condition 2 to Condition 4. When the transition
time was taken from the temperature records, however, Condition 4 did
not start until 1112. The second problem was the small number of
transmitter monitoring sites. Large heat exchange differences often
occurred at different locations on the lizard's body surface. For
example, the anterior portion of the lizard was often in the shade
while the dorsal black body probe was in the sun. Also, fairly large
internal temperature differences occurred, as between the head and the
gut. It is undoubtedly technically feasible to double or triple the
number of temperature monitoring sites on a free-ranging lizard the
size of an adult male CyaZura. Thirdly, the surface probes should
have been implanted subcutaneously, since temperatures there were
probably of greater biological significance. The temperature difference
was probably minimal between the subcutaneous area and the ventral
surface as measured. However, dorsal black body temperature probably
differed somewhat from the dorsal skin surface temperature. Hopefully,
since the thermal lag time of the transmitter probe was less than the
subcutaneous area, the calculated times for the heat transfer con
ditions were not grossly inaccurate.
Several patterns of heat transfer were consistently observed during
the morning heating and regulating phase. Only Conditions 1 and 2
occurred during the morning heating phase on clear days. Typically,
lizards emerged from the plywood shelters and, after a short delay,
walked to a sunlit patch in the still partially shaded pen, where sand
surface temperature exceeded gut temperature. The resulting Condition
1 lasted a variable portion of the morning heating phase, depending on
whether, how soon, and how many times the animal moved to another sunlit


82
area. Condition 4 typically alternated with Condition 2 during the
early regulating phase, when lizards shuttled at a high rate. As
substrate temperatures rose and lizards tolerated these higher tempera
tures, Conditions 3 and 1 became more common. Lizards probably
tolerated higher body surface temperatures than core temperatures,
as evidenced by the mean maximum temperature of the dorsal black body
transmitter (X = 45.8 C, SD = 2.0, Range = 40.6-50.0) exceeding the
mean maximum of the gut (X = 38.3 C, SD = 1.0, Range = 36.1-39.7).
However, the mean maximum black body temperature did not exceed the
critical thermal maximum of the core (X = 46.2 C). Lizards commonly
returned to the plywood shelters for lengthy periods during midday hours,
where nearly stable body temperatures were eventually temporarily
attained (Figure 17). Lizards usually avoided the shaded sand avail
able underneath the shelters during this time in favor of the higher
temperatures in the shelters. The same shade choice selectivity with
resultant stabilization was observed in the field. Condition 4
predominated when the pen became shaded in the afternoon, since lizard
surface temperatures declined faster than gut temperature. By seeking
out relatively warm, recently shaded substrates and assuming a
prostrate posture, lizards temporarily stopped their core temperature
decline (Figure 17, 1630-1642). They were not usually thigmothermic
during this time, however, since ventral surface temperature did not
usually rise above core temperature.
The most common heat transfer condition was Number 2, occurring
an average of 51.3 percent of the morning heating plus regulating phases
(Figure 18, left graph). Condition 3 occurred only 2.5 percent of the


Figure 18. Partitioning of heat transfer in active penned Cyoluva aavinata. Three
large males were monitored sequentially in August for a total of eight
days. The times the lizard spent in each of the four heat transfer
conditions during morning heating and regulating phases are graphed on
the left. In the right graph, dorsal and ventral heat gains are compared
relative to dorsal and ventral heat losses, respectively. Circles
symbolize lizard core temperature. An arrow points toward or away
from a circle when dorsal or ventral surface temperature is greater
or less than core temperature, respectively. Horizontal lines are means,
vertical lines are ranges, and rectangles are plus and minus two standard
errors of the mean.


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FILES



THE THERMAL BIOLOGY CF THE TURKS AND
CAICOS ISLANDS ROCK IGUANA CYCLURA CARINATA
By
DAVID LESLIE AUTH
ISSERTATION PRESENTED TO THE GRADUATE COUNCI
THE UNIVERSITY OF FLORIDA
PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1980

ACKNOWLEDGEMENTS
I am grateful to numerous members of the faculty of the University
of Florida: to Walter Auffenberg, Lewis Berner, Robert DeWitt, Thomas
Enrmel, and Howard Wallace, for their faith in my competence as a zoolo¬
gist; to Archie Carr, John Kaufmann, Brian McNab, and Hugh Popenoe, for
their forebearance, patience, and time as members of the Ph.D. committee
to John Anderson, John Brookbank, William E. S. Carr, Thomas Hollinger,
David Johnston, James Nation, Michael O'Rand, and Ronald Wolff, for the
pleasure of being a laboratory teaching assistant for their courses; to
Gerald Olsen, for introducing me to the anesthetic halothane; to Robert
DeWitt, John Swel, Brian McNab, and Horst Schwassmann, for use of equip¬
ment; and to James Heath, whose work and brief personal contact started
my interest in lizard thermoregulation.
I thank the New York Zoological Society for the financial support
necessary to accomplish the research. The University of Florida has
provided extensive support throughout my graduate career, in the form of
teaching assxs tantships in the Zoology Department ar.d the College of
veterinary Medicine. Educational assistance through the G.I. Bill and
the Naval Reserve proved essential.
John Iverson introduced me to the habitats and habits of Cy atura
aavinata and continued to aid me in many ways during the study. Diderot
Gicca, Dagmar Werner, and Thomas Wiewandt willingly shared their knowl¬
edge of other species of Cyatuva.
ii

C. W. (Liam) Maguire and Bill and Ginny Cowles of the Meridian
Club, Pine Cay, permitted me to stay for practically nothing in the lap
of luxury while working in the field. They were most hospitable hosts.
Gaston Decker, Chuck and Cathy Hesse, and Francoise de Rouvray, also of
Pine Cay, helped me in many ways,
Alan Bolten, Jon Baskin, and William Link were superior companions
and considerate office mates. Alexis Arends, Susan Barnard, Hugh Ellis,
Michael May, Ken Prestwich, and Perran Ross gave needed technical
assistance. Karen Bjorndal, David Deitz, Dale Jackson, Howard Kochman,
Anne and Peter Meylan, and Michael Marshall all provided moral support.
I owe the greatest debt to my mother, Mildred G. Auth, my father,
Eugene F. Auth, and my brothers, Ronald and Dennis.
iii

TABLE OF CONTENTS
ACKNOWLEDGEMENTS ii
LIST OF TABLES vii
LIST OF FIGURES ix
ABSTRACT xiv
INTRODUCTION
1
METHODS AND MATERIALS 7
ECOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CYCLURA CARINATA ... 24
Climate of Water Cay 24
Diel Body Temperature Cycle of a Heliothermic Lizard 31
Regulating Phase Body Temperature in the Field 34
Temporary Hyperthermia and Hypothermia 44
The Relationship of Pertinent Environmental Variables
to Regulating Phase Body Temperatures 55
Season
Habitat Complexity . .
Cloud Cover
Shaded Air Temperature
Substrate Temperature.
55
62
67
73
75
Patterns of Heat Transfer between Penned Iguanas and
their Environment: The Three-Transmitter Study 78
Statistical Analyses of Regulating Phase Body Temperature. . . 89
Body Temperature Maxima and Minima ..... 39
Skewness of Body Temperature Frequency Distribution. . . 107
Standard Deviation of Mean Body Temperature 112
Rates of Lizard Heating and Cooling in the Field 119
Morning Heating and Regulating Phase 119
Lizard Cooling in Burrows 125
ETHOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CYCLURA CARINATA. . . 132
Lizard Activity 132
Burrow Movement 132
Control of Lizard Emergence and Submergence 132
iv

Morning Heating Phase 146
Regulating Phase 149
Foray Movement and Body Temperature 154
Feeding, Transmitter Passage Time, and Defecation. . . . 166
Drinking 168
Nasal Salt Gland Secretion 170
Time-Motion Study of Penned Lizards 171
Behavioral Thermoregulation 174
Description of Thermoregulatory Movements and Postures . 174
Shuttling Behavior 184
Sun Orientation . 187
Thigmothermia 200
Low Body Temperature Behavior 213
High Body Temperature Behavior 216
Head-Body Temperature Differences 224
The Integument: Metachromatism and Reflectivity 227
The Integument: Sensory Spots 231
PHYSIOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF CYCLURA CARINATA. . 236
Heating and Cooling Rates 236
Aerobic Metabolism 248
Heart Rate 253
Recovery from Maximum Exercise 259
DISCUSSION 270
Body Temperature Relations 270
Comparison of Cyclura carinata's Mean Body
Temperature with Other Diurnal Lizards 270
Seasonal Variation in Body Temperatures of
Active Lizards 276
Stability of the Mean Preferred Body Temperature .... 277
Late Afternoon Maximum Body Temperature 279
Thermal Safety Margins 281
Panting Threshold 282
Body Temperature and Reproductive State 284
Ethological Considerations 285
Lizard Activity 285
Eurythermy vs. Stenothermv 288
Comparative Thermoregulatory Behavior of
Heliothermic Lizards 296
Complexity of Social Behavior in Cyclura carinata
as a Function of Latitude 305
Nocturnal Lizard Aggregations 306
v

Physiological Considerations 307
Aerobic vs. Anaerobic Metabolism 307
Neural Basis for Shuttling Behavior in Lizards 312
SUMMARY OF RESULTS AND CONCLUSIONS 326
LITERATURE CITED 347
BIOGRAPHICAL SKETCH 367
vi

LIST OF TABLES
Table
1 Daily variation in relative humidity 30
2 Summary of regulating phase body temperatures of
lizards in the field 35
3 Hyperthermia in Cyolura aarzruzta in the field 47
4 A seasonal comparison of some environmental variables
at Site II on Water Cay 56
5 Seasonal changes in regulating phase body temperature
in the field and in the laboratory thermal gradient .... 57
6 Vegetational cover at Sites I and II on Water Cay 66
7 Body temperatures of Cyclura car-Lnata confined to
cages with two structurally different habitats 68
8 Skewness of frequency distributions of minimum and
maximum body temperatures 91
9 Means and standard deviations of minimum and maximum
body temperatures of individual lizards and correlation
of maximum vs. minimum and minimum vs. maximum pairs . . 92
10 Linear regression analysis of daily minimum vs.
maximum 's and mean of daily Tfo minima vs. mean of
daily Tjy maxima for individual lizards 101
11 Skewness and kurtosis of Tfo frequency distributions .... 118
12 Rates of lizard heating and cooling in the field during
the morning heating and regulating phases 120
13 Lizard cooling in burrows in the field 124
14 Lizard body temperature at emergence in the field 131
15 Emergence and submergence times, temperatures, and
light intensities for free-ranging lizards in the
field 146
16 Possible acclimatization of lizard activity 165
vii

17 Effect of body temperature on the feeding rate and meal
size of penned Cyctura oavina.ta 167
18 Time-motion study of Cyclura aarinata in the Gainesville
pen 172
19 Skin patch heating 198
20 Estimated thermal conductance during heating and cooling
of lizards in still air 240
21 Correlations of lizard behaviors with body temperature
during heating and cooling in the cage and on the jig . . . 245
22 Recovery oxygen for Cyclura carinata after seven minutes
of maximum exercise 265
23 Comparison of actual mean recovery time after maximum
exercise with recovery times at three hypothetical rates
of oxygen utilization 267
24 Field body temperatures of diurnally active lizards .... 271
viii

LIST OF FIGURES
Figure
1 Black body temperatures in the field 25
2 Air temperature in the shade at Gainesville, Florida
and Grand Turk, British West Indies 26
3 Mean maximum wind speed at Site II in July 28
4 Monthly precipitation on Grand Turk, British West Indies. . 29
5 Body temperature record of Lizard 2 on February 10, 1976. . 32
6 Summary of field body temperatures during the regulating
phase 45
7 Lizard hyperthermia and hypothermia during the regulating
phase 46
8 Patterns of regulating phase body temperature and distance
traveled in Cyctura oarznata in a laboratory thermal
gradient 51
9 Hypothermia in a free-ranging lizard 54
10 Seasonal change in Cyclura regulating phase body
temperature in the Caicos and Gainesville, Florida 59
11 Relationship of mean body temperatures in the field and
the thermal gradient for active lizards from several
families 61
12 Influence of habitat on body temperature 65
13 Effect of cloud cover on mean regulating phase body
temperature 69
14 Effect of overcast duration on lizard body temperature
change 72
15 Correlation of lizard body temperature during the
regulating phase with shaded air temperature 74
16 Thermoregulatory flexibility 77
ix

17 Three transmitter record of a large male Cyclura
carinata in the Gainesville pen on August 21, 1975 80
18 Partitioning of heat transfer in active penned
Cyclura carinata 84
19 Rate of core body temperature change during the four heat
transfer conditions 86
20 Correlations of the sum of daily heat transfer conditions
with lizard activity and mean body temperature 88
21 Time distributions of four combinations of maximum and
minimum body temperature 97
22 Correlations of maximum vs. minimum Tfo and the mean of
Tfo maxima vs. minima for individual lizards 100
23 Hourly change in minimum and maximum body temperatures. . . 104
24 Timing shift of daily minimum and maximum T¿>' s 106
25 Skewness of daily Tq frequency distributions 110
26 Comparison of the level and precision of thermoregulation
of Cyclura carinata and Dipsosaurus dorsalis in continuous
thermal gradients Ill
27 Correlation of the standard deviation of mean daily
with mean daily Ty 113
28 Effect of body weight and mean Tfo on the standard
deviation of mean 116
29 Hourly change in the mean and standard deviation of lizard
body temperature and shaded air and sand temperatures . . . 118
30 Correlation of mean rates of heating and cooling in the
field with lizard body weight 124
31 Correlations of parameters describing regulating phase
body temperatures 126
32 Lizard movements in a burrow 134
33 Seasonal change in lizard emergence and submergence times . 136
34 Correlations of times of emergence and submergence with
body weight in penned Gainesville lizards 140
35 Gradual reentrainment of lizard emergence time after
release 145
x

36 Lizard activity during the morning heating phase 148
37 Daily and seasonal variation in activity 151
38 Variable inhibitory effectiveness of cloud cover on
lizard activity 155
39 Influence of cloud cover on the length of lizard forays
and the mean rate of travel during forays 159
40 Correlations of foray length and rate of travel with mean
body temperature 161
41 Body temperatures during forays 163
42 Defecation body temperatures and timing 169
43 Correlation of lizard mobility with confinement area. . . . 173
44 Thermoregulatory motions and postures of Cyolura oarinata . 175
45 Lizard shuttling in the summer field cages 185
46 Midday depression of shade seeking body temperature .... 188
47 Lizard orientation to a heat lamp during heating and
cooling in the photothigmotron 189
48 Positive and negative orientation during lizard heating
and cooling 192
49 Daily and seasonal variation in sun orientation in the
Gainesville pen 195
50 Left vs. right side sun orientation in the Gainesville
pen 196
51 Daily change in the pattern of travel during forays .... 199
52 Participation of the lizard's head, neck, and legs in the
reduction of substrate contact 203
53 Relative importance of lateral compression of the trunk
vs. head, neck, and trunk elevation 205
54 Conductive and convective heating and cooling of a single
Cyolura oar-inata 206
55 Conductive heating and cooling of a single Cyolura
oarinata, with and without simultaneous radiative
heating with a heat lamp 209
xx

56 Thigmothermic responses of a single Cyclura caminata to
periods of increasing, stable, and decreasing substrate
temperature 212
57 Behavioral responses of Cyclura carznata at low body-
temperatures 214
58 Daily adjustment of the panting setpoint to changing
heat input 220
59 Correlation of total panting time with head and gut
thermal lag times 222
60 Head-body temperature differences during heating and
cooling of a 1790 g male Cyclura carznata in the
Gainesville rooftop arena 226
61 Integumental and substrate reflectivities 228
62 Density of sensory spots on different parts of the lizard
body surface 234
63 Heating and cooling rates of a 1435 g male Cyclura
carznata in a constant temperature chamber 239
64 Breathing rates of lizards during heating and cooling in
a constant temperature chamber 247
65 Correlation of resting and maximum oxygen consumption
with body temperature . 250
66 Analyses of aerobic metabolism 252
67 Heart rate as a function of body temperature 254
68 Heart rate analyses 256
69 Relationship of oxygen consumption to heart rate at
different body temperatures 257
70 Temporal pattern of body movements while struggling to
escape 260
71 Effect of body temperature on the rate of recovery from
seven minutes of maximum activity 263
72 Early and late phase recovery from maximum exercise in
Cyclura carznata at different body temperatures 269
73 Summary of body temperatures of selected behaviors and
physiological maxima of Cyclura carznata 293
xii

74 Effect of increasing body temperature on frequency
distribution skew and standard deviation of the mean
for minimum and maximum body temperatures
75 Hypothetical neural basis for changes observed during
increasing mean body temperature in Cyolura carinata. . .
. 319
. 324
xiii

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
THE THERMAL BIOLOGY OF THE TURKS AND
CAICOS ISLANDS ROCK IGUANA CYCLURA CARINATA
By
David Leslie Auth
June , 1980
Chairman: Walter Auffenberg
Major Department: Zoology
The ecological, ethological, and physiological thermal biology of
Cyclura carinata, a large, tropical, herbivorous iguanine lizard, was
investigated on Water Cay in the Caicos Islands and in Gainesville,
Florida. The study lasted from June, 1974, until September, 1976, and
included four field trips totaling twenty weeks.
Far-field transmitters were placed in the coelom or gut of free-
ranging adult iguanas (675-1864 g) for long-term monitoring of core
body temperature (T^) in the field. Mean regulating phase T^ ranged
from 38.0 to 39.7 C (February-October), quite high compared with most
other iguanids. Stenothermia was pronounced, with a mean, regulating
phase range of only 3.3 C. Maximum voluntary T^, 43.8 C, was only
2.4 C below the CTMax. The CTMin, 12.8 C, almost equalled the lowest
recorded environmental temperature.
Iguanas were initially hyperthermic in both the field and a
thermal gradient, the former the possible result of bacterial infection
or a transmitter meal and the latter of thermoregulatory learning.
xiv

Individual differences in regulating phase T^ were partially due
to variable vegetational cover and, perhaps, egg development in females.
Mean operant T^'s did not correlate perfectly with seasonal change in
the ambient thermal environment. Mean regulating phase field T^
exceeded mean preferred T-^. The latter did not change with Gainesville
seasonal acclimatization, whereas winter variance of mean T^ was less
than summer variance in both field and thermal gradient.
Mean regulating phase Th was inversely correlated with cloud cover
time, the correlation improving with seasonal cooling. T^ was nearly
independent of shaded air temperature.
Heat transfer through the dorsal and ventral body was examined
using three transmitter placements on penned iguanas. Heat was usually
gained dorsally and lost ventrally; iguanas were thigmothermic only 20.7
percent of their active period, usually in combination with heliothermia.
Individual frequency distributions of regulating phase maximum and
minimum T-,' s were commonly skewed positively and negatively, respectively,
whereas inclusive individual distributions were usually negatively
skewed. Standard deviations were slightly but not significantly greater
for mean minimum than mean maximum T-'s. Change in T^ between any two
consecutive maxima was positively correlated with change in two
consecutive minima, with the first minimum immediately before or after
the first maximum. Individual mean maximum and minimum T^' s were
directly and linearly correlated, the regression line extrapolating
to approximately the maximum voluntary tolerance. These results are
discussed in terms of dual-limit thermoregulation.
Iguanas were diurnally active all year and rarely escaped into
burrows due to overheating. Mean regulating phase heating was 1.15

times faster than cooling and 3.0 times slower than heating during
morning basking. Iguanas also heated faster than they cooled in a
constant temperature chamber, both when restrained and when permitted
to move about. Maximum T^ commonly occurred during late afternoon
basking. This and other results indicate that surface rather than
core body temperature is regulated.
Both length and rate of travel were independent of mean T^ between
35.0 and 40.5 C. Arboreal climbing, burrow excavation, defecation,
drinking, feeding, nasal salt gland secretion, and panting were also
examined in relation to mean T-, .
b
Thermoregulatory behaviors and postures were described. A
photothigmotron was constructed to quantify iguana light shadow and
substrate contact in a controlled thermal environment. Shuttling and
sun orientation were examined. A time-motion study revealed the low
activity rate of penned iguanas and activity partitioning between
foraging, social, and thermoregulatory behaviors. Thermoregulatory
behavior was quite similar to other heliothermic iguanids.
Resting and maximum oxygen consumption, and heart rate and
oxygen consumption during recovery from maximum exercise were determined
at different s. Iguanas fatigued rapidly during maximum exercise,
yet rarely developed large oxygen debts in the field. Heart rate
increment, initial rate of recovery, and oxygen scope were greater
in the preferred T^ range.
xvi

INTRODUCTION
Although much is known regarding the thermal biology of small
lizard species (see Heatwole 1976 for a review), large lizard species,
because of their rarity in numbers and usual distribution in isolated
areas, have received lesser attention. This is unfortunate, for large
mass confers certain advantages and disadvantages due to low heating
and cooling rates. This study investigates the thermal biology of a
species of rock iguana in the genus CycZura Harlan, one of nine genera
in the subfamily Iguaninae (Avery and Tanner 1971). Although the genus
includes some of the largest lizards in the world, CycZura caminata is
one of the smaller members (maximum weight of 1,864 g for males,
1,135 g for females).
Preliminary observations by Auffenberg (personal communication) in
1973 indicated that CycZura caminata probably had a fairly high diurnal
operant or eccritic body temperature (2^). This, coupled with large size and
a herbivorous diet, uncommon in lizards, suggested that its thermal biology
could in some ways be unique. A great deal of thermally related data had
already been collected for the desert iguana, Dip sosaurus dorsalis, a small
herbivorous lizard of the North American Southwest with a very high
preferred (Porter et at. 1973). Thermally related comparisons of
CycZura oarinata with this species as well as with the larger iguanines,
Amblyrhynohus cristatus (White 1973), Iguana iguana (McGinnis and Brown
1966), and Sauromalus obesus (Johnson 1965) seemed worthwhile.
Continuous monitoring of the 2V of free-ranging lizards in the
field using radio-telemetry forms the basis for the present study. This
.1

2
approach has only rarely been followed for large lizards (Stebbins and
Barwick 1968, McNab and Auffenberg 1976).
The natural history of Cyalura aarinata was examined simultaneously
from four directions by Dr. Auffenberg (feeding strategy), John Iverson
(autecology and ethology) and myself, under a grant from the New York
Zoological Society. The field site was located primarily on Pine and
Water Cays, on the northwest side of the Caicos Island Bank. Since the
goal of this research was to gain as much information as possible which
could be useful in guaranteeing the species' future survival, I chose to
attack the problem of thermal biology broadly, rather than emphasizing
any one aspect such as thermoregulation. Ecological, ethological, and
physiological approaches were taken, working only at the organismic
level and using free-ranging field, controlled field, and laboratory
studies. This synthetic approach has been utilized rather successfully
by Grenot (1976) for Uromatix aaanthinurus, another fairly large
herbivorous lizard.
The field ecological approach to Cyalura aarinata's thermal
biology depends on a thorough understanding of the thermal environment
in which the lizard lives. This means that cloud cover, wind speed,
black body temperature, shaded air temperature, and substrate tempera¬
ture have to be monitored daily and seasonally as well as the lizard's
body temperature and its thermally related behavior. Then, environ¬
mentally related questions can be answered. How does the lizard
respond to the rather constant easterly winds? Are these xeric habitat
animals able to stay out of their burrows even during the hottest part
of the summer day or are they forced underground, as is the smaller

3
Dipsosaurus dorsalis (Porter et al. 1973)? Is T^ independent of shaded
air temperature, as expected in a heliotherm? The populational dis¬
tribution on the islands should be examined in terms of habitat types
available and any variation in mean T\ with vegetational density
determined. The iguana distribution (before the beginning of the study)
was minimally affected by introduced mammalian competitors, since the
human population was small and the only other mammal was the introduced
Norway rat.
Continuous T^ records of free-ranging lizards provide a large
amount of analyzable data, as shown by Stebbins and Barwick (1968) from
their monitoring of a single lace monitor lizard. Maximum and minimum
T-0's during the regulating phase, times of emergence and submergence,
and heating and cooling rates can easily be extracted from the record.
Then, many interesting questions can be addressed. How do the maxima
and minima vary during the day on clear days? Does body temperature
variability decrease as body weight increases? Does this variability
decrease as mean T^ increases? How is mean Th related, if at all, to
reproductive state, sex, and size?
Heath (1964) and DeWitt (1967) were two of the first investigators
to report a significant temperature differential between the head and
body of lizards during heating and cooling, calling into question the
significance of measuring just cloacal temperature and bringing up the
question of what body temperature is regulated. Since then, many
workers have simultaneously measured more than one body temperature in
lizards (Johnson 1972, Webb, Johnson, and Firth 1972, Crawford 1972,
Spray and Belkin 1972, 1973, Crawford and Barber 1974, Parmenter and

4
Heatwole 1975, Pough and McFarland 1976, McNab and Auffenberg 1976,
Crawford et al. 1977). For a large lizard like Cyolura earznata,
multiple body temperature measurements are advisable as part of the
study. Head-gut and dorsal black body-gut-ventral body surface tempera¬
tures were measured to determine internal temperature differences and
dorsal and ventral heat transfers with the environment, respectively.
Lizards had to be confined in outdoor pens to maintain continuous visual
contact, a serious problem with free-ranging lizards.
Two aspects of the ethological thermal biology of Cyotupa
savznata are investigated. The first addresses the question of how
daily variation in the thermal environment and T^ influence lizard
activity. Lizard movement in burrows, emergence and submergence timing,
movement during the morning basking and regulating phases, feeding rate,
total food consumed, passage time of transmitters through the gut,
defecation timing, and secretion timing of the nasal salt gland are
considered. Lizards vary considerably in the amount of time spent
in actual motion. "Widely foraging" predators, such as varanids and
teiids, are generally more active than "sit and wait" insect feeding
iguanids (Pianka 1973). Herbivorous lizards should fall somewhere in
between, if they feed on a variety of plants, since they must go to
where the plants are located and then actively pick items up. The
partitioning of the lizard's active time between feeding, social, and
thermoregulatory movements should vary consistently between each of
these three dietetic categories. Ideally, a continuous time-motion
study should be conducted in the field, as accomplished for Egevnia
aunnznghami by Wilson and Lee (1974). However, this proved impossible

5
from a fixed blind, due to vegetational density and the considerable
distance traveled by adult iguanas. Penned lizards were observed
instead.
The second ethological approach is to discover how Cyclura
carinata regulates its body temperature and then to compare this
behavior with that of other lizards, reported on extensively in the
literature. This has traditionally been the emphasis of lizard thermal
biology studies, since the time of Cowles and Bogert (1944). The roles
of thermoregulatory postures were comprehensively catalogued in a
qualitative manner, similar to Brattstrom's work on Amphikolurus
barbatus (1971) and Greenberg's on Sceloporus cyanogenys (1977).. Some
of the behaviors were quantified under controlled field and laboratory
conditions. A laboratory device was built to quantify the degree of
sun orientation and thigmorhermia, using a photographic technique
similar to Heath's for Phynosoma (1965). I did not attempt to find out
to what extent body posture controlled T-r> under a defined set of
thermal conditions, as Muth (1977b) and Porter and James (1979) have
recently done.
Lizards usually heat faster than they cool (Bartholomew and
Tucker 1963, 1964, Morgareidge and White 1969, Weathers 1970, Weathers
and Morgareidge 1971, Weathers 1971, Baker et oil. 1972, Ellis and Ross
1977). This is attributed to increased peripheral blood flow during
heating. The ratio of heating to cooling rate increases as body mass
increases in the American alligator, to the extent that while hatchlings
heat and cool at about the same rate, a 1000 kg adult would theoretically
heat nearly three times faster than it cools (Smith 1979). The testing

6
of Cyalura aar-inata is justified on comparative grounds and because of
its large size, especially since the data obtained complement the
heating and cooling rate results for the species in the field.
Many rate processes become somewhat temperature independent in
reptilian preferred T^ ranges (see Dawson 1975 for a review). Selected
physiological parameters—resting and maximum oxygen consumption, rest¬
ing and maximum heart rate, and oxygen consumption during recovery from
maximum exercise—were measured in Cyalura aari'riata to see how they
varied with The study of recovery from maximum exercise was
especially interesting, since the amounts and kinds of voluntarily
tolerated, strenuous activity the species engaged in were observed in
the field. Bennett's methodology (1972) for his comparison of oxygen
consumption, oxygen debt, and heart rate in Varanus gouldii and
Sauromalus hispidas was closely followed, permitting comparison
with results obtained for Cyalura.

METHODS AND MATERIALS
The field study site was located in the Caicos Islands, between
21° and 22° North latitude and 71° 30' and 72° 30' West longitude,
approximately 150 km north of Hispaniola and 890 km southeast of Miami,
Florida. Pine Cay was the base of operations, a privately owned island
with excellent lodging, potable water, an air strip, and a large popu¬
lation of Cyalura cari,nata (as of August 1973). Field work was conducted
primarily on Water Cay, connected by a traversable sand bridge to Pine
Cay. Iguana density was adequate here at the beginning of the study, no
human habitation was present, and bulldozed paths provided easy access to
most areas at the northeast end of the island. The two islands are
located on the northern perimeter of the triangularly shaped Caicos
Bank, a shoal with a northern base of 125 km and a southerly directed
altitude of 75 km. Water Cay has an area of approximately 225 ha, with
a leeward northwestern sandy beach, a windward eastern rocky limestone
coast, and a southern mangrove swamp. The height and density of the
vegetation increases with island age and soil depth and quality from the
northwestern to the southeastern side of the island. The highest point
is approximately five meters above sea level.
I spent a total of 140 days in the field during four trips:
June 1-July 28, 1974; September 30-November 4, 1975; February 4-23, 1976;
and August 17-September 11, 1976.
Two field sites were chosen on the. eastern end of Water Cay. Site I
was in level, sandy, Semiopen Scrub, 600 m from the NE side and 275 m from
7

8
the NW side of the island. Site II was in Dense Scrub, with Rocky
Coppice bordering the Pine-Water Cay inlet, 200 m from the SE corner of
the island (see Auffenberg, in prep., for descriptions of vegetation
types). Work was confined to approximately one hectare at each site.
Site I bordered a bulldozed road and Site II was crisscrossed by an
interconnected road network.
A moderately large hotel complex was completed on Pine Cay in the
summer of 1973. By the time of my first trip, iguana populations had
already declined on both Pine and Water Cays due to predation by man,
dogs, and feral cats. The subsequent collapse of the population on Pine
Cay has been documented by Iverson (1977). Cyelura density at Site II
was estimated at 10 adults per ha in July, 1974. Density was somewhat
less at Site I. Iguanas had practically disappeared from Site I by
September, 1975, necessitating a concentration of effort at Site II.
Iguanas had been extirpated from Site II by August, 1976, resulting in
a shift of field work further southwest along Water Cay and to Port
George Cay, just north of Pine Cay. Due to the small number of hatchling
and juvenile iguanas, the study had to be restricted to adults.
Eleven male (X = 993 g, 435-1715) and four female (X = 748 g,
528-1135) iguanas were brought to Gainesville, where they were kept in
two outdoor galvanized sheet metal pens. One pen was circular, enclos-
? 2
ing an area of 16 m , and the other was rectangular, enclosing 18 m .
Eight plywood boxes, each measuring 40 cm wide by 40 cm long by 10 cm
high, were perched on concrete blocks and provided shade, shelter, and
basking platforms. Substrates were beach sand and grass in the circular
and rectangular pens, respectively. From October 1 to March 30, the

9
pens were partially or completely covered with 4 ml polyurethane plastic
to increase temperatures. An electric forced air heater kept temperatures
above freezing on exceptionally cold nights. The captives were fed
bananas, green beans, white grapes, and fish-flavored catfood.
Several environmental variables were monitored in the field.
Measurements were taken hourly at a single location near the blind at
Sites I and II, where an open, sun exposed area bordered an area of
fairly dense vegetation. Changing temperatures between readings were
estimated to occur linearly. Substrate temperatures were taken on dry
sand with a Number 421 banjo probe attached to a Model 46 Tele-Thermometer
(Yellow Springs Instrument Co.). Air temperatures were taken at a height
of 2-3 cm with a Number 405 air probe. A black body for estimating
daily variations in solar heat input was constructed by placing a
mercury thermometer in a groove carved in a balsa wood block. The block
was then covered with a thin brass plate (0.4 mm) and coated with flat
black paint. A record was kept of the times the sun was obscured by
clouds. Light intensity was measured with a Type 213 light meter
(General Electric) directed from a height of 20 cm at a piece of white
painted cardboard placed on the ground. Instantaneous wind speed was
measured with a Dwyer wind meter. Average wind speed was determined
over a one minute period with an Air Meter Model W131 (Weather Measure
Corp.). Relative humidity was measured with a sling psychrometer.
Core body temperatures of free-ranging iguanas were measured with
Model L far-field transmitters (Mini-Mitter Co.). The transmitter was
a crystal-controlled blocking oscillator operating in the citizen's
band range (27 MHz). Osgood and Weigl (1972) describe the circuit diagram

10
and construction. The signal, a series of beeps increasing in frequency
with temperature, was received with a Model HA-420 three-channel walkie-
talkie (Lafayette) with an attached beat frequency oscillator. Trans¬
mitter range was approximately 200-300 feet, depending on the topography.
Accuracy was +0.5 C. The transmitter was powered by two hearing aid
batteries (Mallory MS 76), which had a working life of one to two months.
The transmitter was embedded in epoxy resin and enclosed in a plastic
capsule. The capsule was then coated with a mixture of 50 percent
paraffin and 50 percent beeswax, followed by dental acrylic (NuWeld,
I.D. Caulk Co.). The package weighed about 15 g and was smaller (1.7 by
3.7 cm) than a Cuolura egg. A single mercury thermometer (Scientific
Products), checked for accuracy against a NBS certified bomb calorimeter
thermometer (Parr Instrument Co.), was used to calibrate all transmitters
used in this study. Most transmitters were recovered after the monitor¬
ing period. Drift never exceeded 0.5 C.
During the summer field trip of 1974, a transmitter was implanted
in each of eight adult iguanas. The operation was performed during the
night of the day of capture. The lizard was fastened to a wooden jig
similar to that used by Bartholomew and Tucker (1963). The individual
was then placed in a large freezer cabinet until body temperature had
fallen to 2-5 C. T^ was monitored intestinally with a Tele-Thermometer
Probe Number 701 inserted approximately 10 cm into the distal colon
and taped to the tail. The individual was kept on ice during the
operation. The transmitter was inserted through a dorsoventral
incision made on the right lateral abdominal wall about 1 cm anterior
to the. rear leg. The incision was closed with nylon fishing line.

11
Individuals were released the day following the operation and monitored
immediately. During subsequent field trips, transmitters were placed
in the gut immediately after capture. A gauze pad soaked with halothane
(Ayerst Laboratories) was positioned at the bottom of a 250 ml beaker.
See McDonald (1976) for a discussion of this inhalation anesthetic. The
beaker was then slipped over the lizard's head. A rubber apron taped
to the mouth of the beaker extended over the forelegs, reducing the rate
of loss of gas anesthetic. Lizards were usually apneustic when the mask
was initially placed over their heads, the duration of apnea being
inversely correlated with body temperature. The drug took effect within
a minute after breathing commenced. Complete anesthesia lasted between
five and ten minutes, and recovery was seemingly complete within an hour.
All individuals were weighed (Ohaus Dial Spring Scale), measured (snout-
vent and total length), examined for scars and ectoparasites, and marked
on both sides of the trunk with a bright orange Arabic numeral (Aero-
gloss paint, Pactra Industries).
The body temperatures of eleven different, free-ranging adult
iguanas were monitored during the study (10 males, 1 female). Two males
were monitored on two separate trips. Recording periods extended from
three to twenty days. Temperature readings were taken every five to
fifteen minutes from before the individual emerged up to two hours
after it submerged for the day.
I observed and monitored iguanas from a blind during the first
field trip. Lizards were attracted to the vicinity with a sardine bait
and noosed from inside the blind. Less timid individuals were noosed
without bait and blind.

12
Iguanas were less timid during che second and third trips, per¬
mitting more thorough observation of specific individuals by continuous
tracking (6 adult males tracked 1-6 days, X = 4 days). Following initial
transmitter placement, iguanas were monitored without further visual
contact until the start of the second or third day. Since iguanas
usually retreated into their burrows when disturbed during the initial
basking period, visual contact was always delayed for a time after
emergence. I crawled along behind the lizard at a distance of approxi¬
mately five meters during its feeding forays. Visual contact was
intermittently lost when the individual moved off into dense scrub. The
lizard was usually quickly resighted by following the same direction of
travel. Signal strength, as perceived by the human ear, reached a
maximum approximately three meters from the transmitter. Bushes along
the route of travel were flagged with orange plastic tape marked with the
dace and time. A crude map of the route was recorded in a notebook,
along with observations of the individual's behavior. The locations of
the flags were later marked on a detailed map of the site. Average
rates of travel and distances traveled could then be calculated at
leisure.
Near-field transmitters, built according to Mackay (1970), were
used during the study. They were cylindrically shaped, measuring approxi¬
mately 1 by 3 cm and weighing 4 to 6 g. The signal, a series of clicks
increasing in frequency with temperature, was received at a maximum
distance of one meter with an AM transistor radio (50-160 KHz). The
package was sealed in the same manner as the far-field transmitters.
Iguana movement in the burrow and T^ were monitored
simultaneously during the second and third trips by placing both a

13
near-field and a far-field transmitter in the gut of a single large male.
After the lizard submerged for the night, its position was located by
moving the AM radio to where the signal was strongest. The radio was
again placed directly over the lizard early the next morning prior to
emergence. The signal was monitored at the blind six to nine meters away
from the burrow by listening through an earplug connected to the radio.
When the lizard moved, indicated by fading or loss of the signal, I
quietly moved the radio to the lizard's new position and returned to the
blind. This procedure was repeated until the lizard emerged. To see
how far it would submerge, the lizard was deliberately scared into its
burrow at different times of day. The location of the burrow terminus
was determined by digging at the end of the monitoring period.
In June of 1974 body temperatures of four iguanas were monitored
two at a time in two field cages at Site I. The cages were constructed
of quarter inch hardware cloth in the shape of a cylinder 1 m high
by 2 mlong. Two, 15 cm diameter, 1 m long cylinders were attached to
one end of each cage and covered with sand to simulate burrows. Shellac
coated copper wire was wrapped around the entire cage to act as an
antenna for the near-field transmitters. Strands were approximately
10 cm apart. Each two leads were attached to a circuit breaker connected
to two similar leads from wire wrapped around the ferrite antenna of the
AM radio. Thus, monitoring the 2^ of one lizard could be immediately
followed by monitoring the second by switching circuits. Individuals
were kept in a complex habitat during the first four to seven days.
One-half of the cage was covered with thatch palm fronds and branches
were arranged beneath the fronds inside the cage, a 11owing the iguana

14
to get nearly one meter off the ground while still remaining in the
shade. The burrows provided the only shade during the next two to three
days of monitoring, in what was called the simple habitat. The body
temperatures of two caged and two free-ranging iguanas could be monitored
simultaneously from the blind.
Preferred 2V'S were determined in two thermal gradients constructed
of plywood, each 2.2 m long, 0.4 m wide, and 0.6 m high, with tops
covered with quarter inch hardware cloth. Two 250 w heat lamps were
suspended at one end of each gradient; their height was adjusted so that
the black body temperature immediately beneath them was 60 C, slightly
below the lowest monthly maximum recorded in the field (63 C in February).
The gradients were placed in a controlled environmental room (Environ-
Room, Lab-Line Instruments, Inc.) set at an air temperature of 25 C (+ 1.0
C accuracy, air velocity next to the lizard less than 0.25 m/second).
The air temperature in the cool end of the gradients gradually increased
to a maximum of 28-30 C after four hours of the eight hour photoperiod.
The photoperiod was centered at 1200 E3T. A 40 w fluorescent bulb
provided less than ten footcandles of illumination. Preferred 's
were determined for eight adult iguanas, a group of four acclimatized
to Gainesville temperatures both in July-August and December-January.
Summer and winter lizards were monitored individually, for four to
eight consecutive days, respectively. Monitoring commenced the day
following transmitter placement. Lizards received neither food nor
water while in the gradient.
Low body temperature behavior was examined in six adult Cyelura
acclimatized to Gainesville temperatures in May. Starting at 1200-1600
(all times reported in this study are EST), the individual was cooled in

15
the environmental room set at an air temperature of 5-8 C. Body tempera¬
ture was monitored colonically with the Tele-Thermometer. The losses of
seven behavioral responses were noted as T^ decreased: 1. crawling,
2. turning over when placed on the back (critical thermal minimum),
3. maintaining a horizontal head orientation when the body was rotated
about its long axis, 4. maintaining a constant field of view by rotating
the head or eyes when the body was rotated laterally, 5. moving the legs
when pinched while resting on the back, 6. bloating, and 7. closing and
opening the eyes. Each lizard was cooled an additional 3 C, then exposed
to room temperature outside the environmental room. The lizard's body
temperature was again recorded as each of the seven behavioral responses
was regained.
To examine the timing and degree of heat transfer through the dorsal
and ventral surfaces of the iguana's body, three-transmitter experiments
were run in a circular Gainesville pen in late August and early
September. Two far-field transmitters were fastened with electrician's
tape to the outside of a piece of television antenna cable. When the
cable was wrapped around the lizard's waist just anterior to the rear
legs, the transmitters were situated ventro-laterally on either side of
the trunk. The cable ends were held together dorsally with a rivet.
The temperature-sensitive thermistor bead, embedded in epoxy resin, was
located at the end of a short probe wire extending from one end of the
transmitter. One probe tip was taped to the middle of the ventral side
of the cable, to monitor the body-substrate interface temperature
immediately under the lizard. The other, painted flat black to monitor
dorsal heat input, was suspended free in the air about 1 cm anterior
to the cable and 5 mm above the middle of the lizard's lower back.

16
A near-field transmitter was placed in the lizard's gut. Three large
males (1195, 1240, and 1260 g), monitored one at a time for a total of
nine days, moved freely about the pen. Seven other iguanas were also
present. Readings from the three transmitters were taken every five to
ten minutes. To obtain gut temperature readings, the AM radio was taped
to the end of a wooden pole and extended from outside the pen to within
a meter of the lizard. Shaded air, sun-exposed sand, and black body
temperatures were also measured. The lizard's behavior was observed
and notes taken for later correlation with the transmitter records.
To determine whether feeding rate or meal size was significantly
less immediately after morning emergence than during the higher T-^'s of
the regulating phase, iguanas in the Gainesville pen were presented with
known weights of white grapes. Food was either placed on trays on the
ground, before individuals emerged for the day, or on trays affixed to
the tops of posts, after individuals had reached regulating phase body
temperatures. The elevated trays were large enough for only one
individual to sit on, preventing feeding by several animals on a single
known weight of grapes. Also, by placing grapes on three post tops
simultaneously, one feeding lizard was rarely displaced by another.
Animals were fed on clear days after at least two days with no food.
Sun orientation directions of lizards were measured with a hand
held Silva compass. Records were obtained in the field, the Gainesville
pen, and a 2 m circular arena placed on top of the zoology building. To
determine the importance of vision in sun orientation control, lizards
were blindfolded by placing ellip.tically shaped plastic disks over their
eyes and securing the disks with several layers of electrician's tape

17
wrapped around the head. A stretch sock, with an opening for the
nostrils, was then placed over the head and taped at the neck to prevent
the lizard from scratching off the tape.
A photothigmotron was built to quantify thermoregulatory postural
changes in Cyclura. A square plywood substrate box, measuring 65 by 65
by 5 cm, was lined with Plexiglas and sealed internally with fiberglass
cloth and resin. A 0.3 mm thick aluminum sheet formed the top of the
inner Plexiglas box and was affixed to it with Al-40 adhesive (Devcon).
A grid of 1 cm squares was scored on the exposed surface of the
aluminum. The substrate box rested on a 15 gallon water bath
(Precision Scientific Co.). The water bath pump was connected via
Tygon tubing (Norton Plastics and Synthetics Division) to two ports
drilled in the side of the substrate box. Water at the desired tempera¬
ture was pumped through the entry port in the substrate box and circulated
evenly via six inner exit ports pointed in different directions. A small
hole was made at one corner of the aluminum top to permit pressure
equalization when the pump was initially turned on. A short tube was
attached externally to the rim of the hole to prevent water overflow.
To confine the lizard to the aluminum substrate, a one meter high hard¬
ware cloth cage, reinforced with wooden frames at top and bottom, was
placed on the substrate box. The cage top was covered with a Plexiglas
sheet, on which was placed a 35 mm camera directed downward toward the
aluminum grid. A single 250 w heat lamp, attached to a tripod resting
on the floor, was directed slightly downward toward the side of the cage.
The observer's side of the cage was covered with vinyl plastic provided
with a small observation port. The photothigmotron was placed inside
the Environ-room.

18
Before each iguana was introduced into the photothigmotron,
several body areas were determined. The lizard was anesthetized with
halothane and placed on a Plexiglas sheet. Outlines of the outer edge
of the body and the inner area of substrate contact were marked on the
sheet and later transferred to a piece of paper. Three separate
measurements of each area were made planimetrically and the mean taken.
Contact area of the anesthetized lizard was assumed to equal maximum
contact of the free-ranging lizard. The day before the first experiment,
the lizard was placed on an insulative piece of plywood resting on top
of the aluminum substrate of the photothigmotron. One experiment was
performed each day, starting at midday. The photoperiod was ten hours.
Animals were kept in the cage for three to five experiments, during
which they received neither food nor water. An hour before heating or
cooling of the lizard began, the water bath pump was turned on to bring
substrate temperature to the desired level. The plywood was then
gently removed, shifting the individual onto the metal substrate. The
heat lamp was also turned on in some experiments. Photographs of the
dorsal aspect of the lizard were taken at intervals. Core body tempera¬
ture was measured with a near-field transmitter placed in the digestive
tract. The extent of leg and tail contact was noted, as well as how
much of the anterior trunk was elevated off the substrate. The black
and white photograph negatives were projected with a slide projector
onto a piece of white paper and the body outline traced. The dorsal
outline area was determined planimetrically. Head, anterior trunk, leg
and tail noncontact areas as well as a standard correction based on the
difference between the inner and outer outlines of the anesthetized

19
lizard were subtracted from the dorsal outline area to get the approxi¬
mate contact area. This value could then be converted to a percentage
of the maximum possible contact.
Head temperature was measured at two different locations, using
probed near-field transmitters. The transmitter was taped to a piece
of television antenna cable fastened with a rivet around the lizard's
neck. In a large male monitored in the photothigmotron, the thermistor
probe was inserted into the orbital venous blood sinus through a 1 mm
diameter hole drilled in the skull. The probe wire was glued to the
center of a circular cork disk, which was glued to the top of the head
with epoxy resin. Initial placement of the probe resulted in a temporary
eye bulging (also noted during cannula placement in Phi’ynosoma eomutwn
by Heath 1966). In a second large male, monitored in the arena atop the
zoology building, the probe tip was placed just inside a slit made in
the tympanic membrane. The probe wire was glued to a plastic disk, which
was glued to the outside of the tympanum. Lizards were allowed to
recover from implantation for three days before testing.
Skin reflectivity was determined for two Cyclura adults from Water
Cay, one in the light and one in the dark color phase. The light phase
lizard was killed by heating it above the lethal maximum body tempera¬
ture and the dark phase lizard died of starvation. Epidermal-dermal
squares, measuring approximately 25 mm on a side, were removed from the
lizards immediately after death and placed on water-moistened filter paper.
Patches were taken from the mid-dorsal, ventral pectoral, and ventral
abdominal trunk and the dorsal and ventral tail just distal to the anus.
Integumental reflectivities were immediately determined, using a Bausch

20
and Lomb Spectronic 20 Spectrophotometer with reflectometer attachment.
Monochromatic reflectance was obtained at a 90° angle to the skin surface,
using a magnesium carbonate block as a standard. Readings were taken
between 340 and 960 nm at 20 nm intervals. Reflectivity over the same
range was also determined for a common substrate on Pine and Water Cays,
fused corralline sand covered with two species of blue-green algae.
To check for possible reflectance of near-infrared light (700 to
900 nm) from the integument of live Cyotura, two adults were photographed
during daily activity in Gainesville, using Kodak Infrared Ektachrome
film (Schwalm et al. 1977). When the camera lens is covered with a
yellow filter, reflected near infrared light appears red in the developed
film.
Skin patches were heated from different angles to determine whether
the rate of integumental heating decreased as the radiant heat shadow
caused by scale imbrication increased. The patches were removed from
various locations on a single adult iguana, tacked to separate pieces
of plywood, dried, and cooled to 20 C in a controlled environmental
room. A YSI Probe 421 was placed under the patch to monitor the time
required to heat the dried skin to 50 C with a 250 w heat lamp. Each
patch was heated 11 times, with the lamp at a constant distance of 17 cm,
oriented normally to the skin and at 15, 30, 45, 60, and 75° from normal,
both directly anterior and posterior to the imbricate edges of the
scale rows.
The histological structure of integumental "sensory spots"
(Miller and Kasahara 1967) were examined in tissue sections prepared for
light microscopy. Material had been fixed in formalin and transferred

21
to isopropyl alcohol for preservation at the Florida State Museum.
Tissue blocks were taken from the lizard's rostral, labial, parietal
eye, and dorsal tail regions, dehydrated in ethyl alcohol, embedded in
methacrylate plastic, and sectioned at 5 micrometers thickness vertical
to the skin surface. Sections were mounted serially and stained with
Mayer's hematoxylin and eosin-phloxine.
Six adult iguanas were heated and cooled in a controlled environ¬
mental room (Environette, Lab-Line Instruments, Inc.). Each individual
was habituated for one day to its small hardware cloth cage, placed in
one Environette set at an air temperature of 20 C. Polyurethane panels
enclosed three sides of the cage, reducing the air flow over the lizard's
body surface to practically zero. The cage floor was also composed of
polyurethane. On the second day a YSI rectal probe, coated with petroleum
jelly, was inserted into the lizard's large intestine to a depth of 7-10
cm and taped to the tail. The cage was then rapidly moved to another
similarly wind-baffled box in another Environette set at an air tempera¬
ture of 40 C. When within 1-2 C of ambient air temperature, the lizard
was rapidly transferred back to the 20 C Environette and cooled. During
the third and final day in the chamber, the individual was secured in a
supine posture to a wooden jig. The lizard's legs were free to move.
A rectal probe was again inserted into the intestine and a banjo probe
glued middorsally to measure body surface temperature. The individual
was then heated and cooled again. Animals were observed continuously
during heating and cooling through an observation port in the chamber
door. Breathing rates were recorded intermittently. The photoperiod
was ten hours. Heating began at 1300 and cooling was always over by

22
2100. Lizards were weighed at the start and the end of the test period
and given neither food nor water.
Heart rate and oxygen consumption at rest, during maximum activity,
and during recovery from exhaustive exercise were determined for five
adult iguanas (two females and three males; X = 1252 g, range = 834-1722).
Each lizard was tested on six consecutive days, starting at an air
temperature of 15 C and increasing at 5 C increments to a maximum of
40 C. Experiments were conducted after 2100 on individuals which had
received neither food nor water for two days prior to the first run.
After each run the chamber air temperature was elevated five degrees
for the next night's run. When not being tested, animals were permitted
to move freely about the 1.4 by 2.5 m Environ-Room, kept on a 10L:14D
cycle.
The lizard's exhaled air was trapped in a rectangular, black-
painted Plexiglas mask fitted over the head. The mask had a neoprene
apron glued to its open posterior end, which fitted over the lizard's
anterior trunk and forelegs. Room air was sucked into the open posterior
end of the apron, past the lizard's head, and out the front of the mask
via Tvgon tubing through columns of soda lime and Drierite, to remove
carbon dioxide and water vapor, respectively. Flow rates ranged from
280 to 2440 cc/min. The gas then passed through a flowmeter, a pump,
another flowmeter, and finally through a Beckman C-2 paramagnetic oxygen
analyzer, located outside the Environ-Room. Gas flows were measured
with rotameters calibrated with a Brooks Vol-U-Meter. The decrease in
percent oxygen resulting from the lizard's respiration was recorded
continuously, except during the exercise period, with a Honeywell
Electronic 15 stripchart recorder. Lizards were kept in total darkness

23
during all oxygen consumption measurements. Oxygen consumption was
calculated from oxygen content and flow rate data using the appropriate
equation given by Depocas and Hart (1957).
For each run, a rectal YSI probe was inserted into the lizard's
intestine. The animal was then fastened in a supine posture to a wooden
jig, with its head resting inside the mask, taped to the jig. Oxygen
consumption was monitored until a stable resting rate was achieved.
The lizard was then removed from the jig and mask and exercised by hand
manipulation for several minutes, sufficient to completely exhaust the
animal. The lizard was immediately refitted into the mask and refastened
to the jig. ECG needle leads were pushed beneath the skin of the shoulder
and the base of the tail. Heart rate was measured every 4-10 minutes of
the recovery period with a Physiograph Four A with high gain preamplifier
(E and M Instrument Co.), located outside the Environ-Room. When
oxygen consumption again fell to the resting rate, the lizard was
manipulated by hand for another two minutes to get a maximum heart rate
reading.

ECOLOGICAL ASPECTS OF THE THERMAL BIOLOGY
OF CYCLURA CARINATA
Climate of Water Cay
The Caicos Islands are in the northern tropical zone, slightly
below the Tropic of Cancer. They are also in the tropics under the
Koppen System (Koppen 1923), since mean monthly shaded air temperature
never falls below 18 C. Water Cay has a trade wind littoral climate,
with fairly uniform high temperature and distinct wet and dry seasons.
Figure 1 summarizes the black body temperature measurements taken
during three trips to Water Cay. Cyclura had to remain intermittently
in the shade on sunny days during a large portion of its activity period
or face lethal body temperatures due to radiant heat input. Adult
lizards provided with some well timed overcast periods could theoret¬
ically survive in an exposed location all day in at least October to
February. Yearly photoperiod varied by almost three hours, reflected
in the difference in width of the heating curves for February vs. July.
Figure 2 illustrates the annual air temperature cycle near the
two primary locations where Cyclura was observed. The range of mean
shaded air temperature was only 4.5 C on Grand Turk. Of course, both
daily and seasonal microclimatic extremes at ground level were greater.
Gainesville, located in the subtropics, was too cold part of the year
to support a Cyclura population. From the beginning of April until
the end of September, penned Gainesville lizards ate readily and gained
weight. Feeding was negligible during the other six months of the year.
24

25
on —i ‘ ‘ ‘ -
7 8 9 10 II 12 13 14 15 16 17 18
HOUR INTERVAL
Figure 1. Black body temperatures in the field. February, June, and
October measurements are represented by solid, dashed, and dotted
lines, respectively. Maximum temperatures during the recording
periods are represented by open squares, mean temperatures for the
hourly intervals by open circles, and minimum temperatures by
solid circles. All weather conditions encountered were included
in the calculations. Horizontal lines are means and vertical lines
are ranges of regulating phase s of lizards at Site II for
February, July, and October, respectively.

AIR TEMPERATURE (°C)
26
Figure 2. Air temperature in the shade at Gainesville, Florida (Agronomy
Farm Weather Station; 29° 40' N, 82° 20' W; Prine 1977; and
Grand Turk, British West Indies (Auxiliary Air Field; 21° 26' N,
71° 8' W). Vertical lines extend from the maximum to the
minimum recorded Ta each month (dashed line for the year 1976
at Gainesville, solid line for the years 1900-1968 at Grand Turk).
Horizontal lines are mean monthly temperatures on Grand Turk.
Circles and squares represent mean maximum and mean minimum air
temperatures, respectively. Air temperatures were measured in
a standard NOAA shelter (Stevenson screen) at a height of 1.5 m.

27
Special precautions had to be taken to prevent the lizards from freez¬
ing to death between November and February.
An easterly wind blew almost continually over the islands. The
mean surface wind speed on Grand Turk was 24.6 km per hour (15.3 mph).
Figure 3 illustrates the mean maximum July wind speeds on a road pro¬
tected on the windward side by a limestone ridge and dense scrub at
Site II. Wind speeds were greatest during the middle of the day, a
typical pattern (Geiger 1957). Wind speed was greater at 50 cm than
at 3 cm above the ground, also a typical pattern. Thus, lizards basking
or feeding above the ground were usually cooled at a greater rate by
forced convection than lizards on the ground. Wind speed was consider¬
ably higher facing the inlet between Water and Pine Cays just a few
meters away. Since lizard burrows were common near the top of the
ridge, some interesting wind related behavior was expected there,
perhaps similar to the seeking out of minimal wind exposure at low
Tfo and maximum exposure at high Tj, as seen in Amblyrhynohus ovistatus
of the Galapagos Islands (Bartholomew 1966, Bartholomew et al. 1976).
Rainfall was distinctly seasonal in the Caicos (Figure 4). Mean
annual rainfall on Pine and Water Cays was probably slightly greater
than on Grand Turk (597 mm or 23.5 inches). Four seasons characterize
the western islands of the Caicos; a cool dry winter from January to
March, a warm dry spring from March to May, a hot, increasingly wet
summer from June to October, and a warm, wet autumn from October to
December.
Water Cay was fairly humid (Table 1). Daily relative humidity
decreased as air temperature increased, minimizing near midday. Of

MEAN MAXIMUM WIND SPEED (M/SEC)
28
Figure 3. Mean maximum wind speed at Site II in July. Ten consecutive
wind speed maxima were measured each hour for seven days at
3 cm (open rectangles) and 50 cm (solid rectangles) above the
ground. Horizontal lines are means, vertical lines are ranges,
and rectangles are plus and minus two SE of the mean.

MONTHLY RAINFALL (MM)
29
190
Figure 4. Monthly precipitation on Grand Turk, British West Indies. The
line represents the mean monthly rainfall for the years 1900-1968.
Total rainfall per month for several months on Pine Cay, Caicos
Islands, are represented by dots.

30
Table 1. Daily variation in relative humidity. Measurements
taken during hours of precipitation were not included.
Data were collected at Site I (Jun) and Site II (Feb,
Jul, Oct) on Water Cay.
MEAN HOURLY
RELATIVE
HUMIDITY
HOUR INTERVAL
FEB
JUN
JUT.
OCT
N=4
N=12
N=15
N=7
DAYS
0700-0759
-
75
74
86
0800-0859
78
74
71
77
0900-0959
67
69
64
69
1000-1059
66
62
64
64
1100-1159
66
59
61
62
1200-1259
60
59
64
62
1300-1359
61
64
63
59
1400-1459
62
61
64
64
1500-1559
66
61
65
66
1600-1659
70
67
71
70
1700-1759
73
72
71
78
MINIMUM DAILY READING
56
46
52
62
MAXIMUM DAILY READING
79
80
80
87
RANGE
23
34
28
25
MEAN MONTHLY RH
66.
0
64.6
66.9
68.
0
2SE
2.
1
1.4
1.5
2.
4

31
course, daily absolute humidity did not vary as much as relative
humidity. Mean monthly relative humidity did not differ significantly
between February, June, July, and October.
Piel Body Temperature Cycle of a Heliothermic Lizard
The body temperature cycle was divided into four phases by Wilson
and Lee (1974). Some slight alterations were made for the following
scheme (Figure 5).
1. Morning Heating Phase The lizard emerged from its burrow and
heated rapidly in direct sunlight. Cyelura usually started its first
feeding foray below the first maximum Ty, upon reaching the "normal
activity range" (Cowles and Bogert 1944). Wilson and Lee's "basking
phase" ends here. This term is confusing since basking did not cease
when the first feeding foray began. Also, Cyotura was so wary during
the June-July trip that it was very difficult to observe the start of
the first foray unless I was sitting in a well positioned blind. There¬
fore, the first phase has been called the morning heating phase, ending
at the first maximum TThis endpoint is obvious on a continuous T^
record.
2. Regulating Phase fluctuated between a series of maxima and
minima, staying within a fairly narrow range.
3. Burrow Cooling Phase It started at the lizard’s evening
submergence and ended during the night, when had fallen to the 24
hour minimum. The decline in Tfollowed a Newtonian cooling curve,
with an ever decreasing rate of T^ decline as the stable deep burrow
temperature was approached.

BODY TEMPERATURE
9 10 II 12 13 14 15 16 17
TIME
Figure 5. Body temperature record of Lizard 2 on February 1Ü, 1976. The Arabic numerals at the
top of the graph refer to the four phases of the cycle. The broken line at the
bottom indicates the periods when the sun was obscured by clouds. Capital letters
refer to data points discussed in the text.
Co
NJ

33
4. Stable Tfo Phase The lowest T^ in the 24 hour cycle occurred
just before emergence in two lizards larger than Cyclura carlnata, the
Lace Monitor, Vai^anus varius (Stebbins and Barwick 1968) and the Komodo
Dragon, Varanus komodoensis (McNab and Auffenberg 1976). However,
stabilized considerably before emergence in Cyclura, necessitating a
fourth descriptive phase. The term "inactive phase" used by Wilson and
Lee would be confusing since Cyclura was also inactive in the burrow
cooling phase.
A continuous body temperature record provides a substantial amount
of information about a lizard. It was sometimes difficult to determine
the exact time of emergence from the record, since deep burrow tempera¬
ture was slightly greater than surface temperature early in the morning.
The lizard in Figure 5 may have lost heat on the surface just after
emergence (A) or near the burrow mouth prior to emergence (B). Slight
differences in emergence T^ could sometimes be used to determine the
burrow the lizard spent the night in, without having to locate the
burrow the preceding night by following the increasing signal strength
with the receiver. The time of submergence (F) is easily discerned in
the February record. However, determination of submergence time from
the summer records was commonly more difficult, since often fell
prior to submergence and at a rate not easily distinguished from the
initial decline after submergence. Body temperature shifted abruptly
upward as the overcast cleared temporarily at (D), a change maintained
until submergence. Shifts of this sort were fairly common in the records.
Several further terms concerning lizard T-^ need to be defined.
All refer to measurements on a single individual taken during its

34
regulating phase. Unless otherwise specified, each T^ reading used in
a calculation occurred during a period of sunshine or overcast (10
minutes or less) which did not cause a decline in T^.
1. Mean Daily Ty Once the continuous record for a single day
was plotted, the first and last maxima plus points at fifteen minute
intervals, on the hour, were averaged.
2. Daily Minimum, Maximum Ty The lowest, highest T^ recorded
for the day (C and E, respectively, in Figure 5).
3. Mean Daily Minimum, Maximum Tfa An average of the daily
minimum, maximum for a specified number of days.
4. Mean of Daily Ty Minima, Maxima An average of all minima,
maxima for one day.
5. Mean of Ty Minima, Maxima An average of all minima, maxima
for a specified number of days.
Regulating Phase Body Temperature in the Field
Cyclura aarinata is a stenothermal lizard during the regulating
phase (Table 2). The mean of the regulating phase T\ ranges for all
the monitoring days during clear weather was only 3.3 C (SD = 1.3,
range = 0.7-6.4, N = 106 days). The frequency distribution of these
Tjj ranges was positively skewed toward greater stenothermality, with
a median of 2.9 C and a modal class of 2.0-2.4 C.
Adults rarely went into their burrows at Sites I and II because
of extremely high surface temperatures, although this may have been a
common behavior in more sparsely vegetated areas. Free-ranging lizards
prematurely submerged for the day on only 19 occasions during 129
individual monitoring days: 45 percent due to rain, 40 percent due to

Table 2. Summary of regulating phase body temperatures of lizards in the field. Data are
arranged by season, field site, sex, increasing body weight, and individually by
decreasing daily mean Tfo, each line equaling one day's monitoring. Mean Tfo's were
calculated from the entire record, irrespective of weather conditions or lizard
location. The letters (B) and (SU) signify that the lizard was in a burrow or the
sun was obscured by clouds (sun under) when the minimum Twas recorded, respectively.
Lizards 2 and 3 were monitored in February and October.
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
DAILY
MAXIMUM
Tb
RANGE
LENGTH OF
REGULATING
PHASE (MIN)
% TIME
SUN
BEHIND
CLOUDS
% TIME
LIZARD IN
BURROW
1(F)
F
1135
II
2-19-76
37.7
0.82
36.2
39.6
3.4
293
4
0
17
37.5
1.85
33.4(B)
39.8
6.4
280
15
36
18
36.8
1.38
34.0(B)
38.8
4.8
240
30
19
2(F)
M
1665
II
2-05-76
40.8
0.46
40.2
41.5
1.3
142
10
0
6
38.6
0.72
36.0
39.2
3.2
288
14
0
7
38.4
0.68
37.1
40.0
2.9
264
8
0
12
38.3
0.52
37.0
39.2
2.2
315
0
0
11
38.3
0.27
37.5
38.6
1.1
187
28
0
9
38.0
1.40
34.7(SU)
40.0
5.3
332
8
0
8
37.7
0.94
35.4
39.0
3.6
333
0
0
10
37.6
0.79
35.9
38.8
2.9
244
41
0
3(F)
M
1775
II
2-12-76
40.0
0.60
38.1
41.0
2.9
427
0
0
13
38.5
0.53
37.1
39.4
2.3
363
40
0
11
38.2
0.50
37.0
39.2
2.2
288
20
0
14
38.2
0.63
36.6
39.5
2.9
426
8
0
15
38.1
0.61
37.0
39.2
2.2
327
19
0
19
37.8
0.48
36.4
38.6
2.2
330
3
0
17
37.4
0.79
35.6
38.5
2.9
505
38
0
18
37.4
0.69
35.8
38.7
2.9
360
43
0
20
36.0
1.81
32.6(SU)
38.6
6.0
313
63
0
Cn

Table 2. Continued.
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
4(C)
F
645
I
6-10-74
40.8
2.03
37.6
9
40.4
1.14
38.2
8
40.3
1.63
37.4
11
39.9
3.00
35.8
7
39.1
0.93
37.1
5(C)
F
723
I
6-24-74
41.9
0.81
40.4
25
41.5
0.42
40.7
22
40.6
0.95
38.5
26
40.6
0.58
39.3
27
40.0
0.76
38.2
28
39.3
0.96
37.2
29
39.3
0.68
38.0
7-01
35.3
3.79
28.6(B)
6(C)
F
815
I
6-17-74*
39.0
0.48
37.8
9
38.8
1.29
37.2
8
38.3
0.40
37.4
18
38.3
4.18
33.9(SU)
10
38.3
1.10
36.4
7
37.6
1.29
35.7
14
37.0
1.37
34.8
11
36.5
1.36
34.9
7(C)
M
1290
I
6-24-74
40.7
0.63
39.4
22
40.3
0.63
38.0
29
38.7
0.96
36.3
25
38.1
1.24
35.3
26
38.1
1.01
35.7
DAILY LENGTH OF
MAXIMUM REGULATING
Tb RANGE PHASE (MIN)
% TIME
SUN % TIME
BEHIND LIZARD IN
CLOUDS BURROW
43.8
6.2
419
30
41.8
3.6
260
23
42.6
5.2
288
7
42.0
6.2
312
24
41.0
3.9
360
0
43.7
3.3
325
0
42.5
1.8
501
0
41.5
3.0
272
29
41.7
2.4
481
3
41.4
3.2
507
0
41.4
4.2
525
10
40.2
2.2
459
0
39.8
11.2
481
72
40.0
2.2
315
50
40.8
3.6
291
29
39.4
2.0
268
11
41.3
7.4
242
42
39.8
3.4
430
29
39.8
4.1
407
2
38.6
3.8
326
31
37.8
2.9
315
24
41.9
2.5
325
0
41.8
3.8
332
24
40.8
4.5
465
0
41.3
6.0
465
0
39.7
4.0
488
3
NOT
APPLICABLE
to

Table 2. Continued.
LIZARD
(FREE OR
CAGED) SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
7(Continued)
27
38.0
1.16
36.0
28
37.8
1.11
35.3
7-01
35.0
3.22
29.3(B)
8(F) F
675
I
6-26-74
41.9
0.44
40.7
25
40.9
0.87
39.3
14
40.6
0.97
37.7(SU)
16
39.8
0.66
38.5
24
39.8
0.46
38.4
22
39.6
0.60
37.3
17
39.6
1.38
36.7(SU)
23
39.2
0.71
37.5
18
38.4
1.93
30.3(SU)
12
35.2
3.04
30.4(B)
15
33.8
4.45
29.7(B)
9(F) M
1450
I
6-29-74
42.4
0.37
41.6
28
42.4
0.46
41.6
17
42.3
0.39
41.5
27
42.2
0.66
40.9
24
41.7
0.42
40.8
16
41.6
0.70
40.5
23
41.0
0.48
39.9
25
40.1
0.57
39.4
7-02
40.0
1.05
38.1
6-26
39.9
1.08
38.0
7-01
39.2
0.51
38.1
4
37.2
3.82
29.8(B)
6-15
35.0
4.31
28.9(SU)
DAILY LENGTH OF
MAXIMUM REGULATING
Tfo RANGE PHASE (MIN)
% TIME
SUN / TIME
BEHIND LIZARD IN
CLOUDS BURROW
40.1
4.1
507
0
NOT
39.9
4.6
417
12
APPLICABLE
39.7
10.4
507
69
II
42.9
2.2
420
0
0
42.1
2.8
493
0
0
42.1
4.4
321
21
0
42.3
3.8
457
0
0
40.6
2.2
360
7
0
40.3
3.0
378
22
0
42.1
5.4
393
51
0
40.2
2.7
339
10
0
40.5
10.2
300
34
0
40.9
10.5
429
0
43
42.1
12.4
330
45
170
43.3
1.7
490
0
0
43.5
1.9
525
0
0
43.0
1.5
390
54
0
43.2
2.3
486
0
0
42.6
1.8
378
7
0
43.3
2.8
457
0
0
41.6
1.7
351
10
0
41.2
1.8
450
0
0
42.5
4.4
510
14
0
41.4
3.4
480
3
0
39.9
1.8
267
47
0
40.9
11.1
455
26
17
41.0
12.1
358
35
0
U>
-a

Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(C)
SITE
DATE
MEAN
*b
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
10(F)
M
787
II
7-18-74
40.8
0.85
38.8
13
39.9
1.79
36.9
15
39.5
0.58
38.2
12
38.6
1.07
37.1
16
38.5
1.07
35.2(B)
20
38.4
1.43
32.1(B)
22
38.4
1.39
35.9
21
38.1
1.21
35.3
19
37.4
2.39
31.9(E)
23
37.1
1.30
34.9(SU)
11
36.9
1.60
33.3(B)
17
34.4
3.30
30.1(SU)
11(F)
M
1598
II
7-09-74
41.4
0.87
39.0(SU)
10
41.3
0.33
40.3
11
40.8
0.36
40.1
12
40.1
0.59
38.8
18
38.9
0.66
37.4
13
38.8
0.78
37.1
6
38.5
0.25
38.2
22
38.2
1.11
36.6
15
38.1
0.66
33.7(SU)
19
38.0
0.86
36.5
20
38.0
0.64
36.9
21
37.6
0.88
35.7
23
37.1
1.49
33.8(SU)
25
36.7
1.27
35.1
7
36.2
1.11
34.1(SU)
17
34.2
3.31
29.5(SU)
16
28.0
0.10
28.0(B)
DAILY LENGTH OF
MAXIMUM REGULATING
Tfc RANGE PHASE (MIN)
% TIME
SUN / TIME
BEHIND LIZARD IN
CLOUDS BURROW
42.5
3.7
408
10
0
42.9
6.0
500
0
0
40.6
2.4
375
6
0
40.8
3.7
486
6
0
40.6
5.4
384
2
9
40.2
8.1
450
19
8
40.7
4.8
487
12
0
40.2
4.9
435
5
0
40.6
8.7
431
0
20
39.5
4.6
318
16
0
39.9
6.6
460
0
31
38.9
8.8
405
62
0
42.9
3.9
396
35
0
42.1
1.8
420
30
0
41.5
1.4
481
0
0
41.1
2.3
509
6
0
40.1
2.7
480
8 •
0
40.4
3.3
339
0
0
38.9
0.7
178
20
0
40.3
3.7
574
14
0
39.6
5.9
550
27
0
39.6
3.1
489
42
0
39.3
2.4
470
24
0
39.8
4.1
517
22
0
39.6
5.8
575
26
0
39.7
4.6
285
5
0
38.4
4.3
234
100
0
39.8
10.3
381
65
0
28.3
0.3
0
2
100
U)
CO

Table 2. Concinued
LIZARD
(FREE OR
CAGED)
SEX
WEIGHT
(G)
SITE
DATE
MEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
12(F)
M
1420
I
8-28-76
39.7
0.28
39.4
30
39.5
0.57
37.9
29
36.3
2.31
32.2(B)
13(F)
M
950
III
9-03-76
39.3
0.69
38.2
5
38.7
0.59
37.5
4
37.8
1.14
35.2
O
L
36.8
2.63
31.8(SU)
14(F)
M
1864
II
10-25-75
39.9
0.84
38.2
21
39.7
0.36
38.5
23
39.3
0.53
38.2
15(F)
M
1784
II
10-29-75
39.2
0.85
37.5
28
39.2
0.82
37.3
27
38.3
0.81
37.6
3(F)
M
1672
II
10-06-75
40.1
0.65
37.9
5
38.6
3.30
31.0(B)
7
38.2
1.44
35.0
8
38.1
1.45
34.6(B)
16
37.8
1.09
35.9
21
37.7
0.95
36.2
4
37.6
0.89
35.1
20
37.5
1.29
35.2
18
37.5
0.82
35.6
19
37.3
0.76
34.5
15
34.8
2.82
30.6(B)
DAILY LENGTH OF
MAXIMUM REGULATING
Tjy RANGE PHASE (MIN)
% TIME
SUN % TIME
BEHIND LIZARD IN
CLOUDS BURROW
40.2
0.8
114
0
0
40.2
2.3
327
27
0
39.9
7.7
387
38
25
40.4
2.2
195
35
0
39.6
2.1
315
3
0
39.4
4.2
213
62
0
39.3
7.5
417
45
0
40.8
2.6
186
24
0
40.7
2.2
300
18
0
40.2
2.0
357
6
0
41.0
3.5
281
0
0
40.7
3.4
313
31
0
40.4
2.9
330
30
0
41.5
3.6
445
5
0
42.3
11.3
487
10
20
40.6
5.6
345
30
0
40.4
5.8
380
0
12
39.4
3.5
159
19
0
39.7
3.5
255
13
0
39.4
4.2
312
73
0
39.7
4.5
270
22
0
39.0
3.4
310
59
0
38.3
3.8
354
5
0
39.2
8.6
285
21
49
to

Table 2. Continued
LIZARD
(FREE OR
CAGED)
SEX
WEICHT
(G)
SITE
DATE
KEAN
Tb
S.D.
OF
MEAN
DAILY
MINIMUM
Tb
DAILY
MAXIMUM
Tb
RANGE
LENGTH OF
REGULATING
PHASE (MIN)
% TIME
SUN
BEHIND
CLOUDS
% TIME
LIZARD IN
BURROW
2(F)
M
1507
II
10-16-75
40.1
0.83
39.3
41.4
2.1
90
33
0
19
39.5
0.37
38.5
40.1
1.6
308
6
0
18
38.6
0.76
37.2
40.2
3.0
360
51
0
Laid eggs in the cage during this day.
•t-
o

41
human disturbances, and 15 percent due to extreme cloudiness. Rates of
cooling during burrow excavation and shade sitting were not consistently
different. Thus, the cause of the cooling could not be determined by
examination of the T^ record alone.
Figure 6 summarizes the regulating phase data for all the field
lizards. The rise in mean T^ over the 3.5 C range was steady. Minimum
T^'s fell into two groups having no correlation with increasing mean
2%/s, one group above and one below 39 C mean This probably is of
no significance. The correlation of maximum T^ with mean T- was better.
The lowest maximum T^ recorded was 1.6 C above the highest minimum. Thus,
all ranges of individuals overlapped somewhat, no matter what the
sex, habitat type, or season of measurement.
The sample sizes were too small to determine conclusively whether
or not mean T7 differed significantly with sex. Lizard 9, a dominant
male, had a significantly higher T^ than Lizard 8 (p <_ 0.05), an adult
female monitored simultaneously. However, two of three caged females
in the simple habitat, Lizards 4 and 5, had significantly higher body
temperatures than the caged male, Lizard 7. Mean 's were not
significantly different in two free-ranging males and a female monitored
at Site II in February.
Gravid females may have lower regulating phase 2^'s than spent
females and males. The gravid June female (Lizard 6) had the lowest
mean of all those monitored. The means were significantly higher
(p _< 0.05) for the other two caged females (Lizards 4 and 5) and the
free-ranging female (Lizard 8) at Site I, all of which had probably
recently laid eggs, since they actively defended their burrows against
both encroaching males and females. However, Lizard 6's T^
was not

Figure 6. Summary of field body temperatures during the regulating phase. The lizards were
monitored from 3 to 16 days. Mean regulating phase T¿ was calculated from values
taken at 15 minute intervals from the monitoring record. The number of days used
to calculate each mean was variable and usually equal to the total monitoring time
(Table 2). Periods of hyperthermia were included in calculation of the means,
whereas periods of declining Tfo due to overcast or burrow submergence were not.
Vertical lines are ranges, horizontal lines are means, and rectangles are 99 per¬
cent confidence limits of the mean. Sex is indicated by an M or F. The four
caged lizards in the complex habitat at Site I in June are represented by
rectangles filled with vertical lines; the same four lizards in the simple habitat
by dots; three free-ranging lizards in February at Site II by horizontal lines;
two lizards in June at Site I by diagonal lines sloping downward to the right; two
lizards in July at Site II by open circles; a single lizard in August at Site I by
open squares; a single lizard in September on Fort George Island by an open rectangle;
four lizards in October at Site II by diagonal lines sloping downward to the left.

44
43
42
4í
40
39
38
37
36
35
34
33
M
^ L 1 1 1 L 1 1 - 1 -■ ! 1 1 ■■ ■ j l—
6 ! 3672 13 10 7 II 3 i! !2 5 14 2484 5 9 £
INDIVIDUAL IDENTIFICATION NUMBER

44
significantly lower than the caged male, Lizard 7. On the single day
Lizard 6's Tu was recorded after egg laying, the maximum was 0.5 C
higher than the highest maximum of the seven monitoring days previous
to laying. The highest mean T^ recorded was on the day she laid her
eggs, but the mean was not significantly higher than the next two
highest means recorded prior to laying. The differences probably
would have been significant had not the day of laying been overcast
50 percent of the time.
Temporary Hyperthermia and Hypothermia
All eleven iguanas monitored in the field passed through a period
of hyperthermia soon after release (Figure 7 and Table 3) . Tâ– < then
declined to a more stable, presumably normal range. The T^ decline
was either rather rapid as for Lizard 3 or more gradual as for Lizards
8 and 11. The extent of the hyperthermic response, as measured by tne
sum of the ranges of the maximum daily T^ and the mean daily T^'s
between the hyperthermic peak day and the return to normalcy, was
usually slightly greater in coelomically than enterically implanted
lizards. However, the response may vary with season as well as with
transmitter location. The summer animals were probably suffering
from a bacterial infection resulting from transmitter implantations
(see Vaughn et at. 1974, Kluger et al. 1975, and Bemheim and Kluger
1976a, b for related laboratory studies). The second Th rise observed
in Lizards 8 and 9 may have been due to reinfection of the wound or
a secondary response to a chronic infection. In the latter case it
may be that lizards could not tolerate a high T^ for more than a few
days even though infectious agents were still present (Cowles and

Figure 7. Lizard hyperthermia and hypothermia during the regulating phase. Lines
connect T£, maxima, omitting those during periods of heavy overcast (open
circles). Open squares are daily means. Lizard 11 was prevented
from emerging on Day 3 by a land crab (Cardisoma guanahumi) ensconced
for a single day just inside the burrow mouth. The transmitter was in
the gut of Lizard 3 and in the coeloms of Lizards 8 and 11.


Table. 3. Hyperthermia in Cyatura oavinata in the field. The ease of graphically distin¬
guishing a Tfo peak and a recovery from hyperthermia was classified as high, medium,
or low. The maximum Tfo was the highest body temperature recorded for a lizard.
The low maximum TB was the lowest maximum recorded on the day recovery was com¬
plete or before a second rise in occurred (Lizards 8 and 9).
LIZARD
2
3
4
5
6
7
8
9
10
11
3
MEAN S.D.
MONTH
FEB
FEB
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JULY
JULY
OCT
LOCATION OF
TRANSMITTER
GUT
GUT
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
COELOM
GUT
CAGED OR FREE
RANGING LIZARD
F
F
C
C
C
C
F
F
F
F
F
DAYS LIZARD MONITORED
8
10
6
11
13
11
15
12
13
20
19
CLARITY OF TB TREND
MEDIUM
MEDIUM
HIGH
HIGH
LOW
HIGH
HIGH
HIGH
HIGH
HIGH
MEDIUM
MAXIMUM Tb
41.5
41.0
43.8
43.7
39.9
41.9
42.1
42.9
42.6
43.5
42.9
42.9
42.3
42.3 1.2
DAY OF MAXIMUM TB
1
2
5
4
1
4
3/15
2/6
3
4
3
HOURS AFTER RELEASE
TB MAXIMUM RECORDED
4
28
100
75
4
72
49/339
25/125
55
73
48
48 31
LOW DAILY MAXIMUM TB
37.6
37.4
—
38.7
37.5
36.3
39.4
38.7
35.9
37.8
37.6
37.7 1.1
DAY OF LOW DAILY
MAXIMUM Tb
5
8
9
6
8
11
4
13
10
6
TIME OF Tb MAX - TIME
OF LOW TB MAX (HOURS)
91
145
117
117
91
216/73
48/50
235
140
72
116 60
RANGE OF DAILY
MAXIMUM Tb'S
3.9
3.6
5.0
2.4
5.6
2.7/3.5
3.9/4.8
7.0
5.1
4.7
4.4 1.3

Table 3. Continued.
LIZARD 23456
MEAN Tp ON DAY OF
MAXIMUM Tp
40.8
40.0
40.8
41.9
38.8
MEAN Tp ON DAY OF
LOW DAILY MAXIMUM Tp
38.0
37.4
—
39.3
36.5
RANGE OF MEAN
2.8
2.6
—
2.6
2.3
7
8
9
10
11
3
40.7
40.6
41.9
41.7
42.4
39.9
41.4
38.6
38.0
39.2
39.9
37.1
38.1
38.1
2.7
1.4/2.7
1.8/2.5
2.8
3.3
0.5
MEAN S.D.
40.7 1.2
38.2 1.1
2.3 0.8
4>-
oo

49
and Burleson 1944, Licht 1965b, Licht and Basu 1967, Kluger et at.
1975). The body cavity internal to the wound sites and surrounding the
transmitter suppurated only slightly or net at all at the time trans¬
mitters were removed. The mean time between release and the maximum
hyperthermic response was 2.4 days in summer Cyclura and 1-2 days in
Dipsosaurus injected with live Aeromonas bacteria (Bernheim and Kluger
1976a). Recovery was complete in Dipsosaurus by the sixth day after
injection and, on the average, by the seventh day after the operation
in summer Cyclura. These similarities may signify that similar dis¬
ruptions were in force in the brain thermoregulatory centers of both
species. Hyperthermia in enterically implanted lizards may have been
a feeding response to a transmitter meal or, somehow, a reaction to
handling or the use of halothane. The lizard with the weakest response
was Number 6, the gravid caged female. Embryonic development in Cyclura
carinata begins within the female's body and continues after nesting
at burrow temperatures of 28-29 C (Iverson 1979). Regulating phase
T^'s in the range of nongravid lizards may be fatal to developing
embryos in Cyclura, as they are in Iguana -iguana (Licht and Moberly
1965). Thus, the hyperthermic response to bacterial infection may
have been counterbalanced somewhat by a hypothermic response to
gravidity.
Hyperthermia also occurred in the laboratory thermal gradient
(Figure 8). The high daily mean T^ was significantly greater (p _< 0.05)
than the low in six of eight animals. The hyperthermic responses in
the field and gradient were fairly similar; that is, a latent period
of variable duration was followed by a maximum Tj^ and a decline. High
mean T^ occurred in three species of Phrynosoma when first housed in

Figure 8. Patterns of regulating phase body temperature and
distance traveled in Cyclura oarínata in a laboratory
thermal gradient. Data are presented for eight
lizards, each monitored for four consecutive days.
Lizard 3 was included in both samples. Horizontal
lines are Tfo means, vertical lines are ranges, and
open rectangles are plus and minus two standard
errors of the mean. The total distance each lizard
walked (open circles) was measured during the full
eight hour photophase.

DAY IN GRADIE
BODY TEMPERATURE
DAILY DISTANCE TRAVELED (M)

52
terraria (Heath 1965) and in Anotis oavolinensis when first introduced
into a thermal gradient (Licht 1968). The authors believed the hyper¬
thermia, which ceased after two days, was due to lizard inexperience in
new surroundings. Learning the new thermal environment resulted in a
decline in T^ to the preferred level. The same explanation probably
applies to Cyolura. Four of eight lizards did not move voluntarily
during the first day. Two of the lizards were located under the heat
lamps when the photophase began. It was thought best to push them
toward the cool end of the gradient when they began to pant. The steady
increase in walking distance per day corresponded in part to learning
how to maintain preferred Tfo by shuttling.
The hyperthermic response, as measured by the difference between
the maximum mean daily T^ minus the mean preferred Tis similar in the
iguanid lizards studied to date: 1) 2.0 C in Anotas oavolinensis by the
third day in a thermal gradient (Licht 1968), 2) 2.3 C in field monitored
and 2.2 C in gradient monitored Cyolura oavinata, 3) 3.7 C in Dipsosauvus
dorsalis on the second day after injection of dead Aeromonas hydrophila
(Bernheim and Kluger 1976a), and 4) 2.9 C in Phrynosoma by the third
day in a thermal gradient (Heath 1965). One explanation for the
similarity in the responses is the constancy of the small temperature
gap between the preferred Tâ– , and the CTM in these lizards.
Voluntary hypothermia during the regulating phase was a rare event,
observed in only two free-ranging summer lizards (8 and 10) on the day of
their release. Lizard 8 never ventured further than a meter from its
burrow the first day, retreating six times to the burrow for lengthy
cooling periods (Figure 9). As T^ maxima increased, T^ minima increased
also, resulting in a series of ascending peaks ending with the highest

Figure 9. Hypothermia in a free-ranging lizard. Lizard 8 was in its burrow (solid
line) or lay partially exposed in the sandy depression in front of its
burrow mouth (dotted line) for a considerable part of its first day
after release.

BODY TEMPERATURE

55
peak late in the afternoon. The same pattern was observed in only 2 of
51 thermal gradient records. Hypothermia in newly released field
lizards may have been due to a temporarily heightened thermal sensitivity
due to cooling to 3-12 C during the implantation operation the previous
night.
The Relationship of Pertinent Environmental Variables
to Regulating Phase Body Temperatures
Season
Table 4 summarizes the environmental temperature data accumulated
for Site II. Mean black body temperature was not significantly different
in February and October. The mean range was only 3.1 C between February,
nearly the coldest month, and July, nearly the warmest. The mean
differences between months for the other three temperature measures
in the table were significant. The ranking was also consistent. The
February-July mean temperature difference increased from 4.2 C for sun
exposed sand and 4.7 C for shaded sand to 6.5 C for shaded air. Although
the monthly percentages of time that clouds obscured the sun did not
differ significantly, the thickness of the cloud cover and the percent¬
age of the sky in overcast were greater during the rainy season in
October than in February and July.
Mean monthly 2^ did not correlate directly with changing monthly
environmental temperature. Mean Tâ– , at Site II was significantly higher
in October than in February and July (p < 0.01, student's t test) and
mean T^rs in February and July did not differ significantly from each
other (Table 5) . The seasonal variation in mean Ti was probably only
slightly greater than 0.5 C (July minus February mean). The significant

56
Table 4. A seasonal comparison of some environmental variables
at Site II on Water Cay. The data were collected at
approximately hourly intervals from 0900 to 1700.
Values are for a variety of weather conditions in
February (15 days), July (16 days), and October (16
days).
N
X
X+2SE
X-2SE
HIGH
LOW
BLACK BODY TEMPERATURE
October
191
44.1
45.4
42.8
64.4
27.0
February
109
44.2
46.1
42.3
63.0
24.1
July
181
47.2
48.8
45.6
67.4
28.3
SUN EXPOSED SAND
TEMPERATURE
February
101
33.4
34.9
31.9
>52.0
23.7
October
190
35.8
36. 7
34.9
>52.0
25.1
July
177
37.6
38.5
36. 7
>52.0
26.8
SHADED SAND TEMPERATURE
February
109
26.3
26.9
25.7
36.3
21.4
October
192
29.1
29.5
28.7
39.6
25.1
July
182
31.0
31.3
30.7
35.5
26.5
SHADED AIR TEMPERATURE
(HEIGHT 6 CM)
February
109
26.3
26.7
25.9
32.6
22.1
October
193
30.5
30.9
30.1
38.1
24.2
July
181
32.8
33.1
32.5
37.2
27.8
PERCENT OF TIME SUN
OBSCURED BY CLOUDS
February
15
21.5
31.2
11.8
63
0
October
16
22.1
32.3
12.3
73
0
July
16
24.2
37.5
10.9
100
0
N
TOTAL (mm)
PRECIPITATION
February
15
5
July
16
36
October
16
83

57
Table 5. Seasonal changes in regulating phase body temperature in
the field and in the laboratory thermal gradient. Only
periods when the sun was not obscured by clouds were
included in the field calculations. Only third and fourth
day monitoring records were used in the calculation of mean
T-n's for gradient lizards. Values were taken from the T^
records at 15 minute intervals. All lizards were non-
hyperthermic.
MONTH
DAYS
FEMALES/
MALES
SITE
N
X Tb
X+2SE
X-2SE
SD
LOW
HIGH
FREE-1
RANGING LIZARDS
February
13
0/2
II
280
38.0
38.1
37.9
0.82
34.7
40.0
June
9
1/1
I
231
39.7
39.8
39.6
0.89
37.3
42.5
July
13
0/2
II
365
38.1
38.2
38.0
1.24
33.7
40,7
October
13
9/3
II
268
38.5
38.7
38.3
1.25
34.5
41.0
THERMAL
GRADIENT LI
ZARDS
July-Aug
8
2/2
222
36.9
37.1
36.7
1.86
31.6
41.8
December-Jan
8
2/2
174
37.1
37.3
36.9
1.08
34.0
39.3

58
difference between June and July T^ means was probably due to the
difference in the amount of vegetation at Sites I and II. The variance
of mean T^ in February was significantly lower (p <_ 0.01) than in July
and October. The winter variance was lower primarily in the upper
portion of the T^ range (Figure 10). Lizards had been acclimatized to
the more thermally variable Gainesville climate before testing in the
thermal gradient (Table 5). Yet, summer and winter sample means did
not differ significantly on any of the first four days in the gradient.
As in the Caicos, the variance of the mean T^ for winter animals was
significantly less than for summer animals (p < 0.01) and the lower
winter variance was more evident at high T^ (Figure 10). Thus,
acclimatization to low environmental temperatures in Cyclura appears to
involve an increased avoidance of high and low regulating phase T^'s
without change in mean T
Mean T^ was significantly lower for Cyclura monitored in the
gradient than in the field (Table 5). The July-August gradient mean was
2.8 C lower than the June field mean and 1.2 C lower than the July field
mean. The difference was only 0.9 C for the winter field and gradient
samples. In Figure 11 mean field 's of active lizards have been
plotted against mean gradient 's for lizard species from several
different families. Mean gradient 2%/ s are nearly always lower than
mean field T7„'s above a mean field Tof 37 C. Allowing T^ to rise
above the preferred gradient T^ is probably adaptive in diurnal helio¬
therms temporarily living in hot environments, such as Cyclura in
summer, since morning and afternoon activity periods are thus lengthened
(DeWitt 1967, Licht et al. 1966).

PERCENT OF SAMPLE
59
BODY TEMPERATURE INTERVAL
Figure 10. Seasonal change in Cyclura regulating phase body temperature in
the Caicos and Gainesville, Florida. The February and July field
distributions and the October field distributions are for two and
three nonhyperthermic free-ranging male lizards,respectively.
Each month's field sample consisted of a total of 13 monitoring
days. The two thermal gradient distributions consisted of third
and fourth day results for two males and two females.

60
There are at least three other reasons why lizards commonly have
higher field T^'s than gradient i^'s. First, Regal (1971) found that a
dominant Klauberina rivers-iana provided with a subdominant lizard in the
same gradient box maintained a higher T^ than the same dominant lizard
monitored separately. Since gradient lizards are usually tested singly,
Tb's may be lower due to missing social interaction present in the field.
Secondly, field ' s measured by mercury thermometer rather chan
telemetry, as was the case for practically all the points in Figure 11,
may be biased to those lizards in full sun and plain view as opposed
to sequestered individuals in deeply shaded surface areas (McGinnis and
Dickson 1967). Thirdly, lizards are commonly not fed while in the
thermal gradient, although reptiles often have elevated active temperatures
after feeding: lizards (Cogger 1974), snakes (Regal 1966), and turtles
(Gatten 1974c).
The single anguid plotted in Figure 11, Gevrhcnotus multicarinatus,
which has such a low field temperature mean compared to its gradient
mean, is probably forced to be eurythermal due to its habitat choice.
It is a secretive, ground dwelling lizard living amongst shaded vegetation.
The lizard does occasionally bask, reaching a voluntary maximum of
approximately 35.7 C, considerably higher than the mean for active
lizards (Cunningham 1966). The field T-^' s for the five gekkonids in
the figure were measured while the animals were in their diurnal retreats.
Even though their s during nocturnal activity must necessarily conform
to air and substrate temperatures, gekkonids in their diurnal retreats
commonly seek out T^'s as high or higher than their preferred gradient
Tb's. The high diurnal T-^' s may be necessary for digestion of food
collected during the night (Bustard 1970, Piar.ka and Pianka 1976).

61
Figure 11. Relationship of mean body temperatures in the field and the
thermal gradient for active lizards from several families.
Solid triangles represent agamids, open triangle,anguid; open
squares; gekkonids; solid circles, iguanids; circles enclosing
dots, Cyctuva at Site I in June and Site II in July vs. the
summer gradient mean and at Site II in February vs. the winter
gradient mean; open circles, scincids. Data were taken from
several sources: Bradshaw and Main 1966, Brattstrom 1965, Cowles
and Bogert 1944, Cunningham 1966, DeWitt 1967, Fitch 1954,
Fitzpatrick et al. 1978, Grenot 1976, Heatwole 1976, Huey 1974,
Lee and Eadham 1963, Licht and Basu 1961, Licht et at. 1966,
McGinnis and Brown 1966, Norris 1953, Parker and Pianka 1974,
Pentecost 1974, Pianka 1971, Pianka and Parker 1972, Pianka and
Pianka 1976, Spellerberg 1972b, Vance 1973, Wilhoft 1958,
Wilhoft and Anderson 1960.

62
Egg laying was delayed in the Gainesville pen. A female captured
on April 4 laid in Gainesville on June 12, within the normal Caicos
nesting season of the first two weeks in June. However, another female,
which had been captive in Gainesville since July 30, 1974, laid in the
pen on July 25, 1975. Yet another female, captive since February 25,
1976, laid on July 25. The rate of egg development may have been slower
in Gainesville's cooler climate or shorter photoperiod (Mayhew 1961).
However, the delays may also have been partially due to a lack of nesting
burrows in the pen, resulting in egg retention. For example, a caged
female on Water Cay, captured on June 5, did not lay until June 17.
Habitat Complexity
A foraging Cyoluva aarinata, a semiarboreal lizard sometimes
traveling up to 1600 ft (488 m) a day, should receive a radiant heat
input from direct and filtered sunlight that is inversely proportional
to the density and height of the plants in its territory. If the habitat
niche of the species is sufficiently broad, the regulating phase Tfo of
individual lizards should increase significantly as vegetational cover
decreases. This has been demonstrated for the eurythermal lizard, Anolis
aculatus (Ruibal and Philobosian 1970). Cyalura burrows were absent
from both beach and swales, infrequent in Mixed Woodland, perhaps due
to low food plant diversity, and infrequent in Mesic Coppice, perhaps
due to insufficient open areas for basking. Iguanas once lived in all
the other vegetation zones on Pine and Water Cays, reaching the highest
density in Rocky Coppice. This generalist habitat niche is not surpris¬
ing considering the lack of significant vertebrate competitors.

63
Monitoring sites were carefully chosen in order to find the maximum
mean Tâ– , difference due to differences in vegetational cover. The highest
regulating phase 2^'s were expected in the sparsely vegetated Open Scrub
nearest the beach. However, man and his dogs had nearly extirpated the
lizards from this zone by the time my study commenced. Therefore, Site I
had to be located further inland in semi-Open Scrub bordering the Dense
Scrub. Lizard T^'s were expected to be lowest at Site II, a thickly
vegetated area covered with a mixture of Dense Scrub and Rocky Coppice,
bordering a rocky coast. The site was crisscrossed by a network of
bulldozed roads. On the average, bushes and trees were 63 percent taller
at Site II than at Site I (Table 6). Site II also had approximately 82
percent more shade, when roads were excluded from the transects.
As expected, mean T. was significantly higher at Site I, although
the difference was not very great (mean difference = 1.6 C, Table 5).
The small lizard sample sizes were compensated for somewhat by the long
monitoring times. Inland environmental temperature and wind velocity
differences between sites were negligible. Thus, the T^ difference
was probably due to the vegetational cover difference. The site
difference was significant in hyperthermic as well as nonhyperthermic
lizards (Figure 12). The mean T^ difference was slightly less for
maximally hyperthermic than for nonhyperthermic lizards. The high
Tj^ distributions were negatively skewed while the low 2^ distributions
were more normally distributed.
Caged lizards were monitored in an attempt to produce a significant
difference in mean Tâ– , by varying the amount and kind of shade available
in the lizard's habitat. It was hypothesized that, given only two
burrows as heat sinks, each with relatively constant air and substrate

Figure 12. Influence of habitat on body temperature. Each upper
frequency distribution is for three days of monitoring
of a maximally hyperthermic adult male Cyclura earinata
(Site I, solid line; Site II, dashed line). The lower
distributions are for the same two lizards cn the three
days of lowest mean Tunaffected by long periods of
overcast. Horizontal"lines are ranges, vertical lines
are mean Tfo's, and rectangles are plus and minus two
standard errors of the mean.

65
t5 interval

66
Table 6. Vegetational cover at Sites I and II on Water
Cay. The height of a bush or tree was
measured at approximately 7.5 m intervals along
the transects, which were 30 m long.
SITE I
SITE II
N (PLANTS)
HEIGHT OF VEGETATION
20
20
MEAN HEIGHT (M)
1.77
2.88
2 SE
0.39
0.53
RANGE
0.79-3.88
1.12-4.57
N (TRANSECTS)
PERCENT SHADE
5
5
MEAN PERCENT SHADE
43.3
79.0
2 SE
7.5
8.3
RANGE
33.1-51.7
68.9-90.4

67
temperatures below the lizard's minimum regulating phase Tmean T^
and the standard deviation of mean T^ would be greater than for the
same lizard given vegetational shade, two burrows, and branches,
which allowed greater convective cooling of lizards perched off the
ground. Overall, mean T^ was significantly higher in the complex
habitat (p < 0.01), but individually, this was true only for Lizard 5
(Table 7). The results may have been opposite to those expected
because of lizard hyperthermia. Lizards 5-7 had their hyperthermic
peaks while in the complex habitat, perhaps overriding the effect of
the vegetational shade and the branches. The standard deviations of
overall mean T- in the two habitats were about the same, again contrary
to expectation. The mean of the four lizards caged in the complex
habitat did not differ significantly from the two free-ranging lizards
also monitored at Site I in June (Table 5). Clearly, caged Cycluva
required only a very simple habitat to thermoregulate as well as free-
ranging lizards.
Cloud Cover
Mean regulating phase T, decreased linearly as overcast time
increased (Figure 13). The correlation coefficient was significantly
less than zero (p 0.0001 for v = -0.042, X = -0.042 Percent Cloud
Cover + 39.582). As expected, the correlation coefficient increased as
the weather got colder (June-July, r = -0.193, N.S.; Oct, r = -0.344,
N.S.; February, r = -0.541, p <_ 0.01). Mean T^ per day was never
depressed below 36.5 C by cloud cover times of less than 30 percent
of total time. Continuous overcast periods had to be fairly lengthy
to depress mean T^ below the mean minimum T^ for clear weather, since

68
Table 7. Body temperature of Cyclura cavinata confined to cages
with two structurally different habitats. Lizards were
first monitored in the complex habitat for three to
four days, then in the simple habitat for two to five
days.
COMPLEX
HABITAT
SIMPLE HABITAT
LIZARD
DAYS
N
MEAN
rp
1B
SD
2SE
DAYS
N
MEAN
tb
SD
2SE
4
3
60
39.9
1.17
0.30
2
48
40.4
1.40
0.40
5
4
101
41.1
0.85
0.17
3
95
39.6
0.87
0.18
6
3
64
38.1
1.10
0.28
5
97
37.7
1.54
0.31
7
4
104
38.8
3.54
0.69
3
90
38.2
1.15
0.24
Overall
14
329
39.7
1.63
0.18
13
330
38.8
1.60
0.18

MEAN BODY TEMPERATURE
43
° 0 10 20 30 40 50 60 70 80
PERCENT OVERCAST TIME
Figure 13. Effect, of cloud cover on mean regulating phase body temperature. Practically all field monitoring
is included (N = 124 days). Each symbol represents one day's monitoring; February (solid tri¬
angles), June (solid circles), July (open circles), Angust-September (solid squares), and October ^
(open squares). The regression line was fitted by the method of least squares.

70
large adult Cyclura cooled slowly and radiative heat input continued
during light overcast. The high mean T^ of 42.3 C during June, achieved
with an overcast time of 54 percent, was possible due to the thinness of
the cloud cover.
Lizard T-^ sometimes increased rather than decreased during over¬
cast periods, especially when the overcast lasted only a short time
(Figure 14). When the lizard had been heating prior to the overcast,
the post overcast T- rise can be explained as a result of thermal lag.
However, T. also commonly rose initially during overcast after decreasing
before overcast, the maximum T^ during overcast being higher than the
previous maximum before overcast. Iguanas sought out hot substrates
avoided while the sun was shining and assumed a prostrate posture
(Figure 44, Illustrations 4 and 11). It appears that higher core Tj_
was tolerated under these conditions, since overheating, as indicated
by body surface temperature, was not possible. The noises of lizard
activity; that is, walking, chasing, arboreal feeding, and dropping to
the ground after feeding, nearly ceased during extended overcast periods.
Except for short walks, iguanas remained practically motionless until a
few minutes after an extended overcast cleared. The abruptness of the
activity cessation varied directly with the severity and duration of
the overcast. Short periods of overcast, frequently encountered,
resulted in little or no curtailment of activity. The T^ change during
overcast differed slightly with season. On the average, decreased
2.0 C per hour during February, 1.3 C during June-July, and 2.6 C
during October, the month of heaviest overcast. A large increase in
2^ was most likely during summer overcast.

Figure 14. Effect of overcast duration on lizard body temperature change. Data
from February are indicated by open triangles, June and July by solid
circles, and October by open circles. Regression lines were calculated
by the method of least squares. All lizards were probably out of their
burrows during the full overcast period.

ro

73
Shaded Air Temperature
In Huey and Slatkin's cost-benefit model for thermoregulation
(1976), the value k represents a lizard's "thermoregulatory strategy,"
varying from perfect thermoregulation (k = 0) to passive nonregulation
(k = 1). The linear regression coefficient r for a graph of T^ vs.
shaded T' at the lizard's body surface is a good estimate of k. A lizard
living in a tropical forest would be expected to have a high r value,
since direct solar radiation near the forest floor is only patchily
available. On the other hand, regulating phase T- should be practically
independent of shaded T^ in a heliothermic lizard like CycZuva, living
in fairly open habitats. Cyolm'a, with its low r values, is a nearly
perfect thermoregulator (Figure 15; v = 0.151 in Graph 1, 0.013 in
Graph 2, 0.131 in Graph 3, and 0.154 in Graph 4),
Mean T^-T^ during the regulating phase at Site II was least in
July (X = 5.9 C, SD = 2.00, Range = 1.2-10.4 C, N = 68), greater in
October (X = 7.2 C, SD = 1.65, Range = 4.2-12.2 C, N = 67), and greatest
in February (X = 11.0 C, SD = 2.11, Range = 5.5-15.6 C, N = 58), since
shaded air temperature decreased faster seasonally than 2b,, again
expected in a heliotherm.
One problem with this analysis is that the air temperature was not
measured at the surface of the lizard, but rather at a site near the
blind. An attempt was made to select the lowest shaded air temperature
at the site. The assumption was made that this air temperature was also
available to the lizard. If this assumption is correct, the value of r
is still a measure of the degree of heliothermy. Of course, the
correlation of any instantaneous thermal input with core T^ declines as
body size increases in a lizard moving rapidly through a varied thermal
environment.

BODY TEMPERATURE
74
22 26 30 34 38 22 26 30 34 38
SHADED AIR TEMPERATURE
Figure 15. Correlation of lizard body temperature during the regulating
phase with shaded air temperature. Each graph represents the
data for one dominant male (Graph 1, Lizard 3; Graph 2, Lizard 9;
Graph 3, Lizard 11; Graph 4, Lizard 3). was independent of
shaded air temperature in all four lizards. Data were collected
on five to seven clear days in February (Site II), June (Site
I), July (Site II), and October (Site II), respectively.

75
Substrate Temperature
Heliothermic lizards are able to behaviorally maintain body tempera¬
ture within narrow limits, somewhat independently of ground surface
temperatures, which fluctuated with changes in daily and seasonal
thermal flux, cloud cover, and wind speed. McGinnis (1970) coined the
term "thermoregulatory flexibility" for this ability of heliothermic
lizards. Even during the hottest days, shaded substrates other than in
burrows were available at temperatures well below lizard T^ (Figure 16).
Iguanas rested quietly in the shade during maximum midday insolation
rather than retreating to a burrow, in a posture somewhere between
prostrate and upright (see Figure 44). was usually considerably
less than the sand surface temperature in the sun (T ) on clear
days, except during the early morning and late afternoon (upper and
middle graphs). Hyperthermic lizards in June decreased the Tqsn~^-u
difference more than February nonhyperthermic lizards by spending con¬
siderable time in exposed areas. Iguanas remained above ground during
mildly heavy overcast on partly cloudy days, receiving maximum insol-
ative heat input with minimum convective heat loss by sitting in exposed
areas protected from the wind. T-, was commonly higher than T , under
these conditions (lower graph). Nonetheless, T^ eventually decreased
during extended overcast. 2V decreased more slowly than T in the
late afternoon on clear days due to the longer heat retention time of
the lizard's body than the surface of the windswept sand. The negative
T -±i difference in the late afternoon was not as great as shown in
the middle graph, however. Large males, the last lizards to submerge,
sought out the last remaining sunspots in the habitat, unavailable to
me for temperature measurements without disturbing the lizards. Also,

Figure 16. Thermoregulatory flexibility. Tfo's of emergent lizards
have been subtracted from the sand surface temperature in
the shade (solid circles, Tssh - ^Jj) and from the surface
temperature of sand continually exposed to the sun (open
circles, Tssn - . Solid and dashed lines encompass the
Tgsh - Tj, and Tgsn - Tfo values, respectively. Solid and
dashed lines also indicate the hourly mean values (upper
two graphs only). Data for the upper, middle, and lower
graphs were collected on 14 clear days in February at
Site II, 8 clear days in June at Site I (lizards hyper¬
thermic), and 5 partly cloudy days in June at Site I,
respectively. Mean Tss}i, Tfo, and Tssn values are given
on the left side of each graph.

77

78
iguanas kept their ventral body surfaces in contact with the substrate
during this period. The sand surface temperature important in con¬
ductive heat exchange, lying beneath the motionless lizard's body, did
not decrease as rapidly as T
^ J ssn
Patterns of Heat Transfer between Penned Iguanas and
their Environment: The Three-Transmitter Study
Heat transfer occurs everywhere on the lizard's body surface.
Dorsal heat exchanges are primarily radiative and convective while ventral
exchanges are also conductive. Heat loss due to evaporation of water from
cutaneous and respiratory surfaces and heat gain due to metabolism and
condensation of water vapor will be assumed to be approximately equal
in the following analyses and thus, ignored. Based on temperature
received from the dorsal surface, gut, and ventral surface transmitters,
four heat transfer conditions were possible: 1. The lizard simultaneously
received a net heat input from both dorsal and ventral body surfaces,
that is, both the dorsal and ventral transmitter temperatures were
higher than the gut temperature, 2. the lizard received heat dorsally
and lost it ventrally, 3. lost heat dorsally and gained it ventrally,
and 4. lost heat both dorsally and ventrally. All four heat transfer
conditions usually occurred sometime during an emergent lizard's day,
as illustrated in Figure 17 for a large male.
Three problems concerning the limitations of the transmitter
technique should be pointed out. Ideally, the time constant for
transmitter sensing of a rapid temperature change in the environment
should be infinitely small. However, this was not the case. For
example, the lizard in Figure 17 moved to the shade at 1109, resulting

Figure 17. Three transmitter records of a large male Cyc'luva aavinata in the Gainesville pen on
August 21, 1975. The lizard's black body temperature at five mm above and immediately
anterior to the rear legs, gut temperature, and ventral surface temperature, also just
anterior to the rear legs, are represented by the dot and dash line, solid line, and
dotted line, respectively. The lizard was in the shade much of the day (rectangles),
occasionally entering different plywood shelters (S2, S3, etc.). Transitions from one
heat transfer condition to another are demarcated by vertical lines and labeled by
number at the bottom of the graph. The heat transfer conditions are: 1) dorsal and
ventral surface temperatures higher than the gut (core) temperature, 2) dorsal surface
higher, ventral surface lower than the core, 3) dorsal surface lower, ventral higher
than the core, and 4) both surfaces lower than the core. The day was almost perfectly
clear.

og/l 0£,9i Qggl Qg.H Qggl Qggl QgJI Qg.OI Q£, 6
TEMPERATURE
08

81
in a rapid change from Condition 2 to Condition 4. When the transition
time was taken from the temperature records, however, Condition 4 did
not start until 1112. The second problem was the small number of
transmitter monitoring sites. Large heat exchange differences often
occurred at different locations on the lizard's body surface. For
example, the anterior portion of the lizard was often in the shade
while the dorsal black body probe was in the sun. Also, fairly large
internal temperature differences occurred, as between the head and the
gut. It is undoubtedly technically feasible to double or triple the
number of temperature monitoring sites on a free-ranging lizard the
size of an adult male Cyalura. Thirdly, the surface probes should
have been implanted subcutaneously, since temperatures there were
probably of greater biological significance. The temperature difference
was probably minimal between the subcutaneous area and the ventral
surface as measured. However, dorsal black body temperature probably
differed somewhat from the dorsal skin surface temperature. Hopefully,
since the thermal lag time of the transmitter probe was less than the
subcutaneous area, the calculated times for the heat transfer con¬
ditions were not grossly inaccurate.
Several patterns of heat transfer were consistently observed during
the morning heating and regulating phase. Only Conditions 1 and 2
occurred during the morning heating phase on clear days. Typically,
lizards emerged from the plywood shelters and, after a short delay,
walked to a sunlit patch in the still partially shaded pen, where sand
surface temperature exceeded gut temperature. The resulting Condition
1 lasted a variable portion of the morning heating phase, depending on
whether, how soon, and how many times the animal moved to another sunlit

82
area. Condition 4 typically alternated with Condition 2 during the
early regulating phase, when lizards shuttled at a high rate. As
substrate temperatures rose and lizards tolerated these higher tempera¬
tures, Conditions 3 and 1 became more common. Lizards probably
tolerated higher body surface temperatures than core temperatures,
as evidenced by the mean maximum temperature of the dorsal black body
transmitter (X = 45.8 C, SD = 2.0, Range = 40.6-50.0) exceeding the
mean maximum of the gut (X = 38.3 C, SD = 1.0, Range = 36.1-39.7).
However, the mean maximum black body temperature did not exceed the
critical thermal maximum of the core (X = 46.2 C). Lizards commonly
returned to the plywood shelters for lengthy periods during midday hours,
where nearly stable body temperatures were eventually temporarily
attained (Figure 17). Lizards usually avoided the shaded sand avail¬
able underneath the shelters during this time in favor of the higher
temperatures in the shelters. The same shade choice selectivity with
resultant T^ stabilization was observed in the field. Condition 4
predominated when the pen became shaded in the afternoon, since lizard
surface temperatures declined faster than gut temperature. By seeking
out relatively warm, recently shaded substrates and assuming a
prostrate posture, lizards temporarily stopped their core temperature
decline (Figure 17, 1630-1642). They were not usually thigmothermic
during this time, however, since ventral surface temperature did not
usually rise above core temperature.
The most common heat transfer condition was Number 2, occurring
an average of 51.3 percent of the morning heating plus regulating phases
(Figure 18, left graph). Condition 3 occurred only 2.5 percent of the

Figure 18. Partitioning of heat transfer in active penned Cyotuva aavinata. Three
large males were monitored sequentially in August for a total of eight
days. The times the lizard spent in each of the four heat transfer
conditions during morning heating and regulating phases are graphed on
the left. In the right graph, dorsal and ventral heat gains are compared
relative to dorsal and ventral heat losses, respectively. Circles
symbolize lizard core temperature. An arrow points toward or away
from a circle when dorsal or ventral surface temperature is greater
or less than core temperature, respectively. Horizontal lines are means,
vertical lines are ranges, and rectangles are plus and minus two standard
errors of the mean.

HEAT TRANSFER CONDITIONS
DAILY UNITS (MIN C)

85
time, restricted to the regulating phase. Lizards gained heat dorsally
more of the time than they lost heat and were thigmothermic only 20.7
percent of the day, 9.3 percent during the short morning heating phase.
To crudely compare dorsal heat gain with loss and ventral heat gain with
loss, the area between gut and dorsal black body temperature lines and
gut and ventral surface temperature lines in graphs like Figure 17 were
measured with a planimeter. Cyclura gained approximately five times
more heat than it lost dorsally and lost five times more heat ventrally
than it gained (Figure 18, right side). This is a quantitative way of
saying the lizard was a heiiotherm rather than a thigmotherm.
Average core body temperature increased most rapidly during
Condition 1, followed by Conditions 2, 3, and 4, respectively (Figure 19).
Morning heating rates were significantly higher than heating rates of
the same Conditions 1 and 2 during the regulating phase. The maximum
rate recorded, +20.5 C/hr, occurred during Condition 1 during morning
heating. The variance of mean T% change was highest for Condition 2
during both the morning heating and regulating phases, due to the wide
range of radiative and convective heat inputs vs. conductive heat losses
possible during this condition. Iguanas occasionally cooled during
Condition 2 when substrate heat loss exceeded dorsal gain.
The number of daily heat transfer conditions was linearly and
directly related to daily lizard activity (Figure 20, left graph). The
correlation was nearly perfect. The line extrapolated to practically
zero heat transfer conditions at zero activity. The relationship is
intuitively obvious; the more the lizard moves in the pen, the greater
the number and variety of thermal environments it encounters. The
number of heat transfer conditions was also linearly and directly related

86
2 ¡ 2 3 "4
1—M.H.P—> 1 R.P 1
HEAT TRANSFER CONDITION
Figure 19. Rate cf core body temperature change during the four heat
transfer conditions. Rates during both the morning heating
and regulating phases (M.H.P. and R.P.) are given. Three
male lizards were used, weighing 1195, 1240, and 1260 g.
Horizontal lines are means, vertical lines are ranges, and
rectangles are plus and minus two standard errors of the
mean.

Figure 20. Correlations of the sum of daily heat transfer conditions with lizard
activity and mean body temperature. An activity unit is any behavioral
sequence; a walk between two points, a feeding bout, digging sand at the
pen wall, etc. The mean TjJs were calculated for the morning heating
plus regulating phases. Correlation coefficients for the regression
lines are given at the bottom of each graph. Data were collected for
three males in the Gainesville pen over a period of eight days.

DAILY ACTIVITY UNITS
70
L¿l J I I I
29 30 31 32 33 34
DAILY MEAN Tfa
35
00
00

89
to mean T^ (Figure 20, right graph), but the r value was not as great
(p < 0.05). The regression line extrapolated to 29.1 C mean T^ at
zero heat transfer conditions, approximately equal to the mean Th of
summer lizards in the field just prior to emergence. This relation¬
ship is intuitively obvious; as overcast skies decrease per day, lizard
mean 2b. and activity and the number of thermal environments available
to the lizard all increase.
Statistical Analyses of Regulating Phase
Body Temperature
Body Temperature Maxima and Minima
Heath (1970) proposed that the thermoregulatory behavior of lizards
is controlled by maximum and minimum limit temperatures or setpoints
separated by a nonthermoregulatory zone. Lizards seek shade when the
maximum limit temperature is reached, a mandatory behavior since further
rise in T-^ results first in panting, then paralysis and death. Sun
seeking behavior occurs at the minimum limit temperature and is also
mandatory, since temperature dependent processes such as recovery from
lactic acid debt following activity proceed at such slow rates below
the minimum that lizard efficiency is considerably reduced. Due to
the daily cyclic change in available environmental temperatures,
maximum and minimum limit temperatures may be of little importance to
the lizard early and late in the day and during the middle hours,
respectively.
Maximum and minimum limit temperatures can be accurately determined
only in two-temperature selection experiments, with lizards shuttling
between two chambers at constant ambient temperatures known to lie above

90
and below the limits (Berk and Heath 1975a, b, Barber 1976). Non-
thermoregulatory movements have no effect on T^ maxima and minima in
two-chamber experiments. However, in continuous gradient experiments
and in the field, the same movements can add intermediate minimum and
maximum 2^'s, which inaccurately narrow the temperature range between
mean value estimates of the limit temperatures. Also, maximum and
minimum 's often occur in motionless lizards in the field; that is,
at the start and end of overcast periods and when radiant input wanes
late in the day. However, these nonthermoregulatory values were easily
identified on the T^ records and omitted from analysis.
Frequency distributions of minimum regulating phase i^'s were
commonly negatively skewed, whereas maximum 2^'s were commonly positively
skewed (Table 8). The statistic a^ (Remington and Sc’nork 1970) was used
to estimate the degree of skewness and test for it in samples larger
than 25. Values were commonly negative (10 of 15) for minimum T^'s
and positive (12 of 15) for maximum T-^' s. However, the results showed
disconcerting inconsistencies, even when sample sizes were adequately
large for significance testing. All minimum and maximum T, frequency
distributions were normal for Dipsosaurus dorsalis monitored in a two-
chambered gradient (Barber and Crawford 1977), in contrast to the
CycZura results in a continuous gradient and in the field.
The standard deviation of the mean of the 2V maxima (SD ) was
b max
not significantly different from the SD of the mean of the minima (SD . ,
° mm
F-test, p < 0.05) in any lizard monitored in the field or continuous
s
thermal gradient (Table 9). In contrast, Berk and Heath (1975a) found
the average SD of the minima to be significantly greater than the maxima

Table 8
Skewness of frequency distributions of minimum and maximum body temperatures.
Each lizard's mean daily T-^ varied less than 1 C for the days in the sample.
Field minimum and maximum r¿'s occurred during times of unobscured sun. Only
Lizard 9 was hyperthermic.
MONTH
LIZARD
LOCATION
N
MINIMUM Tg S
Skewness (aq)
P
N
MAXIMUM Tb'S
Skewness (a-^)
P
Feb
2
Free-Ranging
18
-0.941
—
19
+0.019
—
Feb
3
Free-Ranging
35
+0.356
NS
48
+0.754
0.05
June
7
Caged
28
+0.221
NS
34
+0.084
NS
June
8
Free-Ranging
22
-0.864
0.05
28
+1.874
0.01
June
9
Free-Ranging
24
-0.439
0.05
30
-0.393
NS
July
10
Free-Ranging
26
-0.884
0.05
33
-0.003
NS
July
11
Free-Ranging
35
-0.250
NS
43
+0.069
NS
Oct
2
Free-Ranging
12
-0.552
—
17
+0.177
—
Oct 3
Field Lizards (X)
Free-Ranging
38
+0.072
-0.365
NS
57
+0.606
+0.354
0.05
Dec
1
Gradient
13
-0.386
—
17
+0.066
—
Jan
3
Gradient
26
-0.808
0.05
31
+0.236
NS
Jan
4
Gradient
43
+0.645
0.05
55
+0.972
0.01
July
3
Gradient
14
-0.596
—
16
+0.922
—
July
5
Gradient
17
-0.498
—
22
-1.173
—
Aug
6
Gradient
17
+].070
—
18
+1.154
—
Gradient
Lizards (X)
-0.096
+0.363

Table 9. Means and standard deviations of minimum and maximum body temperatures of
individual lizards and correlation of maximum vs. minimum and minimum vs.
maximum T-u pairs. An asterisk indicates the correlation was significant
(p < 0.01).
rr UTMTU» m M,VTM. MAX-MIN MIN-MAX NUMBER OF
iT-t MTNTMA i MAXIMA
B jb iiflAiufl SD MINIMA^ CORRELATION CORRELATION SHUTTLE
MONTH
LIZARD
LOCATION
X
SD
X
SD SD MAXIMA
PAIR (r>)
PAIR (r)
PAIRS (N)
Feb
2
Free-Ranging
37.6
1.06
38.5
0.97
1.09
0.802*
0.549*
34
Feb
3
Free-Ranging
38.0
1.06
38.8
1.09
0.97
0.876*
0.842
51
June
4
Caged
38.7
1.18
41.1
1.26
0.94
0. 338
0.383
24
June
5
Caged
39.6
1. 37
40.8
1.17
1.17
0.833*
0.811
36
June
6
Caged
37.2
1.16
38. 5
1.04
1.12
0.714*
0.614*
27
June
7
Caged
37.8
1. 53
39.4
1.25
1.22
0.792*
0.849*
39
June
8
Free-Ranging
38. 7
2.58
40.3
2.07
1. 25
0.925*
0.857*
40
June
9
Free-Ranging
40.8
1.44
41. 5
1.30
1.11
0.937*
0.921
46
July
10
Free-Ranging
38.0
1.84
39.3
1.68
1.10
0.896*
0.861*
56
July
11
Free-Ranging
39.0
1.68
40.1
1.49
1.13
0.864*
0.827*
52
Oct
2
Free-Ranging
38. 7
0.93
39.6
0.81
1.15
0.584*
0.462
16
Oct
3
Free-Ranging
37.8
1.59
39.0
1.41
1.13
0.800*
0.740*
43
Field Lizards (X)
38. 5
1.45
39.7
1.30
1.12
0.780
0.726
July
3
Gradient
35.6
1.47
37.8
2.01
0.73
0.664*
0.057
15
July
5
Gradient
36.5
2.37
38.3
2.00
1.18
0.554*
0.546*
18
Aug
6
Gradient
35.4
1.56
37.4
1.58
0.99
0.708*
0.849*
20
Aug
7
Gradient
36.2
2.35
39.9
1. 54
1.52
0.261
0.287
10
Dec
1
Gradient
36.3
0.86
37.6
0. 74
1.17
0.167
0.647
13
Jan
4
Gradient
36.8
0.74
37.7
0.80
0.92
0.651*
0.587
45
Jan
3
Gradient
37.7
0.83
38.8
0.61
1.36
0.660
0.548
28
Gradient Lizards (X)
36.4
1.45
38.2
1.33
1.09
0.524
0. 503
Gradient Lizards (X)

93
for Dipsosaurus dorsalis confined to a two-chamber thermal gradient
(1.91 + 0.40 SD for the minima, 1.27 + 0.29 SD for the maxima in Dipso¬
saurus, 1.45 + 0.69 and 1.33 + 0.60 SD for Cyolura). SD was
tightly coupled to SD . in individual lizards in the field: SD =
y mm max
0.728 SD . + 0.238 (r = 0.970, p < 0.0005, N = 12). The linear regression
mm r —
equation for the thermal gradient data was practically identical: SD =
max
0.694 SD . + 0.316 (r = 0.804, p < 0.025, N = 7). However, the
mm —
correlation was not as good. SD . increased faster than SD
mm max
The linear regression equation for Berk and Heath's data for Dipsosaurus
was SD = 0.316 SD . + 0.673 (r = 0.435, p < 0.005, N = 36). The
max mm —
correlation coefficient was considerably smaller than for the Cyolura
data, yet the correlation was still significant. Again, SD . increased
° ° mm
faster than SD . This is logical, since maximum 7, 's are closer to
max d
the critical thermal maximum than minimum TV's. SD . and SD were
D mm max
uncorrelated with their mean TV's (SD . = 0.060 X 2V . - 0.841, r =
D mxn b mm
0.131, p > 0.05, N = 12, SD = 0.095 Xy, - 2.464, r = 0.283,
max ±b max
p > 0.05, N = 12). That is, lizards regulating at higher overall mean
7^'s did not have significantly higher or lower spread to their minimum
and maximum 7^'s than lizards regulating at lower overall mean Th's.
Frequency distributions of Cyolura minimum and maximum 7^'s over¬
lapped considerably. When a frequency distribution was compiled for
the difference between the lowest daily maximum and the highest daily
minimum T^ for 70 days of summer monitoring, the distribution was normal,
with a mean of 0.8 C overlap (SD = 1.2 C) and only 15 days (22 percent
of the total days) without any overlap. Berk and Heath found no overlap
in their Dipsosaurus maxima vs. minima (maxima X + SD = 41.7 + 1.66 C,
minima = 36.4 + 1.12 C). Again, the difference in overlap may be

94
primarily due to comparing field and continuous gradient data for Cyolura
with two-chamber gradient data for Dipsosaurus.
A Berk and Heath analysis (1975a) was used to determine how well
minimum and maximum 5^'s were correlated with each other (Table 9). The
first daily maximum T^ and subsequent minimum formed one correlation
pair and the same minimum and the second daily maximum formed another.
The pairings were continued until all the regulating phase maxima and
minima for the day had been included, excluding those during overcast
periods. The correlation coefficient for linear regression, r, was
used as the test statistic. Four correlation combinations were possible:
one or both max-min and min-max pairs uncorrelated (0,+; 0,-; 0,0), both
pairs positively correlated (+,+), both pairs negatively correlated
(-,-), and one pair positively, one negatively correlated (+,-). All
the r values were positive in both field and gradient samples and the
majority of the correlations (68 percent) were significantly (+,+). On
tne average, correlations were higher in the field than in the gradient.
In contrast, examples of all correlation combinations except (+,-) were
found by Berk and Heath in their Dipsosaurus sample: 78 percent (0,+;
0,-; 0,0), 8 percent (-,-), and only 14 percent (+,+). The r value
averages for the max-min and min-max correlation pair samples were both
practically zero (0.070 and 0.022, respectively; N = 36 lizards).
The most important reason for the difference between Cyclura and
Dipsosaurus in the Berk and Heath analysis would seem to be the differences
in environmental temperature regimes the two species were subjected to.
Maximum and minimum T^'s in Cyclura were not totally independent of the
cyclic daily rise and fall in environmental temperatures. The lower r
values for gradient than field correlations in Table 9 are in agreement

95
with this. The following data analysis was used to look more closely
at the problem. In a continuous sequence of two maximum and two
minimum I^'s, all different, there are four possible combinations; both
the second maximum and the second minimum are greater than the first
maximum and minimum (+,+), the second maximum is larger than the first
while the second minimum is less than the first minimum (+,-), the
second maximum declines while the second minimum increases (-,+) and
both second maximum and minimum decline Each hour over which
the sequence extends can be assigned a unit quantity of one. Then, a
frequency distribution for each combination can be computed for each
month's sample of field data, as is illustrated in Figure 21. The +,+
combination was the most common (44.6, 41.8, and 43.5 percent of total
hourly units for February, June-July, and October, respectively). This
is expected due to the low rate of cooling in a large, frequently moving
animal encountering higher temperatures as midday approached. As
expected, the +,+ peak came latest in the day in the hottest months of
June and July and earliest in the coolest month of February. The -,-
combination peak fell at the noon hour in the cooler months of February
and October, when the lizard was more likely to sit in a cool spot
capable of reducing the second maximum or minimum The percent of
total hourly units for was lowest in the hottest months (18.9
percent in June-July vs. 27.3 and 32.0 percent for February and October,
respectively). When the +,+ and categories were summed and compared
with the summed +,- and -,+ categories, the former were greater than
the latter for all three months (February, 71.9 vs. 28.1 percent; June-
July, 60.6 vs. 39.4; October, 75.4 vs. 24.6). The results for the
thermal gradient were practically equal (53.2 vs. 46.8 percent), again

Figure 21. Time distributions of four combinations of maximum and
minimum body temperature. In a sequence of two maximum
and two minimum Tj^' s in a lizard's daily record, the
second maximum can be greater than the first maximum and
the second minimum can be greater than the first minimum
(+,+), the second maximum greater and the second minimum
less (+,-), the second maximum less and the second
minimum greater (-,+), or both second maximum and minimum
less When each hour of each sequence (sequences
overlap by one maximum and one minimum) is assigned a unit
quantity of one, a frequency distribution of the timing of
each combination can be tabulated. Solid, dashed, and
dotted lines designate the February, June-July, and
October field samples, respectively.

10
8
6
4
2
0
4
2
0
6
4
2
0
8
6
4
2
0
97

98
indicating the importance of the daily change of environmental tempera¬
ture in the field. Berk and Heath (1975b) concluded that the high and
low temperature set point mechanisms behaved independently. Field
results for Cyalura do not agree with this. Nonetheless, the capability
for independent change probably does exist in Cyclura.
The highest recorded maximum 2^’s for individual lizards were not
significantly correlated with their lowest recorded minimum T^1s (Figure
22: Y = 0.259 X + 32.4, X = Y at 43.7 C, v = 0.290). However, the means
of the daily maximum T, 's were highly and directly correlated with the
means of the daily minimum 's for individual lizards in the population
sample (Y = 0.802 X + 8.8, X = Y at 44.4 C, r = 0.970). Two lizards
were excluded from the latter regression line calculation: a male which
still shuttled at a low rate after four days in the thermal gradient,
and Lizard 4, a caged, recently spent female which exhibited nest
defense behavior, staying in her burrow much of the time except to rush
at approaching free-ranging males and females outside her cage. The
minimum and mean of minima increased more rapidly than the maximum and
mean of maxima as T^ increased, resulting in both regression lines
extrapolating to X = Y lines of perfect thermoregulation at 43.7 and
44.4 C, respectively, approximately equal to the highest maximum
voluntary tolerance recorded, 43.8 C.
Regression lines were calculated for daily maximum vs. daily minimum
2^'s and mean of daily T^ maxima vs. mean of daily T^ minima for individual
lizards monitored a sufficient number of days (Table 10). Two-thirds of
the maximum vs. minimum Tâ– , correlations were significant, in contrast
to the population sample analysis (Figure 22). Only three X = Y values,
for Lizards 3, 8, and 6, were quite far removed from 43.8 C, the maximum

Figure 22. Correlations of the maximum vs. minimum Tfo and the mean of Tfo maxima vs. minima
for individual lizards. Each point represents the results for a single lizard
monitored over several days. Solid and dashed lines encompass minimum-maximum
and mean of minima-mean of maxima correlations, respectively. The horizontal
dotted line indicates the highest voluntary Tfo recorded for Cycluva (43.8 C).
The r values were 0.290 (p > 0.05) and 0.970 (p < 0.0005) for the minimum-
maximum and mean of minima-mean of maxima correlations, respectively. Lizards
monitored in the thermal gradient are represented by open triangles, lizards
monitored in the field in February by solid dots, June-July by solid triangles,
August-September by solid squares, and October by open circles. The two values
marked by asterisks were not included in the regression line calculation of the
mean of minima-mean of maxima correlation. See text for an explanation.

100

Table 10. Linear regression analysis of daily minimum vs. daily maximum T,'s and
mean of daily minima vs. mean of daily T^ maxima for individual lizards.
LIZARD
2
3
8
9
10
11
3
3
4
CONDITION
Free-Ranging
FR
FR
FR
FR
FR
FR
Gradient
C
MONTH
Feb
Feb
June
June
July
July
Oct
Jan
Jan
Tg RANGE
4.6
5.2
5.5
5.5
7.9
7.9
7.8
4.0
4.5
N (DAYS)
7
7
8
12
10
12
11
8
9
DAILY MINIMUM Tn (X)
D
VS. DAILY
MAXIMUM 7L
D
(Y)
SLOPE
0.041
0.992
0.808
0.652
0.466
0.558
0.702
0.414
0. 881
INTERCEPT
37.7
2.8
10.2
16.1
23.5
19.5
14.2
24.0
6.8
X = Y
39.4
353.3
53.3
46.4
44.0
44.1
47.6
40.9
57.1
r
0.066
0.890
0.875
0. 775
0.470
0.923
0.815
0.507
0.741
P
NS
0.005
0.005
0.005
NS
0.0005
0.005
NS
0.025
X OF
DAILY Tb
MINIMA (X)
VS. X OF
DAILY Tn MAXIMA (Y)
D
SLOPE
0.877
0.912
1.059
0.874
0.710
0.912
0.661
0.592
0.811
INTERCEPT
5.45
4.3
-1.1
5.9
12.4
4.2
14.0
16.5
7.9
X = Y
44.3
48.6
19.0
46.9
42.9
48.3
41.3
40.5
41.6
V
0.850
0.969
0.917
0.981
0.954
0.973
0.877
0.834
0.929
P
0.01
0.0005
0.0005
0.0005
0.0005
0.0005
0.0005
0.005
0.0005
101

102
voluntary tolerance recorded for Cyclura. The correlations of the mean
of the daily maxima vs. mean of the daily minima were all significant.
Only Lizard 8's X = Y value greatly differed from 43.8 C. Thus, if the
sample of monitoring days is fairly large for each lizard, the daily
mean of maxima vs. mean of minima lines usually have similar slope and
lesser length than the dashed line of Figure 22. The position of a
lizard's line along the population line varies with season, vegetational
density and height of the habitat, reproductive state of females, and,
perhaps, the level of social dominance of males.
The pattern of hourly change of maximum and minimum T^ differed
seasonally (Figure 23). The hourly maximum recorded T^ at both Sites I
and II and the mean hourly maximum T^ at Site I were highest late in
the summer afternoons, long after air and substrate temperatures passed
their maxima. Th increased until late in the day in 61 percent of the
summer lizard days monitored (last maximum Tu minus first maximum = 0.5-
3.0 C, X = 1.7, SD = 1.0), remained essentially level in 31 percent, and
trended downward in only 8 percent (the maximum). Mean hourly maximum
T-^ shifted to midday in October, but a large male still occasionally
reached maximum 2^ during late afternoon basking (3 of 13 fair weather
records). Half of all daily maxima still occurred after 1500 in
February, although a levelling off of daily body temperature variation
had occurred.
The timing of the maximum daily T-^ for four summer and four winter
acclimatized lizards gradually shifted to the final hour of the photo¬
phase in the thermal gradient, even though photophase air and substrate
temperatures were fairly constant after an initial four hour increase

Figure 23. Hourly change in minimum and maximum body temperatures.
Each month's sample consists of two free-ranging lizards
monitored for several days. Mean minimum and mean
maximum T¡-,' s for June sample are indicated by open circles
and are connected by dashed lines. Means for the other
months' samples are indicated by horizontal lines.
Vertical lines are ranges, open and solid rectangles are
plus and minus two standard deviations of the mean for
hourly minimum and maximum Tfo' s, respectively. June data
were collected at Site I, all other data at Site II.
Cloud-caused minima and maxima were excluded from the
calculations.

43
41
39
37
35
33
43
41
39
37
35
33
43
41
39
37
35
104
830
1030
1230 1430
HOUR INTERVAL
1630
1

105
(Figure 24). The standard deviation of the mean time on the fourth
day was one-third that of the first day. The daily maximum T-^ remained
during the last hour in two lizards monitored an additional four days.
Thus, a unimodal timing pattern for daily maximum 2b 's was clear in both
field and gradient. Cyalura appears to become more tolerant of high
body temperatures as time progresses toward the late afternoon, when
overheating in direct sunlight is less likely or impossible. Con¬
currently, the mean time of the daily minimum 2^ shifted toward midday
in the thermal gradient, but without a corresponding reduction in the
standard deviation.
The timing pattern of minimum T^'s was less clear in both free-
ranging and gradient lizards. Hourly frequency distributions of the
daily minimum Twere weakly bimodal in June, July, and October, with
the expected dip during the hottest part of the day. The pattern was
reversed in February, with the majority of the daily minima occurring
near midday. The reason for this seasonal difference is unknown.
Caged summer lizards had a different pattern than free-ranging summer
lizards in their hourly frequency distributions of 2b minima and daily
minimum 2,^'s. In the caged summer lizards, both frequency distributions
peaked unimodally between 1100 and 1200, an hour before the maximum
depression in the core T-^'s at shade seeking and sun seeking (see
Figure 46 and corresponding text). The number of hourly 2^ minima
for summer free-ranging lizards did not vary much between 1000 and 1600
and, as stated above, the hourly distributions of daily minimum 2^'s
were weakly bimodal. These two differences between caged and free-
ranging lizards are probably related to the harsher thermal environment
of the cages, especially around midday. Barber and Crawford (1979)

DAY IN GRADIENT
I
MINIMUM Tb
2
3
4
1 I l I I L
MAXIMUM Tb
I -
2
3
4
0
J I I I L
2 3 4 5 6
HOUR OF PHOTOPERIOD
i
JL
7
-L
8
Figiire 24. Timing shift of daily minimum and maximum Tfo's. Eight lizards, half acclimatized to summer
and half to winter conditions in Gainesville, were monitored during their first four days
in the thermal gradient. Horizontal lines are ranges, vertical lines are means, and
rectangles are plus and minus two standard errors of the mean.
106

107
found that as the hot environment temperature in a two-chambered
gradient increased, the colonic temperature of Dipsosaurus at exit
from the heat decreased, similar to the caged Cyolura. However, the
maximum dorsal subsurface temperature remáined constant. Thus, the
subsurface temperature appeared to be more important in controlling
the timing of shuttling away from the heat than the core temperature.
The midday depression in core temperature may simply be the result of
the greater temperature lag expected between surface and core in a more
rapidly heating animal. The panting setpoint shewed a similar midday
depression in caged lizards (see Figure 58 and corresponding text),
indicating that it also may be controlled by a skin input.
Skewness of Body Temperature Frequency Distributions
Cyelura field and gradient T, distributions both trended slightly
toward negative skewness (Table 11). The test statistic used was a^
(Remington and Schork 1970); positive and negative values resulted from
positive and negative skew, respectively. Eight of ten significantly
skewed distributions were negative, equally distributed between field
and gradient. The trend was about equal in both locations. Antipodal
results for different lizards were evident in all seasonal samples,
especially in the field in February and in the gradient in July-August.
The test statistic a^ (Remington and Schork 1970) was used to
determine the degree of kurtosis in the Tfrequency distributions
(Table 11). Kurtosis could be tested for in samples of 50 or more
(Pearson and Hartley 1966, Table 34, p. 208). Values significantly
less than three indicate the distribution is platykurtic, significantly
greater than three; leptokurtic, and not significantly different from

108
Table 11. Skewness and kurtosis of Tu frequency distributions.
Data points (N) were taken from the Tb records at 15
minute intervals for a variable number of monitoring
days. C and FR stand for free-ranging and caged,
respectively. Vs during extended overcast periods
were not included. Only third and fourth day monitoring
results were used for the thermal gradient calculations.
See the text for an explanation of the statistical tests.
FIELD
MONTH
CONDITION
DAYS/N
X Tb
SKEWNESS
(23)
P
MONTH
LIZARD
KURTOSIS
(34)
P
Aug
12(FR)
4/44
39.5
-1.329
0.01
Aug
12
4.911
0.05
Feb
2 (FR)
7/130
38.2
-0.848
0.01
Feb
2
4.001
0.05
Feb
1(FR)
3/48
37.8
-0.718
0.05
Feb
1
3.117
NS
Jun
9 (FR)
8/236
41.3
-0.684
0.01
Oct
2
3.009
NS
Jun
4(C)
5/109
40.1
-0.322
NS
Feb
O
J
2.996
NS
Oct
14(FR)
3/54
39.6
-0.240
NS
Jun
4
2.880
NS
Jun
6(C)
5/114
37.9
-0.206
NS
Oct
14
2.871
NS
Oct
2 (FR)
4/66
39.3
-0.179
NS
Jun
6
2.830
NS
Jul
11(FR)
10/283
40.1
-0.056
NS
Jul
10
2.737
NS
Jun
7(C)
7/194
38.7
-0.018
NS
Jun
5
2.661
NS
Jun
5(C)
6/177
40.3
-0.016
NS
Sept
13
2.572
NS
Jul
10(FR)
7/199
38.9
-0.008
NS
Jun
9
2.481
0.05
Oct
15(FR)
3/58
39.2
-0.002
NS
Jun
7
2.442
0.05
Oct
3 (FR)
9/196
38.8
+0.105
NS
Jun
8
2.356
0.05
Sept
13 (FR)
5/81
38.5
+0.209
NS
Oct
3
2.228
0.01
Jun
8(FR)
7/165
40.3
+0.234
NS
Oct
15
2.132
0.05
Feb
3 (FR)
6/143
38.4
+0.865
0.01
Jul
11
1.778
0.01
X (SD) -0.189(0.497)
X (SD) 2.824(0
.725)
THERMAL GRADIENT
Jul
5
2/56
37.5
-0.904
0.01
Jul
3
5.766
0.01
Dec
1
2/37
36.8
-0.687
0.05
Jan
4
3.344
NS
Aug
7
2/55
37.3
-0.503
0.05
Jul
5
3.143
NS
Aug
6
2/57
35.3
-0.500
0.05
Aug
6
2.751
NS
Jan
3
2/51
38.0
-0.423
NS
Aug
7
2.642
NS
Dec
2
2/34
35.8
-0.030
NS
Dec
1
2.593
—
Jan
4
2/52
37.1
+0.258
NS
Jan
3
2.557
NS
Jul
3
2/53
37.0
+0.908
0.01
Dec
2
2.298
—
X (SD) -0.235(0.588)
X (SD) 3.137(1.114)

IOS
three; mesokurtic or similar to the normal distribution. The field
distributions trended toward platykurtosis (six platykurtic and two
leptokurtic distributions) whereas gradient distributions did not.
The T^ distributions of the field lizards monitored in the summer
were examined on a day by day basis to see if skewness changed in a
consistent way with increasing daily mean T^ (Figure 25). The trend
toward negative skewness was most evident in the midrange of mean .
As expected, positive skewness predominated in the T^ distributions of
hyperthermic lizards. However, the expected predominance of negative
skewness at low mean 2^'s was not present. Clearly, the overall
divergence of distributions from normality was small, since only 7 of
52 distributions or 13.5 percent were significantly skewed.
The mean and standard deviation adequately describe a lizard’s
body temperature frequency distribution only if the distribution is
normal. Since distributions were negatively skewed in Dipsosaurus
dorsalis monitored in a continuous gradient, DeWitt (1967) substituted
the median and the central 50, 68, and 95 percent of the total obser¬
vations to describe the level and precision of thermoregulatory control,
respectively. Figure 26 compares the level and precision of thermo¬
regulation of Cyolura and Dipsosaurus, two fairly closely related
herbivorous iguanines. Median T^ of the desert iguana was approximately
1.5 C above the rock iguana (upper graph), probably due to the more
thermally extreme habitat of the desert iguana. The medians of the
three samples have been superimposed in the lower graph, allowing com¬
parisons of relative thermoregulatory precision. Negative skewness was
expressed at each of the three central percentages in Dipsosaurus, whereas

MEAN DAILY Tb
Figure 25. Skewness of daily Tjy frequency distributions. The sample consisted of four caged and four
free—ranging lizards monitored in June or July. Circled dots are 33 values for distributions
with significant positive or negative skew (p < 0.05). The curve follows the mean 33 values
for the mean Tfo of each 1 C interval. Days with extensive cloud cover were omitted from the
analysis. Hyperthermic lizard 33 values were included.
o

PERCENT OF TOTAL
in
-6 ~o -4 -3 -2 -i 0 4 +2 *3 +4
°C FROM MEDÍAN Tb
Figure 26. Comparison of the level and precision of thermoregulation of
Cyclura carinata and Dipsosaurus dorsalis in continuous thermal
gradients. Cyclura data were obtained from four lizards
monitored on the third and fourth days in July-August (dotted
lines) and third to fifth days in December-January (dashed lines),
Dipsosaurus data (Dewitt 1967) from 11 individuals monitored for
three days (solid lines). Horizontal lines are ranges encom¬
passing 50, 68, and 95 percent of the regulating phase Ty s.
Vertical lines are median T-, 's.

112
the Cyctura distributions were negatively skewed only when 95 percent
of the observations were included. Cyctura's precision was greater
than Dipsosaurus below the median, but only for the winter sample above
the median. The greater thermoregulatory precision of the rock iguana
may be partially due to its larger body size, resulting in lower heat¬
ing and cooling rates. Unfortunately, the validity of the species
comparison is impaired by the differing sample sizes, the use of
different kinds of continuous thermal gradients, and acclimatization
of lizards to different thermal environments prior to testing.
Standard Deviation of Mean Body Temperature
The standard deviation decreased as mean T^ increased (Figure 27),
in agreement with the maximum vs. minimum T^ and mean of maxima vs. mean
of minima correlations previously described. The correlation was not
significant for the February data. The regression lines extrapolated
to zero SD at 45.2 C mean T^ (February), 45.9 C (June-July), and 43.8 C
(October), again similar to the previously described results. Since
mean 2^ was inversely related to the total overcast time per day, a
multiple linear regression analysis for trivariate data was run to
determine the relative importance of overcast time and mean T^ on the
SD. The results indicate that overcast time (2* , in minutes) was
unimportant as a correlate of the SD (February: SD = 11.201 - 0.272
Xm- - 0.001 T ; June-July: SD = 8.406 - 0.191 Xm, + 0.010 T ; October:
x p o 1 b o
SD = 6.328 - 0.143 Xm, + 0.001 T ).
1b o
A large lizard should have a lower SD of mean T^ than a small
lizard, even when their means of maxima and minima are identical, due
to the higher heating and cooling rates of the smaller lizard. Cyctura

SD OF MEAN DAILY
Correlation coefficients for the regression lines were 0.345 (February, p > 0.05), 0.688 (June-
July, p _< 0.0005), and 0.535 (October, p < 0.01).
113

114
has a large ontogenetic weight range. However, I monitored the of
only one small individual, weighing 180 g (approximately three years
old). Thus, summer lizards could be lumped into only two weight classes,
with means of 729 and 1446 g (Figure 28). The smaller lizards had a
higher mean SD at all mean intervals, but the differences were not
significant. The equation resulting from a multiple linear regression
analysis of the summer field data was: SD = 10.02 - 0.217 Xy^ - 0.0004
W,n. Thus, for example, the influence of mean T-^ on the SD was 12.9
times greater than body weight for a 1600 g Cyclura with a mean T^ of
38 C.
The inverse relationship between the SD and X T^ still held when
examined on an hourly basis (Figure 29). Early morning heating and late
afternoon cooling just prior to and after submergence resulted in the
greatest SD's, due to the rapid rates of T- change and the day to day
individual variation in times of emergence and submergence. The
female Lizard 8 and the subdominant male Lizard 10 submerged earlier
than the dominant males 9 and 11, usually still emergent through the
final time interval illustrated in the graphs. In contrast to the
inverse relationship of the SD and lizard mean , the SD's and the
means of shaded air and sand temperatures at ten different sites varied
directly. A lizard had the greatest diversity of shade temperatures to
choose from during the hottest part of the day. Of course, some of
these shaded environments were unsuitably warm. Cyclura was quite
selective in choosing its shade, especially during the hottest part
of a summer day when a large portion of the habitat was too hot for the
lizard to remain in for long.

Figure 28. Effect of body weight and mean Tfo on the standard
deviation of mean T-u. The sample consisted of eight
summer field lizards, five in a low weight group
ranging from 645 to 815 g and three in a high weight
group ranging from 1290 to 1598 g. Vertical lines
are ranges, horizontal lines are means, and rectangles
are plus and minus two standard errors of the mean.

S.D. OF MEAN TK/ DAY
116
0.21-
0
729 14 4 6 729 144 6 729 1446
MEAN WEIGHT (G)
38.0-38.9 39.1-40.4 40.6-424
MEAN Tb RANGE

Figure 29. Hourly change in the mean and standard deviation of
lizard body temperature and shaded air and sand tempera¬
tures. The highest and lowest Tj/s recorded each hour
of each day (0730-0829, 0830-0929, etc.) were used to
calculate the mean T]y and standard deviation (8, 9, 9, and
12 monitoring days in June-July for Lizards 8-11, respec¬
tively) . Ta at a height of 2 cm and Ts were measured on
the hour during three days in August in the shade of ten
shrubs chosen at approximately equal intervals along a
200 m line at Site I.

TIME
MEAN S.D. OF MEAN TEMPERATURE MEAN TEMPERATURE
OJ OJ OJ OJ OJ 4* 4^,
O — ro oj 4- oiOrvj-^cnooOrv)

119
It is interesting to speculate on what determines the mean, long
term, operant temperature of a heliothermic lizard which remains
emergent all day. The most likely choice is the mean of all the shade
temperatures possibly encountered by the lizard during the hottest part
of the day of the hottest part of the year. This is substantiated
somewhat by the upper graph in Figure 28. Huey and Slatkin (1976)
reported that the mean preferred T^ of southwest U.S. desert lizard
species was well correlated with mean, long term July environmental
temperature.
Rates of Lizard Heating and Cooling in the Field
Morning Heating and Regulating Phases
The mean morning heating rate and the mean heating and cooling rates
during the regulating phase were calculated for individual lizards using
the following equation:
R =
ETCS’, .. - Tu ... ) • t 1 • 60]
b Max b Mzn J
N
(1)
where R is the mean rate of heating or cooling (C/hr), t is time (min),
and N is the number of increasing or decreasing temjjerature periods in
the sample. The slight differences in the T^ ranges of y ^
were ignored. The mean regulating phase heating rate exceeded the
cooling rate in nine out of twelve lizards, with a mean cooling/heating
ratio of 0.87 (Table 12). However, none of the ratios were significantly
different from one (p > 0.05).
On the average the mean morning heating rate was 3.3 and 4.0 times
greater than the mean regulating phase heating and cooling rates,

120
Table 12. Rates of lizard heating and cooling in the field during
the morning heating and regulating phase.
LIZARD/
MONTH
WEIGHT
(G)
MORNING HEATING PHASE
1
N X RATE
(DAYS) (C/HR) SD LOW/HIGH
1
2
2/Feb
1665
8
11.2
1.9
9.1/14.9
4.1
3/Feb
1775
6
10.3
3.1
7.6/15.0
3.1
4/June
645
3
11.0
-
6.0/21.0
1.8
5/June
723
-
-
-
-
-
6/June
815
7
7.3
1.5
5.4/9.1 •
3.5
7/June
1290
3
6.7
-
6.1/7.3
1.4
8/June
675
6
11.0
4.0
6.4/16.9
3.0
9/June
1450
11
8.7
1.8
5.0/11.6
5.1
10/July
787
8
9.6
3.1
5.8/14.5
3.1
ll/July
1598
16
8.4
2.2
5.8/14.6
3.8
13/Sept
950
5
11.6
4.0
8.9/18.5
4.6
3/0ct
1672
10
7.8
2.3
5.6/12.0
3.0
X
9.4
3.3
SD
1.7
1.1

121
Table 12. Continued.
N/5
DAYS
HEATING
2
X RATE
(C/HR) SD
REGULATING PHASE
N/5
LOW/HIGH DAYS
COOLING
3
X RATE
(C/HR) SD
LOW/HIGH
3
2
28
2.7
1.9
0.4/9.5
28
2.1
1.3
0.6/6.0
0. 76
31
3.3
3.1
0.7/13.2
31
3.0
2.1
0.6/8.0
0.90
22
6.0
4.4
0.8/14.0
22
5.2
3.6
1.3/12.4
0.88
29
2.5
1.9
0.2/7.5
29
2.0
1.5
0.4/7.3
0. 79
20
2.1
1.2
0.2/4.5
20
2.3
1.7
0.4/6.4
1.06
27
4.9
3. 7
0.7/14.2
27
3.4
3.7
0.4/18.0
0.76
29
3.7
4.0
0.2/21.0
29
4.1
4.1
0.4/14.4
1.11
26
1. 7
1.4
0.2/6.0
26
1.1
0.8
0.1/3.7
0.64
36
3.1
2.1
0.2/9.7
36
2.8
2.0
0.4/9.1
0.92
32
2.2
1.4
0.5/7.3
32
2.0
1.8
0.2/12.0
0.92
16
2.5
1.2
0.4/4.6
16
1.8
0.5
0.8/2.7
0.70
34
2.6
1.6
0.4/7.9
34
2.7
1.8
0.4/8.3
1.03
3.1
2.7
0.87
1.2
1.1
0.15

122
respectively (Table 12). Lizards maximized their heating rates
behaviorally for a time before beginning their first long distance
movements away from the home burrow in the morning. Heating and cool¬
ing during the regulating phase commonly occurred in a sun-shade mosaic
under vegetational cover. Various thermoregulatory behaviors, described
later, were intermittently in force during both regulating phase heat¬
ing and cooling. Low rates of regulating phase heating and cooling
were attained for short periods by every lizard in the sample, since
numerous shaded areas were contained in every home range.
The rates of heating and cooling decreased as body weight increased,
due to the decrease in surface to volume ratio. Plots of the logs of
weight specific rates of maximum recorded heating, mean morning heat¬
ing, and mean regulating phase heating and cooling vs. the log of body
weight were linear, and the slopes were approximately equal (Figure
30). The equations, correlation coefficients, and significance levels
for the regression lines were: Y = -1.40 X + 2.40, r = -0.975, p _<
0.0005 for the maximum heating rate, Y = -1.12 X + 1.34, r = -0.925,
p _< 0.0005 for the mean morning heating phase rate, Y = -1.30 X +
1.39, v = -0.839, p <_ 0.0005 for the mean regulating phase heating
rate, and Y = -1.41 X + 1.65, r = 0.831, p 0.0005 for the mean
regulating phase cooling rate. On the average, a doubling of body
weight decreased the maximum heating rate by 62 percent, the morning
heating rate by 54 percent, the mean regulating phase heating rate by
59 percent, and the mean regulating phase cooling rate by 62 percent.
The mean daily regulating phase rates of heating and cooling were
directly correlated with the standard deviation of the mean daily T
mean daily heating rate (C/min) = 11.18 SD of mean daily T^ + 0.35,

Figure 30. Correlation of mean rates of heating and cooling in the
field with lizard body weight. Regression line one
represents the maximum rate of heating recorded (solid
dots, one per lizard), line two; the mean rate of heating
during the morning heating phase (open triangles), line
three; the mean rate of heating during the regulating
phase (solid triangles), line four; the mean rate of
cooling during the regulating phase (open circles). Data
were taken from Table 12 and transformed to weight
specific rates.

MEAN HEATING AND COOLING RATE (°C*G HR )
124
0.032
0.016
0.008
0.004
0.002
0.001
0.0008 -
650 800 1000 1200 ¡400 1600 1800
BODY WEIGHT (G)

125
N = 43, v = 0.746, p 0.0005; mean daily cooling rate = 9.32 SD of
mean daily T^ + 0.51, N = 43, r = 0.563, p _< 0.0005. Figure 31
summarizes the correlations between five parameters which describe
regulating phase body temperatures.
Lizard Cooling in Burrows
The decline in lizard T^ during burrow cooling is best described
by Newton's law of cooling:
ft
dt
-Vi''7’ -
a
a
(2)
Integration yields:
An (Ti - T ) = -at + An(2\ - T ) (3)
oa bs a
where T^ is the lizard core temperature during cooling, is the core
temperature at submergence, T' is the burrow air temperature at the
point of the lizard, a is a cooling constant in minutes \ and t is
time in minutes. By graphing t vs. Ln(- T, the slope can be
calculated by linear regression and is equivalent to -a. Since the
burrows of free-ranging lizards were not disturbed during monitoring,
T^ was estimated by , the lizard's body temperature at emergence.
The equation becomes:
Wb ~ Tbe> = + ln(Tbs - Tbe>
(4)
is a good estimator of Ta since equilibrium between T^ and T' was
attained long before emergence and metabolic heat production probably
caused 2V to be less than 2 C above T .
be a
Lizard core temperature nearly equilibrated with burrow tempera¬
ture after only 19 percent of the nightly submergence time (Table 13).
A mean 2^ decline of 10.4 C required an average of 201 minutes. The

126
Figure 31. Correlations of parameters describing regulating
phase body temperatures.
STANDARD
DEVIATION OF
MEAN BODY
MEAN BODY
MEAN RATE
MEAN RATE
TEMPERATURE
TEMPERATURE
OF HEATING
OF COOLING
BODY WEIGHT
Inverse (June,
July, October
Only)
None
None
? (Insufficient Data for Small Iguanas)
Direct
Direct
Inverse
Direct
Inverse
Inverse

Table 13. Lizard cooling in burrows in the field. Asterisks indicate heavy rainfall
during cooling.
DAY OF
DATA
LIZARD
WEIGHT
(G)
MONTH
BURROW
LETTER
CODE
tb at
SUBMERGENCE
tb at
EMERGENCE
NEXT DAY
COOLING TIME TO
TB ~ TBE = 1 c
(MIN)
COOLING
COOLING TIME CONSTANT
SUBMERGENCE TIME (MIN-1)
1
1
1135
Feb
A
38.0
26.9
272
0.24
0.009
2
1
1135
Feb
A
39.4
26.9
284
0.26
0.009
3
1
1135
Feb
A
37.0
27.0
212
0.19
0.010
1
2
1665
Feb
B
38. 7
25.9
326
0.28
0.008
2
2
1665
Feb
B
38.2
26.2
276
0.24
0.009
1
3
1775
Feb
C
38.4
27.8
251
0.24
0.010
1
8
675
June
D
39.9
29.1
148
0.14
0.016
2
8
675
June
E
42.0
28.6
140
0.13
0.018
1
9
1450
June
F
39.2
29.0
59
0.05
0.051*
2
9
1450
June
F
37.4
29.5
45
0.04
0.053*
1
10
787
July
G
39.6
30.0
224
0.22
0.009
2
10
787
July
G
39.6
30.0
174
0.17
0.013
3
10
787
July
II
39.9
28.0
316
0.33
0.008
4
10
787
July
H
37.9
28.0
242
0.20
0.009
5
10
787
July
H
40.7
29.5
210
0.20
0.011
1
11
1598
July
I
39.8
27.7
216
0. 21
0.012
1
3
1672
Oct
J
36.6
29.6
147
0. 14
0.011
2
3
1672
Oct
J
38.4
29.6
164
0.16
0.013
3
3
1672
Oct
A
37.8
30.2
178
0.16
0.011
4
3
1672
Oct
A
39.7
29.8
128
0.12
0.018*
X 38.9
28.5
201
0.19
SD 1.31
1.36
77
0.07
127

128
cooling constant was not significantly correlated with body weight over
a range of 675-1775 g (log a - -0.17 log body weight - 1.43, N = 17
days, r = -0.287, p < 0.05). Heavy rainfall increased the rate of
lizard cooling by conductively increasing heat loss to the burrow
substrate. The importance of conductive heat loss was indicated by a
comparison of cooling rates in burrows and in the freezer used to
anesthetize summer field lizards. Even though T-, - T values were up
to three times greater in the freezer, cooling constants were considerably
less (log a = -0.49 log body weight - 0.77, N = 5 lizards) than for
animals of equal weight cooling in burrows. Freezer cooled iguanas
sat on a 3 cm wide wooden strip that was at room temperature when
the animal was strapped to it.
The slope of the £n(2^ - Th) vs. t lines remained constant on
only 7 of 29 monitoring days. A single slope decrease occurred on 21
days at a mean T^ of 34.4 C (SD = 2.37, range = 30.0-39.6). A slope
increase occurred during only one night's cooling. Slope decreases
were probably due to a change in lizard posture, probably an increase
in adductive contact of the limbs with the trunk and tail. Except for
this movement, lizards remained quiescent until the hour before emergence,
judging from observations in the laboratory and outdoor pen.
Burrow depth, as measured from the ground surface to the roof of
the burrow terminus, averaged 32 cm (N = 8, SD = 25, range = 19-91) for
adult females and 69 cm (N = 12, SD = 43, range = 25-183) for adult
males (Iverson 1979). The greater depth of male burrows may be due both
to the larger size of males and their greater site fixity over time.
Lizards preferentially dug burrows in windblown, well drained sand along
the sides of limestone ridges and at the edge of clumps of vegetation,

129
where roots helped prevent collapse of the burrow mouth. Lizards
probably retreated nearly or completely to the end of the burrows each
evening (see Burrow Movement).
Both higher nocturnal temperature and greater temperature stability
may be determinants of the rather large mean burrow depth in CycZura.
Diurnal burrow temperature stability is not important since lizards did
not usually use burrows as heat sinks. With no surface vegetation and
except during heavy rain, soil temperature over a 24 hour period in
June varied less than 1 C at depths of 25 cm or more. Soil temperatures
at depths of less than 25 cm exceeded the nearly stable, 25 cm depth
temperature during the day and early night and fell below the 24 cm
level as the next sunrise approached. This depth corresponds well with
the average female burrow depth. An adult lizard chose a ground surface
retreat on only one occasion during the study. He retreated into a pile
of limestone rubble during a late afternoon overcast period, rather than
walking the 40 m to his only sand burrow. T-, was 24.1 C, 4.1 C below
the lowest 24 recorded in his sand burrow. ZV 's at 0730 for Lizard 3,
confined in a field cage in October, were 25.7, 26.0, and 28.0 C, all
below Tbe for the same free-ranging lizard (X = 29.6, range = 28.8-30.2,
N = 9). Winter burrow temperature stability may be even more important
in determining average burrow depth than summer stability. Lizards
equilibrated with minimum surface temperatures during cold fronts would
barely be able to crawl. Seasonal variation in mean T-^& probably did
not exceed 4 C.
Females laid their single clutch of eggs at the end of a burrow
and plugged part of the length with sand, leaving the eggs in an isolated,
air-filled chamber. The depth of the egg chamber was commonly less than

130
other parts of the burrow, providing easier vertical escape for the
hatchlings (Iverson 1979) as well as higher mean incubation tempera¬
tures .
Even though Cyclura did not plug its burrow, except during nesting,
relative humidity should be close to 100 percent at the terminus (McNab
1966), due to the average burrow's considerable length (male X = 4.39 m,
female X = 3.09; Iverson 1979). This has obvious value in water con¬
servation, especially during the dry season. Capillary rise in water
from the fresh water lens under Pine and Water Cays may increase the
percent saturation of the burrow soil in lower lying areas. Since it
requires two inches of rainfall to raise very sandy soil to field
capacity (water weight = 5 percent of soil dry weight) to a depth
of two feet (Meyer and Anderson 1952), the average burrow would rarely
be elevated to this level. However, soil saturation of only one per¬
cent was sufficient to raise relative humidity to nearly 100 percent in
pocket gopher burrows (Kennerly 1964).
T-^&} s varied slightly for the same lizard when using different
burrows. For example, Lizard 8 had mean T^e's of 28.7 (N = 3) and
31.1 (N = 2) for two of its three sand burrows, presumably of different
depths. Different lizards sometimes had slightly different mean 's
(Table 14). The greatest mean difference was 2.3 C (maximum difference =
3.2 C), between Lizard 12 in August and Lizard 13 in September. Com¬
parisons between juveniles and adults would probably prove more striking.
The samples were too small to determine whether burrow temperature
influenced operant T^ during the day.

131
Table 14. Lizard body temperature at emergence in the field.
MONTH
LIZARD
SITE
NUMBER OF
BURROWS
UTILIZED
N (DAYS)
X tbe
SD
LOW
HIGH
RANGE
Feb
■»
X
II
1
4
26.3
1.2
24.5
27.0
2.5
Feb
2
II
1
7
25.7
0.6
25.0
26.9
1.9
Feb
3
II
2
2
-
-
25.2
27.8
2.6
13
26.1
1.0
24.5
27.8
3.3
June
8
I
3
7
29.5
1.2
28.6
31.8
3.2
June
9
I
2
8
29.5
0.1
29.4
29.5
0.1
July
10
II
2
9
30.1
0.8
28.2
31.0
2.8
July
11
II
1
15
28.1
0.2
27.7
28.5
0.8
39
29.1
1.0
27.7
31.8
4.1
Aug
12
I
2
3
30.1
-
29.2
30.2
1.0
Sept
13
III
2
5
27.8
1.0
26.8
28.8
2.0
8
28.6
1.3
26.8
30.2
3.4
Oct
2
II
2
4
27.7
-
24.1
29.5
5.4
Oct
3
II
2
9
29.5
0.3
28.8
30.0
1.8
13
29.0
1.5
24.1
30.0
5.9

ETHOLOGICAL ASPECTS OF THE THERMAL BIOLOGY
OF CYCLURA CARINATA
Lizard Activity
Burrow Movement
The results of monitoring Lizard 2 as it moved in its burrow are
illustrated in Figure 32. Movements toward the burrow mouth prior to
emergence occurred incrementally in up to four steps. The maximum pre¬
emergence activity time was probably greater than the 37 minutes recorded.
When scared into its burrow, the lizard retreated a mean distance of only
1.25 m or 27 percent of the October burrow length. The lizard crawled
far into the burrow in a single movement at the end of the day, some¬
times advancing a short distance further later in the night. However,
the animal went to the end of the burrow on only four of fifteen final
submergences.
Control of Lizard Emergence and Submergence
Mean emergence and submergence times conformed to the seasonal
change in photoperiod (Figure 33). Submergence times at the three
different field sites were more irregularly related to sunset than
emergence times were to sunrise. Submergence did not occur any earlier
in relation to sunset in the dry months of February and March than in
the wet months of October and December, even though the latter months
were slightly warmer. Apparently, activity times were not reduced due
to lack of water during the dry season. Mean activity time decreased
132

Figure 32. Lizard movements in a burrow. Six days of monitoring in
October and six in February are represented for Lizard 2
in a single, unbifurcate sand burrow, 4.6 m long in
October and 3.9 m long in February (horizontal dashed
lines). The October record was seven days long, with
a one day gap on the fourth day. Dotted lines mark
lizard movements in the burrow when I was not present.
The lizard was periodically scared into its burrow
(triangles). The letter E indicates emergence.

LIZARD DISTANCE FROM BURROW MOUTH (M)
m o — m oj -fr m o — oj -A m o - m m o — k> oj -u ai
00
—
1 1 1
1
1 1 Ty 1 r
—i—i—
1 1 1 T“
*T
1 1 1 1 â–  H,"|
►—-“n
/
1
- J
J
- W
^__1
1 j
— i
i
t=r
o
f 1 *
I
i
1
1
oi
- 1
1
I
1 ^
1
1
/
00
/
/
/ s
r ¡
â–º
_*=/.
/
1
|
lO
-n
â–º-4
|
o
rn i
i
O I
d
r= CD 1
CT
—■ 1 d.
r H
£
1
1
1
TL
|
1
|
O)
1
\
00
~f r
/ i
j !
^ |
i— i ^
i
1
o
I t-
.. I—1
i
i
1
I
£
... I
i
L 1 4
i
1
1 =
1
i
i
i
1
1
i
~L |
134

Figure 33. Seasonal change in lizard emergence and submergence times.
The upper and lower curves indicate the times of sunrise
and sunset, respectively. Emergence and submergence times
lie within. Vertical lines are ranges, horizontal lines
are means, and rectangles are plus and minus two standard
errors of the mean. The three sets of numbers refer to
the minutes between sunrise and mean emergence time, mean
emergence and submergence times or the mean activity time,
and mean submergence time and sunset. February, July, and
October data were collected at Site II, June data at Site
I on Water Cay, and March-April, November-December data
were collected on Pine Cay (Iverson 1977).

5
6
7
8
9
LlI 10
2
h—
II
15
16
17
18
136
r M A M J J A S 6 N D J
MONTH

137
three hours and 26 minutes or 37 percent between July and December.
The earliest observed emergence occurred almost two hours after sun¬
rise, at 0640 on Pine Cay in July (Iverson 1979). The latest observed
submergence was by Lizard 11 at 1825 on Water Cay in July, 15 minutes
before sunset and 35 minutes after the sun was no longer visible to
the lizard.
Emergence and submergence times varied with the illumination times
of early morning and late afternoon basking sites, in turn influenced by
variations in relief and vegetational cover. Lizard 3's morning basking
site was located on top of a north-south oriented limestone ridge
adjacent to the Pine-Water Cay inlet. His late afternoon basking site
was about three meters away on the same ridge, on the western side near
the top, facing the late afternoon sun and protected from easterly
winds. The basking sites of Lizard 2, another large male of similar
body weight occupying an overlapping home range further inland, came
into direct sunlight later and went into the shade earlier due to the
lower elevation and obstructing vegetation. As a result, Lizard 3's
mean activity times were 127 and 58 minutes longer than Lizard 2's in
February and October, respectively. Lizard 3 should be a healthier
animal due to having more time at regulating phase T-, 's to digest its
food. Lizard 3 was indeed more robust (43.2 and 47.7 g/cm snout vent
length for Lizard 2, 53.9 and 57.3 g/cm SVL for Lizard 3 in October,
1975, and February, 1976, respectively). However, Lizard 2 increased
in weight by 158 g between October and February, whereas Lizard 3
gained only 103 g. The home ranges of the two lizards were approximately
equal in size, but the relative food abundance in the ranges were not
determined. Relative fitness is best expressed as a reproductive output.

138
The females sharing the home ranges of Lizards 2 and 3 in February
weighed 710 and 1135 g (Lizard 1), respectively. Lizard 1 was the
largest female captured on either Pine or Water Cays and, thus, probably
laid more eggs per clutch than Lizard 2's female. Although these
comparisons are suggestive, a large sample of lizard pairs would have
to be analyzed to determine whether significant differences in activity
times influence individual fitness.
Sex related differences in emergence times did not occur but the
data were contradictory. Lizard 8, an adult female at Site I in June,
emerged earlier than Lizard 9, a dominant male. However, Lizard 1, an
adult female at Site II in February, emerged later than Lizard 3, a
dominant male. Lizards 1 and 3 often used the same burrow in February.
Since Lizard 3 was the larger animal and always submerged last into their
unbifurcate burrow, the order of emergence may have been predetermined.
Dominant males were usually the last to submerge due in part to their
low cooling rates caused by large body size. Also, dominant males had
territorial priority over smaller males at the best late afternoon
basking platforms.
Emergence and submergence times in the Gainesville pen depended
on lizard size (Figure 34). The adult females and the lowest ranking
subadult male emerged later and submerged earlier than the rest of the
adult males. Adult females were subdominant to adult males except dur¬
ing the nesting season and except to the male at the bottom of the male
linear dominance hierarchy, which every lizard chased. Correlation
coefficients for the linear regression were -0.700 for summer emergence
(p _< 0.0005), -0.681 for winter emergence (p _< 0.0005), 0.256 for summer
submergence (p > 0.05) and 0.463 for winter submergence (p < 0.005).

Figure 34. Correlations of times of emergence and submergence with
body weight in penned Gainesville lizards. July-August
emergence and submergence times for eight lizards monitored
for six days are represented by open circles and triangles
and the dashed regression lines; December-February times
for eleven lizards monitored for three days are represented
by solid circles and triangles and the solid regression
lines.

8
9
10
II
12
13
J
15
16
17
18
19
140
C¿í>t> D>

141
Note the lower submergence r values. From the regression equations, a
50 g hatchling was active for 299 minutes in summer and 154 minutes in
winter, 62 and 48 percent of the times for a 1200 g male. The summer
and winter ranges for activity times in the pen were similar to the
seasonal range of mean adult activity time in the field. It should be
pointed out that lizard density in the pen was up to 134 times the 31.1
adults per hectare reported by Iverson (1979) for his Rocky Coppice
field site.
Body temperature commonly declined slightly while a lizard paused
near its burrow mouth prior to emergence or just after emergence prior
to reaching a sun exposed location (25 of 52 records, Figure 5). This
was usually the lowest T^ in the 24 hour period. The largest drop
recorded was 0.7 C for a male on Fort George Island in September.
Usually, the decline was 0.3 C or less.
Body temperature at emergence ranged from 24.9 to 31.0 C in the
field (Table 15). Temperature independent emergence was observed in the
Gainesville pen in January at T-, ' s as low as 15 C, only 2.3 C above the
critical thermal minimum. The minimum substrate temperature recorded
on Pine Cay was 12.5 C (Iverson 1979). Thus, in a very shallow burrow
or in rock rubble at the ground surface, T. at emergence in the field
may occasionally fall as low as 15.0 C.
Final submergence occurred at much higher T^ than emergence (Table
15). 's °f only Lizards 4 and 11 in summer consistently fell below
their regulating phase mean minima prior to submergence, to as low as
31.2 C for Lizard 4 during a fifteen minute rain and 34.3 C for Lizard
11 during late afternoon basking. Iverson (1977) also occasionally-
observed an adult Cyolura emergent during a heavy rain, presumably

Table 15. Emergence and submergence times, temperatures, and light intensities
for free-ranging lizards in the field.
%
EMERGENCE
SUBMERGENCE
MONTH
N
(DAYS)
X
RANGE
SD
2SE
N
(DAYS)
X RANGE
TIME OF DAY
Feb
16
0931
0820-1053
39
20
16
1609
1444-1711
47
24
July
23
0756
0716-0835
19
8
19
1720
1445-1825
67
31
Oct
16
0829
0726-0930
36
18
16
1531
1431-1706
39
20
BODY TEMPERATURE
Feb
12
26.0
24.9-27.2
0.9
0.5
16
38.3
36.3-41.4
1.4
0.7
July
23
28.9
27.7-31.0
1.1
0.5
18
37.6
34.3-42.5
2.5
1.2
Oct
15
28.9
24.1-30.0
1.6
0.8
14
38.7
35.3-41.0
1.5
0.8
LIGHT
INTENSITY (FT-C)
Feb
16
829
477-1298
243
122
15
509
251-972
198
102
July
24
680
390-1069
178
73
19
241
50-900
266
122
Oct
16
810
200-1400
282
141
16
309
88-700
138
69
BLACK
BODY
TEMPERATURE
Feb
16
44.6
36.4-50.5
4.5
2.2
15
37.3
30.4-46.1
3.6
1.9
July
24
41.7
32.0-50.0
4.6
1.9
19
39.0
28.6-49.4
5.2
2.4
Oct
16
43.5
33.6-51.7
4.5
2.2
14
38.5
33.4-43.6
2.6
1.4
SUN EXPOSED SAND TEMPERATURE
Feb
16
36.2
28.8-43.8
4.0
2.0
15
33.3
26.8-39.9
3.2
1.7
July
24
30.4
26.6-42.4
3.3
1.3
19
32.6
27.8-41.9
4.4
2.0
Oct
16
31.4
27.4-37.7
3.6
1.8
16
33.5
30.0-39.9
2.4
1.2
142

143
sometimes with T^ near burrow temperature. Adult lizards would have to
remain emergent after sunset during clear weather for T^ to drop to
near burrow temperature. This was never observed. Neither heat lamps
suspended above the ground nor a plywood box with a metal lid heated
from inside with incandescent lamps could induce Cyclura to remain
emergent after sunset in the Gainesville pen. That is, submergence
could be induced prematurely by foul weather but could not be extended
into darkness even when sufficient heat sources were provided to permit
Tto remain in the regulating phase range.
Both light intensity and black body temperature were usually high
at emergence (Table 15), significantly higher than at submergence
(p <_ 0.05). Thus, Cyclura usually emerged only when its would rapidly
increase above burrow temperature. For example, Lizard 3 reached 29.6 C,
the mean emergence T^ while free-ranging, at 0752, 0804, and 0817 while
confined in a sun exposed field cage. The range of emergence times
while free-ranging, 0727-0839, bracketed these cage times. Medium to
heavy overcast during the normal emergence time of free-ranging iguanas,
constituting 16 lizard days of the study, always delayed the emergence
time. On two occasions in October, when the overcast lasted all day,
none of the four monitored iguanas emerged. On five occasions,
emergence was delayed until after the overcast cleared. On five other
occasions, emergence was delayed yet occurred during overcast. T, rose
appreciably after emergence even though the sky was overcast. On only
two occasions, both during the summer, did a delayed emergence occur
which was not followed by an appreciable rise. In one case T-h rose
from 29.5 C to only 30.3 C in one hour and 59 minutes before the sky

144
cleared and T^ rose rapidly. In the other case, T^ declined from 30.2 C
at emergence to 26.8 C before slowly increasing to 30.9 C, three hours
and 42 minutes after emergence, when the sky cleared.
¿In iguana sleeping a meter or more below the ground near the end
of a four meter long nonlinear burrow probably receives neither light
nor temperature cues revealing the beginning of a new day. Morning
arousal may be due to an internal biochemical timing mechanism or
biological clock. A phase shift (phase delay) of clock timing was
observed after release of all the summer lizards (Figure 35). Re¬
entrainment to the normal emergence time required three or more days.
Submergence times were not affected. Lizards monitored during the three
subsequent field trips did not exhibit delayed emergence. Thus, the
delays must have been triggered by the bacterial infection following
the summer implantation operation or, more likely, by some procedural
step during the operation; for instance, cooling the lizards or keeping
them awake during part of a normal sleep period.
After the biological clock initially gets the animal moving, a
second emergence stimulus probably times the final emergence stage
starting near the burrow mouth. High light intensity, one possible
second stimulus in this two step stimulus-response chain, results in
initiation of the final emergence stage. On the other hand, low light
intensity due to heavy overcast inhibits emergence. The final emergence
stage commonly required two steps. First, the lizard extended its head
above the ground surface. This usually required exposure of most of
the trunk, since in sandy soils the burrow mouth was in the side of a
cone shaped depression. After a pause, the lizard quickly walked to

TIME
145
Figure 35. Gradual reentrainment of lizard emergence times after release.
Emergence times (dashed lines) and submergence times (solid
lines) of Lizards 9 (June), 11 (July), and 10 (July) are
displayed from left to right. Capture time, release times,
and overcast skies are indicated by the letters C, R, and 0,
respectively.

146
level ground. Due to the surrounding vegetation, lizards were still
commonly in the shade. For example, Lizard 11 had to walk four meters
to reach its morning basking site. Thus, Cyolura did not consistently
heat its head prior to complete emergence. Lizards were much more
likely to retreat into their burrows when scared after a recent emergence.
This inverse relationship between shyness and , first reported for
Anolis lineatopus (Rand 1964), is probably a common characteristic of
diurnal lizards.
Since the Gainesville pen in July received only 48 percent as
many minutes of direct sunlight during clear weather as Site II, lizard
activity periods were expected to be commensurately reduced. However,
large males and an occasional adult female emerged and submerged when
the pen was totally in the shade. As a result, the mean activity time
of seven adults was 468 minutes or only 17 percent less than the 564
minute mean for two males at Site II. It appears that Cyolura has an
innate minimum daily activity period as well as a maximum activity
period. When the time can be maintained within the preferred range
is reduced below the minimum activity period, the lizard operates at
a lower mean T^ rather than remaining in its burrow.
Morning Heating Phase
The typical movement pattern during emergence, basking, and the
initial minutes of the first foray is illustrated in Figure 36. The
initial blip on the upper three records corresponds to the short walk
in the Gainesville pen from the burrow mouth to the morning basking site.
Morning and late afternoon basking in the field each occurred at only
one or at most two locations, different for morning vs. afternoon

Figure 36. Lizard activity during the morning heating phase. Results
are displayed for five adult lizards confined either in the
Gainesville pen or in a field cage (bottom two graphs).
Except for brief intervals at high Tfo's, lizards were con¬
tinually in direct sunlight. A movement unit was defined
as any change in lizard location; that is, a cage climbing
bout, a feeding bout, a reorientation to the sun, or a short
walk. Two regression lines were fitted to the data, each
covering a segment of the T^ range.

MEAN MOVEMENT UNITS/I
148
Tb INTERVAL

149
basking and always located quite near the present home burrow. Lizards
which emerged in the shaded Gainesville pen moved from filtered sun
spot to sun spot without increasing their T^ significantly, resulting in
higher initial activity peaks than for the lizards illustrated in
Figure 36. Lizards settled down after arrival at a sunlit location
to a low, essentially temperature independent rate of movement. Animals
occasionally fed or engaged in social interactions for short periods
during this interval, as well as displaying intermittent thermoregulatory
behavior. Strenuous activity was sometimes elicited in the basking
iguana, as when an adult male on his first foray encroached into the
area and was chased away. Activity abruptly increased between 's of
34 and 37 C, corresponding to the start of the first foray.
The duration of the morning heating phase for similarly sized
adult males did not differ significantly from February to October.
Pooled data had a mean of 104 minutes, a range of 57-166, and a SD
of 30 (N = 46 clear days). Lizards in winter sometimes basked up to
300 minutes without even completing the heating phase or starting a
foray (Iverson 1977, November 27-December 21 field trip).
Regulating Phase
Maximum daily activity from February to October occurred during
the first foray (Figure 37). The same timing was observed in the pen,
where lizard movement could be quantified more precisely. Activity
decreased to a minimum in the early summer afternoons due to the high
surface temperatures over most areas. Bimodality nearly disappeared
during the cooler months of February and October and, as observed by
Iverson (1979), disappeared in November and December. Dominant males

Figure 37. Daily and seasonal variation in activity. Lizards were
observed for 15 days in each of three months at Site II.
A walking bout (solid line) observed from the blind (July)
or during tracking (February and October), digging out
a burrow (open triangles), climbing a bush (histograms),
chasing another lizard (dotted lines), and feeding (dashed
lines) on sardines (July only), fruits, or leaves were all
counted as one activity unit and the units summed for the
15 day periods. Total mean hourly activity units are
represented by solid lines. Observations in hours with
more than ten minutes of overcast were not included.
Separate months should not be compared quantitatively
since lizards declined in numbers between trips, supplemental
food was provided in July, and lizards were observed from a
blind in July, tracked in October and February.

HOUR INTERVAL
MEAN HOURLY ACTIVITY UNITS
o ro 4- cncoooro-^CDooooro -fccDooo
HOURLY BUSH CLIMBING, CHASING, DIGGING, AND FEEDING BOUTS
FEB

152
were territorial all year, defending an area around their home burrow
primarily by chasing intruding dominant males and smaller subdominant
males away. The three July chase peaks occurred during the first to
the third forays of the dominant male, when the probability of his
making visual contact with another lizard on a foray was greatest.
Similarly, the two chase peaks in October occurred during the first
foray and the second or third forays, respectively, and the single
February chase peak occurred while lizards were on their first or
second forays. The loss of the second and third maxima in February
was due to a decrease in the length of the activity periods as well
as declines in territorial behavior and the number of subdominant males
from five to three. The daily variation in the number of feeding bouts
corresponded well to the number of chases, declining from three peaks
in July to two in October and one in February. Maximum feeding activity
in July occurred before the mean activity and chase maxima, due to
lizards walking en masse to the sardines immediately after morning bask¬
ing. Wild Dilly fruit (MartiZkara bahamensis) profusely scattered on
the ground produced a similar response in February.
Arboreal activity was observed much more frequently at Iverson's
Pine Cay site, probably due to a number of differences at Site II; that
is, lower lizard density (an estimated maximum of 13.4 vs. 31.1 adults
per hectare), greater average size of adults, lower density of food
species, disturbance of lizards due to tracking, and, perhaps, higher
wind speed. Arboreal activity crudely mirrored chases and feeding
(Figure 37). The absence of climbing during July mornings may be due
to the supplemental sardine feeding. Iverson (1977) observed maximum

153
arboreal activity without concurrent feeding at midday in June to
September and during the morning in November and December. The former
may have been an escape behavior from hot surface microclimates and the
latter a seeking of warmer basking platforms. The mean change in
was +0.4 C (range = -0.6 to +2.0, SD = 0.8) for 11 bouts of adult arboreal
feeding in October at Site II (X time/bout = 7.5 min, range = 1-23,
SD = 7.0).
Lizards were occasionally observed digging out their burrows in
the field and the pen. Excavation almost always occurred between 1000
and 1400 during the first or second forays. The currently occupied
burrow was more commonly excavated than those presently unoccupied or
used by other lizards. An excavation by Lizard 2 in February lasted
from 1128 to 1225 and resulted in a T^ drop from 37.2 to 35.4 C.
Another bout in October by Lizard 14 lasted from 1055 to 1114 and
resulted in an overall T^ rise from 39.6 to 39.8 C. In each case the
lizard disappeared into the burrow for up to six minutes, periodically
kicking sand up to three meters out the mouth, emerging and basking
briefly, then reentering the burrow for more digging. Six submergences
occurred during both the February and October observations, averaging
3.3 and 1.2 minutes per submergence, respectively. Immediately after
laying her eggs in the cage burrow, Lizard 6 performed the same digging
behavior in the reverse direction, plugging the egg chamber with sand
scraped and kicked into the burrow from the area immediately outside
the mouth. The sand for burrow plugging may normally be obtained
entirely from inside the burrow. This was impossible in the cage due
to insufficient sand. Nest plugging began at 0930 and ended at 1339
after 29 emergences. T^ ranged from 36.6 to 39.4 C. Since nesting

154
apparently occurs before morning emergence in free-ranging females
(Iverson 1977), T^ may not normally rise much above burrow temperature.
Heavy cloud cover inhibited lizard activity. The daily activity
unit sum was negatively correlated with the percent daily overcast
time (Y = -0.67 S + 57.9, v = -0.697, N = 15 July days, p 0.005).
The effectiveness of cloud cover as an activity inhibitor varied with
its timing during the emergent lizard's day (Figure 38). The first
peak in inhibitory effectiveness occurred during the hour before maximum
hourly activity, when most lizards were just starting their first forays.
Cloud cover was less effective as an activity inhibitor during the
normal peak activity period. Maximum inhibition occurred in midafter¬
noon. Practically any short period of heavy overcast during this
period delayed and reduced the 1400-1600 activity peak. Lizards which
retreated into their burrow during heavy midafternoon overcast usually
did not reemerge until the next day, even when the skies cleared before
the normal submergence time.
Foray Movement and Body Temperature
After basking for a time near the home burrow following emergence,
iguanas rather abruptly started the first of two to four daily forays
(X = 2.4 for February plus October). Lizards walked for a few meters
at a fairly constant rate, usually moving in nearly a straight line,
frequently licking rocks, organic debris, and potential food items
between strides, then pausing for a short time in either an exposed
or shaded location before walking another few meters. Change in the
direction of travel usually occurred after a pause rather than during
a walk. Lizards paused and tilted their heads sideways upon arrival at

LINEAR CORRELATION COEFFICIENT
155
Figure 38. Variable inhibitory effectiveness of cloud cover on lizard
activity. Regression coefficients were calculated for hourly
linear correlations of percent cloud cover vs. total lizard
activity units. The sample consisted of fifteen days of
observations at Site II in July. Significant correlations
at the five percent level or higher are circled. The curve
was fitted by eye.

156
an arboreal food source, then commonly climbed the bush. A search
pattern restricted to the immediate area was initiated when a preferred
fruit was found on the ground. Forays were circular, elliptical, or
linear, usually starting and ending at the burrow utilized the previous
night. Chases and other social interaction occurred after initial
visual contact, when two lizards met during their forays or when one
lizard was on a foray and another was resting near its home burrow
during an interforay (see Iverson 1979 for details of CyoZuva caminata
social behavior).
Mean distances traveled during forays and mean rates of travel were
greater in February than October. Since the differences were not
significant, the two months of data were pooled. The greater abundance
of fruit in October may have resulted in the slightly smaller home
ranges and slightly lesser distances traveled. The mean distances
traveled during first, second, and third forays on clear or nearly
clear days were not significantly different: means = 132.9 m (436 ft),
145.1 (476), and 121.9 (400); 2 SE = 21.3 (70), 15.2 (50, and 33.2 (109),
respectively. The mean rate of travel declined from first to third
forays, but not significantly: means = 1.41 m/min (4.63 ft./min),
1.24 (4.08), and 1.15 (3.77), 2 SE = 0.30 (0.97), 0.19 (0.63), and
0.61 (1.99) respectively. With the sun obscured by clouds 0-20
percent of the time, the overall mean rate of travel was 1.25 (4.10)
[N = 32, SD = 0.51 (1.68), range = 0.36-2.54 (1.77-8.33)]. Iguana
iguana weighing slightly less than the Cyclura aavinata tracked in
the present study could walk at a maximum rate of approximately 3.4
m/min (2^ = 30 C) and 4.0 m/min (T^ = 38 C, 1.8 C above mean operant
T-I) without incurring a significant oxygen debt (Moberly 1968). Thus,

157
walking in Cyalura was probably an aerobic activity, with little or
no build-up of lactic acid, leaving a margin of safety for more
strenuous activities; such as, bush climbing, chasing, burrow digging,
and infreqently, fighting.
Foray length was not depressed until overcast time reached 45
percent or more of total foray time (Figure 3S). High surface tempera¬
tures during perfectly clear days also depressed the maximum foray
distance attained. This effect was probably greater during the summer.
The mean rate of travel was inversely correlated with overcast time.
Lizards moved to a warm exposed substrate and sat quietly in a prostrate
posture during overcast. Since this nonwalking time increased as cloud
cover time increased, the rate of travel decreased.
Both foray length and the mean rate of travel were independent
of mean body temperature between 35.0 and 40.5 C (Figure 40). However,
no foray occurred outside this mean T-. range. Also, when overcast time
was 30 percent or less, mean was inversely correlated with the mean
rate of travel. Lizards had to spend more time sitting in the shade
avoiding overheating when their mean T- ’s exceeded the preferred range
during clear and nearly clear days, thus decreasing the mean rate of
travel.
Mean T-, was significantly lower at the start of the first foray
than at the start of subsequent forays (Figure 41). Also, the mean
T-, range during a foray declined from Foray I to III (February: I;
X = 5.3 C, SD = 2.6, II; X = 1.6, SD = 0.8, III; X = 0.9, SD = 0.6.
October: I; X = 4.0, SD = 2.5, II; X = 1.7, SD = 1.0, III; X = 1.5 C,
SD = 1.2). Thus, lizards were more eurythermal during the first foray
than during subsequent forays. Mean T^ at the start of the first foray

Figure 39. Influence of cloud cover on the length of lizard forays and
the mean rate of travel during forays. Data for February
and October tracking of large male Cyalura have been pooled
(N = 6, X weight = 1711 g, range = 1507-1864). The mean
rate of travel equaled the total distance walked during a
foray divided by the total time required. Solid and dashed
lines are linear regressions for 0-45 and 0-58 percent over¬
cast times, respectively. Neither correlation in the upper
graph was significant (solid line; r = -0.035, dashed line;
r = -0.111, p > 0.05) while both correlations in the lower
graph were significant (solid lines; r = -0.292, p < 0.05,
dashed line; r = -0.494, p < 0.0005).

RCENT OF TIME SUN OBSCURED BY CLOUDS
MEAN RATE OF TRAVEL (M.FT/MIN)
o _ _ no rv,
O en O Oi O cn
r- "i 1 1 1 r~
001 i
LENGTH OF FORAY (M,FT)
04
o
60
90
120
150
IV)
04
cn
o
o
o
O
o
o
o
O
CD
O
ro
O
ur
o
o
- -
4
• •
*
Ui
kO
oozl

Figure 40. Correlations of foray length and rate of travel with mean
body temperature. The percent overcast times are represented
by solid circles (zero percent), open circles (1.0-9.1), open
triangles (10.7-27.5), and open rectangles (31.2-58.0).
Solid lines are linear regressions for all the data points
(upper graph; r = -0.032, p > 0.05, lower graph; r = -0.062,
p > 0.05). The dashed line is the regression for 0-30 per¬
cent overcast time (r = -0.370, p < 0.01).

MEAN RATE OF TRAVEL (M,FT/MIN) LENGTH OF FORAY (M.FT')
161

Figure 41. Body temperatures during forays. Data are illustrated for
two lizards monitored in both February and October. Arabic
numbers below the abscissa refer to Tfo's at the start
(1) and end (2) of Forays I, II, and III and minimum (3)
and maximum Tj-,'s (4) during the forays. Diagonal lines
connect mean Tfo's, vertical lines are ranges, and rectangles
are plus and minus two standard errors for N > 4 days.
Horizontal lines are the lizard's minimum and maximum
voluntary tolerances during the regulating phase.

163
43
41
39
37
35
£33
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a: 3!
§29
ÃœJ
ü-
2 43
ÜJ^°
I-
>-4i
a
o
“39
37 h
35
33
31
29
LIZARD 2
FEB OCT
LIZARD 3
OCT
1 1 1 L_J
12 3 4
2 3 4
< â–  ' â– 
12 3 4
12 3 4
l i J 1—
12 3 4
12 3 4
I i
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Tb’S DURING FORAY AND FORAY NUMBER

164
in February was usually even below the minimum voluntary tolerance of the
regulating phase. Lizards may become more eurythermal during the first
foray during the time of maximum seasonal activity. Iverson (1977)
observed that Cy atura was most active in March and April, only one month
after my February monitoring.
T^ usually both increased and decreased during a foray rather
than steadily increased or decreased. The T^ at the start of the foray
was lower than the maximum T^ during the foray in 91.4 percent of total
forays and was higher than minimum Th during the foray in 75 percent of
second plus third forays. Except during the opening minutes of the
first foray, T\ was usually near the middle of the regulating phase
range. Thus, lizards had upper and lower thermal buffers to accommodate
less frequent maximal T^ changes, for example, an unavoidable T^ rise
during feeding atop a bush or an unavoidable T\ fall during a long
overcast period.
No heating nor cooling trend was evident when 's at the start
of interforays were compared to those at the start of the next foray.
Thus, lizards didn't consistently return to the home burrow to heat
or cool before the next foray. The interforay was probably important
in reorienting the lizard in its home range prior to further walking.
Cyalura is apparently capable of some seasonal acclimatization of
its metabolic rate. Activity in the thermal gradient, as measured by
the number of shuttling movements and the daily distance walked, was
greater in lizards acclimatized to winter than to summer weather in
Gainesville (Table 16). The difference between samples was greatest
during the second day in the gradient and had decreased considerably
by the fourth day. Except for the second day, the magnitudes of the
differences in shuttling movements were greater than in the distance walked.

Table 16. Possible acclimatization of lizard activity. Lizards were kept in outdoor pens in
Gainesville, then monitored in the thermal gradient for four days. The two samples
each consisted of two males and two females matched for approximately equal size and
weight. Air temperature and photoperiod were held constant. The mean Tfo's for the
two samples did not differ significantly. A one-sided t test was used to test for
differences in the number of daily shuttling movements and the daily distance walked.
DAILY SHUTTLING MOVEMENTS DAILY DISTANCE WALKED(M)
DAY
JULY
X(l) SD
DEC-
X(2)
-JAN
SD
U) mo
(2) ' 100
P
JULY
X(l) SD
DEC-JAN
X(2) SD
• 100
(2) UU
P
1
5.5
3.3
1] .5
14.8
47.8
NS
3.23
1.19
2.71
2.35
119.1
NS
2
9.5
5.4
27.8
15.2
34.2
0.05
3.29
1.52
11.06
6.83
29.8
0.05
3
19.8
6.8
39.5
26.1
50.1
NS
10.70
4.82
16.00
7.13
66.9
NS
4
27.8
8.6
34.7
4.1
80.1
NS
17.25
7.71
20.06
1.16
86.0
NS
1-4
15.6
10.6
28.4
18.7
54.9
0.025
8.63
7.35
12.47
8.20
69.2
NS

166
Feeding, Transmitter Passage Time, and Defecation
Lizards occasionally fed briefly on the ground near their home
burrows before basking or starting their first forays. The mean T-, for
feeding on the ground in February plus October was 36.7 C (N = 14, range
30.0-40.4, SD = 2.42). However, arboreal feeding did not begin until
the first foray, at a significantly higher mean T^ than ground feeding
(N = 13, X = 38.6 C, range = 35.7-40.1, SD = 1.39, p _< 0.01, one-sided
¿-test). Also, both the feeding rate and the amount of food consumed
in the Gainesville pen were significantly less immediately after
emergence (Table 17). The mean feeding rate was 50-52 percent less
and the mean amount consumed was 38-43 percent less than during the
regulating phase. Yet, one recently emerged lizard consumed white
grapes equivalent to 8.4 percent of its body weight. Clearly, the 2V
range for feeding was quite broad, approximately 23.6 C to probably
greater than 40.4 C. One advantage of low T- feeding is the capacity
to quickly exploit new food sources on the ground near the home
burrow; for example, fruits which fall during the night, before other
lizards already on their first forays discover the source.
The time required for passage of transmitters through the digestive
tract varied from 2.8 days for a far-field transmitter in a free-ranging
lizard in October to 66 days (2 Jan-8 Mar) for a near-field transmitter
in a lizard in the Gainesville pen. Passage times increased during the
Gainesville winter, probably due to both the. cessation of feeding and
low 2^’s. Passage times for free-ranging lizards at Site II in October
varied considerably (2.8, 4.8, 7.0, and 11.0 days for far-field trans¬
mitters; more than 13 days for a near-field transmitter). Although

Table 17.
Effect of body temperature on the feeding rate and meal size of penned
Cyclura aarinata. Lizards were provided with white grapes ad libitum
as a food source. Data were collected in July and August in Gainesville.
TIME OF
FEEDING
BOUT
APPROXIMATE
BODY
TEMPERATURE
RANGE
N
(LIZARDS, FEEDING
BOUTS)
IA1 0850-1149
23.6-29.2
Ln
CO
IB2 0850-1439
23.6-40.3
5,23
II1 1150-1300
34.8-40.3
4,11
III1 1458-1555
34.8-40.3
5,13
RATE OF FEEDING MEAL SIZE
(X GRAPES PER MIN) (PERCENT BODY WEIGHT)
X
SD
RANGE
X
SD
RANGE
i.i3
0.6
0.4-2.0
4.04
2.5
0.8-8.4
-
r-.
6.8
2.6
4.2-11.7
2. 3
1.1
1.1-5.0
7.0
0.9
5.7-8.3
2.2
1.1
1.0-5.1
6.4
1.0
3.9-7.6
Grapes consumed in a single feeding bout.
"Grapes consumed during the full day starting with the feeding bout in IA.
^Mean significantly lower than II and III (p <_ 0.01) by a one-sided f-test.
+Mean significantly lower than IB (p < 0.025) and II and III (p < 0.005).

168
feeding was undoubtedly reduced during tracking, transmitters also
probably became lodged for variable times between the septa of the
voluminous proximal colon. See Iverson (1980) for a description of
Cyclura's digestive tract.
Lizards most commonly defecated near the morning basking site
during the regulating phase, on the average, 87 minutes after the first
T- maximum (SD = 104, N = 46, Figure 42). This was only 31.0 percent
of the average time between the first and last maximum T^'s (292.1
minutes). Large solid objects, for example, far-field transmitters
and hermit crab appendages, may be expulsed later in the day, on the
average, than leaves and small seeds. Almost all defecations occurred
at T-^'s between 33.6 and 40.4 C (X = 37.6, SD = 2.17, N = 46). Even
though fecal material was not found in lizard burrows, captive individ¬
uals occasionally defecated when handled at T^'s as low as 25 C. The
defecation at a L of 27.0 C was by an untouched lizard which saw me
while it was confined in the rooftop arena.
Drinking
Drinking was an infrequent behavior in Cyclura, even in captivity,
where water was readily available. Drinking in free-ranging lizards
was observed only once, in the dry month of February. An adult male
licked a few drops of rainwater from the surface of dead sea grape
leaves (Coocoloba uvifera) lying on the ground. Three of four caged
lizards in June didn't drink until five, six, and seven days after
capture and then only infrequently. The fourth did not drink at
all during 12 days in captivity. Lizard 6 drank five times within an
hour and a half, starting immediately after completion of egg laying

169
L_J i i I I i ' ! i i i i i 1 I i I 1 J I I I ! J
-80 -40 0 4 0 80 120 160 200 240 280 320 360
T!ME (MIN) BETWEEN FiRST MAXIMUM Tb AND DEFECATION
Figure 42. Defecation body temperatures and timing. Lizards defecated in
the arena atop the zoology building (solid squares), field
cages (solid triangles), field (free-ranging lizards defecat¬
ing far-field transmitters; open circles), Gainesville pen (open
triangles), and thermal gradients (solid circles). Means, plus
and minus one standard deviation, and ranges for defecation
timing and are represented beside the abscissa and ordinate,
respectively.

170
in the field cage. She drank only twice during the previous twelve
days. Clearly, body water is normally primarily obtained from food
rather than from free-standing sources. Drinking occurred at a mean
2^ of 38.2 C (33.7-41.3, N = 15) in the June field cages.
Nasal Salt Gland Secretion
Nasal salt glands have been studied physiologically in members of
six of nine iguanine genera: Amblyrhynohus3 Conolophus, Ctenosaura,
iKpsosccuruSj Icpuana, and Sauromalus (see Dunson 1976 for a review) and
may in fact occur in all iguanines. Dissection revealed the gland's
gross morphology to be similar to other iguanines. Cyclura could often
be located in the field by listening for the sneezing sound made when the
animals cleared accumulated gland secretion from their nostrils. A large
male could propel secretion droplets up to a meter into the air and two
meters to the side with a quick partial exhalation and somewhat semi¬
circular movement of its head. Iguanas usually sneezed while sitting
still, most commonly just prior to walking. Marine iguanas also
usually sneeze just prior to moving, as before changing basking positions
or before entering the sea after morning basking (Carpenter 1966). The
correlations may be anticipatory of an increased respiratory rate
during movement. The hourly sneezing rates of two large males tracked
in February were fairly constant throughout the day (N = 10 days, X =
2.85 sneezes/hr, ranges of daily X's = 1.98-3.82, SD = 0.73; range of
sneezes/hr - 1-6). Actual rates were undoubtedly slightly higher, due
to brief discontinuities in tracking contact and the limitations of human
hearing in windy locations. Sneezing was restricted primarily to regulat¬
ing phase 21 's (X T= 37.7 C, range = 29.1-40.9, SD = 1.66, N = 144
sneezes).

171
Time-Motion Study of Penned Lizards
Iguanas in the Gainesville pen spent most of their summer days
just sitting. Active time averaged 1.8 percent of emergent time (Table
18). A log-log plot of mean daily distance traveled for individual
lizards vs. confinement area was linear, with a significantly positive
slope (Figure 43). Thus, active time was probably greater in the field.
Percent active time in the field can be estimated using the following
equation, assuming equal rates of movement in the field and pen:
Af ‘ AP ' (Df 'DP'l) <5)
where A_? and A are mean percent active times in the field and pen,
J P
respectively, and D ~ and D1 are the mean daily distances traveled.
The estimated A~, 7.5 percent of emergent time, is about equal to
the 3 percent maximum for the small (X = 240 g) herbivorous lizard,
Egemia aunn-inghami (Wilson and Lee 1974).
Penned iguanas spent a higher percentage of their active time in
thermoregulatory movement than free-ranging Egevni-a (43 vs. 12 percent).
This was probably due to Cyatura's higher mean operant T^ (37.5 vs.
32.9 C), coupled with the small amount of time direct sunlight was
available in the pen, as well as the ease of obtaining food in the pen
(an average of 41 vs. 85 percent of active time in pursuit of food).
Thermoregulatory movements in the pen, consisting of shuttling walks
and sun orientations, took less time per movement and occurred at higher
rates per hour than either feeding or social behavior. Comparison of
individual lizard results is not justified since both weather and
activity periods varied considerably from day to day. However, as
expected, the mean number of hourly thermoregulatory movements does
appear to be inversely correlated with body weight.

172
Table 18. Time-motion study of CyoZura caminata in the Gaines¬
ville pen. Five lizards were observed for one day
each in July. A movement unit is any complete
behavioral sequence, such as a feeding bout, a chase,
or a walk to the shade.
Lizard
1
2
3
4
5
Weight (G)- Sex
1260-M
1240-M
1195-M
600-F
129
X
Emergent Time (Min)
473
281
414
348
188
Fercent Pen Shade Time
45
47
37
33
5
Percent Overcast Time
25
15
40
9
59
Mean Sun Exposed Sand Temperature
40.7
31.4
31.7
38.0
42.2
TOTAL ACTIVITY
Active Time (Min)
10.6
2.6
7.2
6.8
4.0
6.2
Percent Emergent Time
2.2
0.9
1.8
2.0
2.1
1.8
Movement Units
87
49
70
108
101
S3
X Hourlv Movements
11.0
10.5
10.1
18.6
32.2
16.5
FEEDING
Total Time (Min)
5.4
0.9
3.7
2.0
1.6
2.7
Percent Emergent Time
1.2
0.3
0.9
0.6
0.9
0.8
Percent Active Time
51
33
51
30
41
41
Total Movement Units
29
8
17
20
8
16
X Hourly Movements
3.7
1.7
2.5
3.4
2.6
2.8
X Seconds/Movement
11.2
6.4
12.9
6.0
12.2
9.7
SOCIAL BEHAVIOR
Total Time (Min)
3.2
0.2
0.5
1.8
0.3
1.2
Percent Emergent Time
0.7
0.1
0.1
0.5
0.2
0.3
Percent Active Time
31
8
6
26
8
16
Total Movement Units
15
5
3
17
15
11
X Hourly Movements
1.9
1.1
0.4
2.9
4.8
2.2
X Seconds/Movement
13.0
2.4
9.3
6.2
1.2
6.4
THERMOREGULATORY
BEHAVIOR
Total Time (Min)
1.9
1.5
3.1
3.0
2.0
2.3
Percent Emergent Time
0.4
0.6
0.8
0.8
0.9
0.7
Percent Active Time
18
59
43
44
51
43
Total Movement Units
43
36
50
71
78
56
X Hourly Movements
5.5
7.7
7.2
12.2
24.9
11.5
X Seconds/Movement
2.6
2.6
3.7
2.5
1.6
2.6

MEAN DAILY DISTANCE TRAVELED (M)
Figure 43. Correlation of lizard mobility with confinement area. The mean daily distance traveled
was measured for adult lizards in the thermal gradient (N = 4 lizards, Dec-Jan),
Gainesville pen (N = 5, July-Aug), and Field Site II (N = 4, Oct). Confinement area
in the field was defined as the mean home range size. The linear regression equation
fitted to the data was; Log Y = 0.297 Log X + 1.398 (N = 22, v = 0.957, p j< 0.0Ü05).
173

174
Behavioral Thermoregulation
Description of Thermoregulatory Movements and Postures
When a lizard assumes a characteristic posture or moves its body-
in a consistent way in a specific microclimate, resulting in a predict¬
able decrease, increase, or stabilization of peripheral or core T^, the
behavior is probably thermoregulatory. However, care must be taken
when assigning a single feeding, social, or thermoregulatory function
to a behavior, since multiple functions are likely. For example, an
arboreally feeding lizard may originally have climbed a bush to avoid
a hot surface microclimate. Possible nonthermoregulatory functions of
thermoregulatory behaviors will be mentioned where appropriate in the
following descriptions. Numbers in parentheses refer to illustrations
in Figure. 44.
Pre (1) and Partial (2) Emergence Postures. The preemergence pause
occurred as the lizard rested in sight of the burrow mouth, usually just
inside the lip. The pause was a very consistent behavior, never omitted
in the Gainesville pen. Lizards secured information about the outside
world without exposing themselves to unfavorable weather or possible
danger. Emergence was delayed by overcast skies, presumably due to low
light intensity inhibition, and by unfamiliar noises such as a human
voice. Partial emergence permitted a visual scan of the immediate
burrow area, again with minimal danger to the lizard. The behavior
is analogous to partial emergence from the water prior to basking in
semiaquatic turtles. However, no unsolicited head rotations were noted,
as have been observed in Chyrsemys scripta (Auth 1975). When frightened,
lizards either turned quickly around and retreated head first into the

Figure 44. Thermoregulatory motions and postures of Cyolupci carinatci. Illustrations were made from
photographs taken in the field (3, 8, 9, 10; free-ranging lizards, 14; captive lizard),
Gainesville pen (1,2,4,11,12,13), and laboratory temperature control chamber (5, 6, 7).
175

176

177
burrow or backed in tail first. Head heating may occasionally be
significant, for example, during a slow emergence at low T^. Sand
burrow mouths were located below the ground surface in the side of an
inverted cone. Due to greater indirect radiation from the sides of the
cone, higher substrate temperature, and lower wind velocity, lizards
emerging into the cone, especially near midday, probably heated faster
than animals moving immediately to level ground. However, since burrow
mouths were often located in the shade, initial head heating was not a
consistent populational requirement.
Upright Posture (3,5). The forelegs were adducted toward the
thorax and variably extended, keeping the ventral pectoral region and
abdomen off the substrate, as far back as the pelvic girdle. The fore¬
legs were straight in a maximally upright animal, and the head was higher
off the substrate than illustrated in (3). The hindlegs sprawled
laterally. The tail was flat on the ground except when curved directly
upward in an apparent appeasement display by females and subdominant
males. Lizards assumed the upright posture more frequently than any
other. It was observed consistently during the morning heating phase,
commonly during pauses between foray walks, and commonly during summer
middays when lizards sought out areas exposed to the easterly wind.
Lizards appeared to be maximally alert, rarely closing their eyes for
longer than a second. They assumed a maximally upright posture when
hearing an unusual noise, seeing an unfamiliar object moving into the
field of view, or looking over an obstruction. Substrate contact also
varied with air, body, and substrate temperatures (see Thigmothermia).
Tor example, the lizard in (5) minimized contact with a cold photo-
thigmotron substrate while in warm air (2V = 30.2 C, T =21, T =37.6).
O S CL

178
Prostrate Posture (4,6). Maximum relaxation on a flat substrate
was characterized by lateral expansion and dorso-ventral compression of
the trunk. When viewed from the side, the midsagittal crest was
slightly concave to slightly convex. Undisturbed lizards occasionally
relaxed their heads on the substrate. Lizards were less alert than
in the upright posture, as indicated by intermittent eye closure.
The position of the legs varied with radiant input, substrate tempera¬
ture, and probably, wind velocity. For example, the lizard in (4)
was cooling slowly in the late afternoon shade while relaxing on a
concrete block. Substrate temperature averaged 2 C less than the
lizard’s core, which was in the regulating range. Contact of the
legs with the trunk and tail minimized conductive and radiative heat
loss. In contrast, the legs of the lizard in (6) extended out from
the body and made maximum contact with the substrate, since substrate
temperature in the photothigmotron was higher than both the still air
and the body core (T^ = 41, T^ = 29, T^ = 36.2 C). The prostrate
posture was assumed during overcast following clear skies, with
Tu in the regulating range and stable or slowly decreasing, and
during the late afternoon in diminishing sunlight, with commonly
at the highest point during the day. The posture was also commonly
assumed by subdominant males when approached by a dominant male, a
sign of submission which probably reduced the probability of a
chase.
Positive Normal Orientation (7). The lizard maximized the
body surface area exposed to a radiant heat source by orienting

179
perpendicularly (normal orientation) and curving its body slightly toward
the source, in this case, a heat lamp in the photothigmotron. The angle of
incidence of the radiation was normalized by tilting the sun exposed side
toward the heat lamp (positive orientation). The right side of the lizard's
trunk in (7) constituted 67.1 percent of the total trunk photographic area.
Object Utilization During Sun Orientation. Lizards maximized
radiant input by facing away from the sun while propped up anteriorly on a
rock or some other object (positive parallel orientation). This behavior
was fairly common in the early morning during the cooler months of the year. It
was rarely observed during the summer; when lizards usually assumed an up¬
right posture during morning basking, on level ground and approximately
perpendicular to the sun'srays. In the late afternoon at Site II, lizards
commonly sat on the sides of hills which faced the sun. Minimizing radiant
heat input by orienting negatively parallel to the sun was rarely observed
in the field, probably due to the large areas of shade present at both Sites
I and II.
Heath's terminology (1965) describing lizard sun orientation has been used
in the present paper. It is based on whether the radiated dorsal body
surface of the lizard faces toward (positive) or away from (negative) the
sun and whether the long axis of the lizard's body is perpendicular
(normal) or parallel to the incoming parallel rays of the sun. Brattstrom's
(1971) and Greenberg's (1977) schemes were rejected due to their use of
positive and negative orientations to describe how the lizard's head is
oriented with respect to the sun, which seems less logical than using the
terms in relation to the dorsal body surface. Brattstrom also confuses
body posture (the position of the lizard's head, trunk, and tail with
respect to the substrate and the position of the limbs with respect to
the trunk) with body orientation to the sun; for example, in his "extreme
positive orientation." Greenberg's scheme was also rejected because he
had to invent a second set of terms to describe the positive normal
orientation. Muth's scheme (1977b), quite different from the three above,
is the most mathematically precise, taking into account all intermediate
orientations. It is admirably suited to computer simulations but was
not adopted for the present study since solar zenith angles were not
measured and most of the literature descriptions also lack the required
quantification for conversion to his terminology.

180
Arboreal Climbing (9). This behavior was probably thermoregulatory
in Cyotuva, removing the lizard from excessive heat inputs at ground
level in the summer and from low ground surface temperatures in the
winter. However, the mean T^ change of arboreally feeding lizards at
Site II in October was not significantly positive. Unfortunately, I
have no June-July nor December-January data. Iverson (1977) observed
considerable winter arboreal climbing without feeding at his Pine Cay-
field site. Lizards sat quietly on branches in the sun.
Walking (10) and Running. Cyotura moved at essentially three
speeds; a slow walk, a medium speed trot or slow run, and a fast run.
Walking and slow running correspond to Type A locomotion and fast
running to Type B locomotion in Uromastix aegyptius (Dmi'el and Rappeport
1976). Walking was the most common locomotory mode used during forays.
A considerable length of the distal tail made contact with the sub¬
strate, and the ventral trunk was minimally elevated above the ground.
Walking was restricted to substrates with surface temperatures below
approximately 45 C. Slow running occurred when I approached lizards
too closely while tracking, when a female or subdominant male increased
its distance from an approaching dominant male, and when any lizard
crossed large exposed areas with surface temperatures in excess of
45 C. Fast running, which was always quadrupedal, occurred during
territorial chases and escapes from potential predators. The tail
extended horizontally off the ground, and the ventral body surface was
maximally elevated.
Semiprostrate Posture (11). Body contact with the substrate
extended from the pectoral girdle nearly to the tip of the tail, with
considerable lateral expansion of the trunk. The calves were directed

181
diagonally toward the tail (3) or sprawled laterally (11) , especially
when the substrate sloped downward laterally. Judging from results
in the Gainesville pen, lizards assumed a semiprostrate posture during
the morning basking phase only when substrate temperature exceeded core
body temperature. The posture was also assumed during the regulating
phase in either the sun or the shade. However, substrate temperature
was neither consistently higher nor lower than body temperature.
Shade Seeking (12). Lizards sought out shaded microclimates when
body temperatures extended into the maximum body temperature range.
Brief physical contact was made between two or three iguanas during
shade seeking in the Gainesville pen, a behavior never observed in the
field in a thermoregulatory context, where lizard density was much lower
and shade was less limited. In (12) a subdominant adult female had
crawled onto the back of an adult male. Soon after the photograph
was taken the male left the shade to the female by sliding forward and
walking away. The subdominant usually licked the dominant once or
twice on its dorsal surface before mounting. Subdominants moved away
from dominants before body contact could be made.
Lizards sitting partially in the shade during midday hours
eventually moved exposed portions of their bodies into the shade; for
example, curved their tails or their heads, as the male in (12).
A little less than half the Gainesville pen was continually
shaded. It was infrequently traversed during daily lizard movement.
When rock piles and shelters were rearranged, activity considerably
increased for a day or two. Lizards walked further and explored the
entire pen. New areas were subsequently excluded from daily travel.

182
Captives undoubtedly became familiar with the new locations of thermally
suitable microclimates during this exploratory phase.
Elevated Posture (13). This posture was characteristic of heat
stressed lizards and was usually accompanied by intermittent or con¬
tinuous panting. Legs were extended, maximally raising the trunk off
the hot substrate. The trunk was flattened ventrally via lateral
expansion, possibly reducing heating by radiation reflected off the
ground (Norris 1967). Elevation also enhanced forced convective
cooling and moved the lizard into a zone of lower air temperature.
Heat stressed lizards were never seen in the field; the 1900 g dominant
male in (13) would probably never reach the panting setpoint as a
free-ranging animal.
Feeding lizards on the ground often assumed a mildly elevated
posture, perhaps aiding in swallowing. Adult males elevated their
trunks during territorial face-offs. The trunk was laterally compressed
rather than laterally expanded as during panting.
Panting (13,14). The mouth was not opened maximally during
panting. The tongue was elevated, engourged with blood, and extended
to the edge of the mandible. The eyes bulged perceptibly. Air flow
over the tongue and roof of the mouth of lizards staked out in the
summer sun resulted in sufficient evaporative cooling to decrease the
heating rate of the large intestine as well as the head.
Substrate Penetration with the Legs. This behavior was observed
only once, in an adult caged male given no access to shade. While in
a walking position, the lizard repeatedly pushed forward with its rear
legs while extending its forelegs into the cool, loose subsoil (2^ =
38.0, T at the surface = 44.0 C). As body temperature fell to between
s

133
36.9 and 36.1 C during the following overcast period, the behavior was
repeated several times, presumably to reach the now warm subsoil.
Sand Swimming. When crossing hot, loosely arranged sands, such
as on bulldozed roads, lizards occasionally crawled forward (for short
distances) with maximum ventral substrate contact, using both fore and
hind legs for locomotion. Presumably, this behavior maximized body and
limb contact with cooler subsurface sand.
Reflexion of the Digits. While pausing on hot exposed substrates
in either the upright, semiprostrate, or walking postures, lizards
commonly reflexed their digits soon after stopping, making contact
with the substrate with only palmer and plantar surfaces of the feet.
Lizards soon moved to the shade rather than elevating their feet
completely off the substrate.
Substrate Rubbing and Touching. Some of the following behaviors
may have a temperature sensing function: 1. sand swimming. 2. ventral
body surface rubbing. Lizards rubbed their ventral trunks on rocks or
other hard objects, sometimes dragging themselves short distances
forward using their forelegs while extending their rear legs posteriorly.
The behavior was most commonly observed just prior to a walk during a
foray and just before evening submergence. 3. chin rubbing. The sides
of the jaw were alternately rubbed on rocks or branches, most commonly
immediately before walking after a lengthy pause. 4. substrate licking.
Besides its olfactory and gustatory functions, licking may convey
surface temperature information. Lizards commonly licked a site before
sitting on it, especially in the Gainesville pen. The presence of both
cold and heat receptors in the tongue of the domestic cat has been

184
demonstrated by recording spike potentials of the lingual nerve (see
Zotterman 1953 for a review).
Shuttling Behavior
The four caged adults observed in the field in June had mean
shuttling movement rates of 2.31, 2.43, 3.04, and 4.80 per hour (Lizards
6, 4, 7, and 5; body weights of 815, 645, 1290, and 723 g, respectively).
Lizard 5 shuttled at a significantly higher rate than the other three
animals. Since rates were not significantly different in the simple
and complex habitats, all data for each lizard were pooled. Shuttling
rate was inversely related to the standard deviation of mean body
temperature (SD = -0.148 Shuttling Rate + 1.59, r = -0.394, p £ 0.05).
Thus, increased Tstability required more shuttling and, presumably,
a higher energy expenditure. The graphs in the upper part of Figure
45 illustrate a day's shuttling at about the average rate and at the
maximum observed rate.
The time caged lizards spent in direct sunlight, vegetational
shade, and burrows varied with the hour of the day (lower graph, Figure
45). Summer peaks of direct sunlight exposure at 0800-0900 and 1600-
1700 bracketed the free-ranging lizard activity peaks at 0900-1100 and
1400-1600 (Figure 37). Direct sunlight peaks moved closer to one
another as winter approached, with a single midday peak in the coldest
months of December and January. The degree of heliothermy, as measured
by the percent sun exposure time of a lizard on a clear day, ranged
from 27.7 to 65.7 percent (N = 21 June days, X = 42.5, SD = 9.49) and
was directly related to mean body temperature (X T^ = 0.086 Percent
Emergent Time in the sun + 35.58, r - 0.406, p < 0.05). Only 18.9

6 7 8 9 10 II 12 13 14 15 16 17 18 19
TIME
Figure 45. Lizard shuttling in the summer field cages. In the upper graph, movements of an average
shuttler (Lizard 7 on June 26, X = 2.6 shuttles/hr) and a fast shuttler (Lizard 5 on
June 27, X = 7.2 shuttles/hr) between sun exposed, vegetat.tonally shaded, and burrow
portions of the cage are illustrated The mean percentages of each hour spent in the sun
(dotted line), vegetational shade (dashed line), and burrow shade (solid line) are
illustrated in the lower graph (for four lizards each monitored for three days).
185

186
percent of the daily direct sunlight time was distributed between 1000
and 1500. During the hottest part of the day, lizards selected burrow
shade significantly more of the time than the warmer vegetational shade
(1300-1359, p <_ 0.025; 1400-1459, p 0.05, one-sided ¿-test). Air
temperature between 1200 and 1600 averaged 29.5 C (28.0-30.6) in the
burrow and 34.9 C (32.9-37.5) in the vegetational shade.
Pianka (1969) found a direct relationship in fourteen species of
Ctenotid skinks between the ratio of hindleg length to snout-vent length
(HLL/SVL) and the percentage of the time lizards were first sighted in
the open (PTSO). The HLL/SVL ratio decreased ontogenetically in Cyclura.
oavinata (HLL/SVL = -0.00048 SVL + 0.7206, N = 26 lizards, HLL/SVL
range = 0.70 - 0.54 SVL range = 89 to 332 mm, v = -0.804, p < 0.0005).
If Cyclura fits the Ctenotus pattern, its ontogenetic range of HLL/SVL,
0.70 to 0.54, corresponds to a PTSO of 84 to 33 percent (HLL/SVL = 0.0031
PTSO + 0.438, N = 14 species of Ctenotus, v = 0.769, p <_ 0.005).
Unfortunately, this was not checked in the field. However, the 42.5
percent mean time in the sun for the June caged Cyclura corresponds to
a HLL/SVL ratio of 0.57 using the Ctenotus equation. The actual mean
SVL for the four lizards, 275 mm, corresponds to a HLL/SVL ratio of
0.59, about equal to 0.57. The range of adult CyeXuxa HLL/SVL ratios
places the species in the open area forager category of Ctenotus (6 of
14 species).
Free-ranging lizards utilized the wind as well as the shade to
cool themselves during the hottest days. For example, Lizard 3, on
returning from a foray at midday or early afternoon at Site II in
February and October, often walked to a wind exposed ridge near its
home burrow and sat in the shade in an upright posture. stabilized

187
or declined during this period. The lizard did not orient its body
in any consistent direction with respect to the wind.
Core at shade seeking varied in a regular fashion during the
day (Figure 46). Each of the four caged lizards moved into the shade
at a low hourly mean T^ during the hottest part of the day. The daily
ranges of hourly mean 2^'s for shade seeking were 2.4, 1.7, 1.6, and
1.5 C (X = 1.8 C) . The corresponding depressions in the mean ¡Z^'s
for sun seeking were about the same: 2.9, 1.5, 1.5, and 1.3 C,
respectively (X = 1.8 C) and also maximized between 1200 and 1300.
Sun Orientation
Direct radiative heat input from the sun increases as a lizard's
shadow area increases (Heath 1965). With a single heat lamp directed
at 45° below the horizontal onto the photothigmotron substrate, an
iguana could maximize shadow area by assuming an approximately normal
orientation and tilting the closest side of its trunk toward the heat
lamp (positive normal orientation) or facing directly away from the lamp
and maximizing forebody elevation (positive parallel orientation).
Radiant input could be minimized by assuming a normal orientation and
tilting the trunk away from the heat lamp (negative normal orientation)
or by facing directly toward the lamp and maximizing forebody elevation
(negative parallel orientation). Of course, lizards could also change
heat input by moving closer to or farther away from the heat lamp.
Cyalura maximized its shadow area in and below the T\ range for morning
emergence (Figure 47). The greatest shadow area was recorded at the
lowest substrate temperature (the lizard was approximately the same
distance from the heat lamp during the early stage of heating at all

AT ENTERING SHADE OR BURROW
43
Figure 46. Midday depression of shade seeking body temperature. Tfc's have been averaged for hourly inter¬
vals over five day periods for each of four caged lizards monitored in the field in June. Only
which were followed by intervals of five minutes or longer in burrow or vegetational shade
have been used to calculate the means. Horizontal lines are means, vertical lines; ranges, and
rectangles; plus and minus two standard errors of the mean.
188

189
BODY TEMPERATURE
Figure 47. Lizard orientation to a heat lamp during heating and cooling in
the photothigmotron. Three records for a 985 g female are
illustrated, one each at substrate temperatures of 20 C (circles),
31 C (squares), and 38 C (triangles). Air temperature was
constant at 19 C during the three runs.

190
substrate temperatures). The iguana either abruptly or gradually
reduced its shadow area as T^ increased. The greatest decrease in area
per °C rise in T^ occurred as the mean preferred T^ was approached
(37.0 C). During the experimental run at a substrate temperature of
38 C, the lizard walked away from the heat lamp when T, reached 39.1 C,
maximizing its shadow area again as 1^ declined to 36.5 C at the greater
heat lamp distance.
The occurrence and degree of positive and negative tilting
orientations depended on body and substrate temperatures. The lizard
in Figure 48 tilted positively while heating radiatively on the cold
substrate (20 C) and negatively while receiving heat inputs from both
the substrate (31 and 38 C) and the heat lamp. Tilting essentially
ceased when the lamp was turned off. Curving the trunk toward (Figure
44, Illustration 7) or away from the heat source accompanied positive
and negative tilting orientations, respectively. Curving also commonly
occurred during parallel orientation.
Blindfolded iguanas neither tilted their trunks nor rotated their
bodies during heating in the sun. Lizard contact with the substrate
did increase with increasing Thowever. Sun orientation appears to
be mediated through the eyes rather than through temperature receptors
heating at different rates in different areas of the dorsal integument.
White (1973) came to the same conclusion regarding Amblyrhynchus
cri-status. He could get the lizard to disregard the position of the
real sun and reorient toward a concave mirror directing the sun's rays
at the lizard's head.

Figure 48. Positive and negative orientation (trunk tilting) during
lizard heating and cooling. The three experimental runs
were the same as illustrated in Figure 47. Substrate
temperatures were 20 C (I), 31 (II), and 38 (III). The
right and left photographic areas of the neck plus trunk
were measured from photographs taken directly above the
lizard. Points falling on the dotted line signify the
lizard was in a bilaterally symmetrical posture, with
equal right and left side photographic areas. The right
side was either facing the heat lamp (open circles),
facing away from the lamp (solid circles), or the lamp
was turned off (solid triangles). The animal was either
heating (solid lines) or cooling (dashed lines).

BODY TEMPERATURE
192

193
Iguanas changed their pattern of sun orientation both during the
day and the year (Figure 49). Morning basking in the fall, as well as
in the spring and summer, commonly occurred while in a perpendicular
orientation. Lizards more commonly oriented positive parallel in the
winter, presumably due to the increased declination of the sun. Object
utilization during early morning and late afternoon basking also
increased in winter. Negative parallel orientation, the minimum heat
intake position, became significant during the hottest part of the day
in the fall, but remained below chance expectation at all times in
winter. Clearly, heat gain orientation was more common in the winter
than the fall, where it was restricted to the morning hours. However,
had the autumn pen not been totally shaded by 1645, the late afternoon
increase in perpendicular orientations may have been sufficient to
increase the 1500-1600 hour Chi-square to a significant level.
Cyclura has a large proximal colon, usually containing in excess
of 40 percent of the digestive tract volume, located next to the left
anterior abdominal wall (Iverson 1977). Does the animal preferentially
heat this area, by orienting its left side perpendicularly toward the
sun more than its right side? No significant difference was found
between left vs. right side orientations when each full day's, body
half data were simply summed and compared. However, when the frequency
distributions were compared, the left side was favored in orientations
between 68 and 121° in September and between 95 and 166° in December-
January (Figure 50, p _< 0.05, Wilcoxon signed rank test) in the
Gainesville pen. The pen data for the warmer month of August showed
no significant pattern. The left side predominance in the perpendicular
range was greatest during morning heating, both in the September and

Figure 4$. Daily and seasonal variation in sun orientation in the
Gainesville pen. Instantaneous sun orientations were
determined four times per hour for ten adult lizards
during four clear days in September (solid lines, X Ta
in the shade during lizard activity = 30.4 C) and one
day each in late December and early January (dashed
lines, X Ta shade = 23.0 C). Dotted lines are expected
percentages based on chance. Possible significant
differences of percentages from random expectation were
tested for using Chi-square.

PERCENT OF OBSERVATIONS

196
10
SEPT
DEGREES FROM NEGATIVE PARALLEL ORIENTATION
Figure 50. Left vs. right side sun orientation in the Gainesville pen.
Instantaneous sun orientations were determined four times
each hour for ten adult lizards during four clear days in
September (N = 271 measurements) and one day each in late
December and early January (N = 145). The orientations were
arranged according to whether more of the left (shaded bars)
or right side (diagonal hatching) of the lizard's trunk
faced the sun.

19 7
Deceinber-January pen study and in a caged field iguana in October.
Thus, it does appear that the proximal colon was preferentially heated.
However, more data on free-ranging iguanas should be collected.
Another interesting finding, which may be related, comes from
February and October tracking. Iguanas moved predominantly in clock¬
wise, approximately linear, and counterclockwise paths during morning,
midday, and afternoon hours, respectively (Figure 51). The orientation
change was probably related to the changing sun position in the sky.
Thus, with the sun at a southerly declination, a lizard had its left
side facing the sun as it moved clockwise on its first foray, when,
presumably, differential heating would be most necessary, especially
if it were also important in the defecation response.
Integumental scales are imbricate over much of the body surface
of Cyclura carinata. When heated from directly anterior and posterior
to the overlapping scale edges, the posterior heating rates of dried
skin patches often exceeded the anterior rates at equal angles of heat
lamp orientation (Table 19). Skin patches from the ventral abdomen
and the dorsal distal hindleg (1,2), with little scale overlap, heated
at about the same rate at each anterior-posterior pair of equal
radiation incidence angles. A/P ratios were higher in patches with
more imbricate scales (3-6). It is of some interest that the scales
of the ventral proximal tail had the greatest A/P ratios, yet receive
mostly thermal radiation from the substrate rather than direct solar
radiation.
Regal (1975) demonstrated for Dipsosaurus dorsalis and Soeloporus
magister that imbricate scales act as miniature radiant heat shields by
producing a shadow zone underneath the overlapping edges. Unlike

NUMBER OF FORAYS
193
HOUR INTERVAL
Figure 51. Daily change in the pattern of travel during forays. Results
are illustrated for two lizards tracked for a total of 17 days
in February plus October. Lizards walked in either a circular
or elliptical clockwise path (solid line), approximately linear
path (dotted line), or counterclockwise circular or elliptical
path (dashed line).

199
Table 19. Skin patch heating. Six patches of integument
were removed from a single Cyolura carinata adult,
dried, and heated repeatedly from 20 to 50 C (Ta =
20 C). The heat lamp was oriented normally to the
patch or at one of five different, less than normal
angles, both directly anterior and directly posterior
to the imbricate edges of the scales.
HEATLAMP ORIENTATION ANGLE (DEGREES FROM NORMAL)
0 15 30 45 60 75
HEATING RATES (SECONDS/30 C)
1. VENTRAL ABDOMEN
Normal
Anterior
(A)
34.4
33.8
39.3
57.1
116.4
411.1
Posterior
(P)
36.9
42.6
59.7
130.9
391.2
A/P
0.92
0.93
0.96
0.89
1.05
2.
DORSAL
DISTAL HINDLEG
27.2
36.9
45.1
52.0
145.1
490.8
37.3
44.9
53.5
147.1
453.8
0.99
1.00
0.97
0.99
1.08
3. DORSAL ABDOMEN
31.3
32.1
36.0
57.3
143.3
398.6
34.6
35. 7
55.4
110.0
289.8
0.93
1.01
1.03
1.30
1.38
4.
DORSAL
DISTAL FORELEG
39.4
37.7
52.4
75.0
141.8
335.0
38.0
45.9
66.4
123.6
240.2
0.99
1.14
1.13
1.15
1.39
5.
DORSAL
PROXIMAL TAI
:l
55.9
59.2
64.7
90.7
169.0
626.8
52.6
59.6
68.2
120.0
303.8
1.13
1.09
1.33
1.41
2.06
6.
VENTRAL PROXIMAL TAIL
42.7
48.1
50.5
85.8
217.0
780.5
41.8
45.7
61.4
111. 7
289.4
1.15
1.11
1.40
1.94
2.70

200
Divsosanrus and Sceloporus, which have dorsal trunk scales arranged in
rows parallel to the long axis of the body, Cyclura scale rows run
obliquely toward the mid-sagittal crest at an anterior angle of approxi¬
mately 60°. The radiant source should be normal to the scale rows of
the dorsal trunk to maximize heat input. Thus, a maximally heating
iguana should orient its body obliquely away from the sun, at an angle
approaching 120°. Basking iguanas observed in the field and pen, in
both the morning heating and regulating T^ phases, commonly oriented
at slightly more than 90° (Figure 50). However, the degree of difference
from normality for the peak in the frequency distribution of perpen¬
dicular orientations was within the range of error for the technique
used to determine the orientations. Unfortunately, the definitive
test, repeated radiative heating of individual lizards at different
orientation angles in a constant thermal environment, was performed
on only one adult male iguana. The maximum heating rate did occur at
120° (2^ = 13 C, T^ range = 20-25 C, single heat lamp angled 35° below
the horizontal, angles of incidence from 0 to 180° at 30° plane inter¬
vals). Sample size, however, was obviously too small to draw any
conclusion.
Thigmothermia
2
The relationship between surface area (cm ) and body weight (g)
in Cyctiwa. was estimated by using a linear regression equation generated
from Norris' data (1967) for eight species of iguanid lizards (Log
Surface Area = 0.688 Log Body Weight + 1.025, N = 13, v = 0.996, p _<
0.0005). The maximum possible substrate contact of an unanesthetized

201
lizard was assumed to be equal to the value for the same anesthetized
lizard. Maximum substrate contact ranged from 13.4 to 16.0 percent
(X = 14.3, N = 4 adults) of the total estimated surface area. Minimum
substrate contact for a nonmotile lizard (Figure 44, Illustration 13),
consisting of a small patch of distal tail and palmar plus plantar
surfaces of the feet, averaged 1.1 percent of total estimated surface
area. A lizard running across a hot substrate made even less instan¬
taneous substrate contact.
When only the head and neck or only the head, neck, and forelegs
were involved in substrate contact reduction, head elevation, as
measured by the substrate to nostril distance, was linearly and inversely
related to substrate contact (Figure 52). Lizards in the photo-
thigmotron did not make substrate contact with the head, probably due
to disturbances: intermittent moving of the camera as well as the
noise and vibrations of the water bath. The upper regression line did
extrapolate to approximately 100 percent contact at zero head elevation.
Thus, maximum substrate contact must be about equal in anesthetized and
unanesthetized lizards. A second regression line was fitted to the data
for rear leg participation in substrate contact reduction. It had
about the same slope as the upper regression line, as demanded by
simple geometry. Reduction of substrate contact by elevation of only
the posterior portion of the body was never observed. Maximum substrate
contact must be energetically inexpensive, since the posture is a
consequence of only the force of gravity on body tissues and muscular
relaxation equivalent to an anesthetized state. Animals in this posture
had slightly concave vertebral columns when viewed from the side. Buccal
inspiration (guiar pumping) was more common than in the more upright

Figure 52. Participation of the lizard's head, neck, and legs in the reduction of substrate
contact. Data for a single adult female in the photothigmotron are illustrated.
The lizard elevated only its head and neck off the substrate (solid circles),
utilized its forelegs as well (solid triangles), or both its fore and hindlegs
(open circles and triangles; some or no abdominal contact, respectively). The
correlation coefficient for the linear regressions were -0.841 (upper line,
p < 0.0005) and -0.432 (lower line, p > 0.05).

100
90
80
70
60
50
40
30
20
10
203

204
postures, probably due to the decreased efficiency of the thoracic
inspiratory musculature.
Percent reductions in substrate contact by head-neck-trunk
elevation and lateral compression were determined for three adult
iguanas from their photothigmotron photographs. Lateral compression of
the trunk was defined as the area difference between actual thoracic
plus abdominal contact and the maximum possible contact for that area
in the anesthetized lizard. Lateral compression's contribution only
exceeded that of body elevation when the lizards were nearly completely
relaxed on the substrate and never exceeded 16 percent of the maximum
possible contact (Figure 53). Body elevation reductions of approximately
1-7 percent, due to changes in head contact, did not occur. Reductions
of approximately 8-15 and 16-48 percent were due to head-neck and head-
neck-foreleg participations, respectively. Trunk elevation instan¬
taneously increased by 20-30 percent when the hindlegs came into action.
The actual elevation increases were even larger since decreases simul¬
taneously occurred in leg and tail contact. Complete lateral relaxation
did not occur between trunk elevations of 16 and 37 percent; that is,
over much of the range of the upright posture. Complete lateral
relaxation of the hindtrunk vs. foreleg involvement in forebody elevation
appear to be partially mutually exclusive.
Ventral body contact responses to Cyclura in different conductive-
convective environments were usually thermoregulatorily adaptive (Figure
54). The lizard made greater ventral contact while heating in positive
T - T. T - T-, environments (Experiments I - III) than when cooling in
s as d
negative ^ - h environments (Experiments IV, V), with a maximum
heating-cooling contact range of 38 percent. Note how contact declined

PERCENT REDUCTION OF MAXIMUM SUBSTRATE
CONTACT BY LATERAL COMPRESSION
AND TRUNK ELEVATION
Figure 53. Relative importance of lateral compression of the trunk vs. head, neck, and trunk
elevation in reducing substrate contact. Three lizards, represented by solid circles,
open circles, and open triangles, were each monitored in the photothigmotron for
several days under differing environmental conditions.
205

206
Figure 54. Conductive and convective heating and cooling of a single
Cyolura carinata. Experiments were run on five consecutive
days for a 1135 g adult female: I; T= 23 C, Ts = 41, II;
Ta = 29; Ts = 41; III; Ta = 36, Tg = ti, IV; Ta = 38, Ts = 21,
V; Ta = 27, Tg = 19. The lizard was either heating (solid
line) or cooling (dashed line). Maximum substrate contact
was assumed to be equivalent to that of an anesthetized state.
The lizard's mean regulating phase Tfo and range in the thermal
gradient are indicated by the vertical and horizontal lines,
respectively.

207
in III as T-^ increased to above the preferred range. The lizard was
probably attempting to decrease conductive heat input as well as cool
itself convectively. As Tasymptotically levelled at the end of run
III, the lizard relaxed somewhat on the substrate. The lizard did not
immediately minimize substrate contact in IV as expected, but rather
gradually reduced it as burrow T^ was approached. The technique utilized
to start the run, that is, pulling out the insulating plywood substrate,
inevitably startled the animal. Immediate minimization of contact may
have been learned had each experiment been repeated several times.
Also, the high T^ at the beginning of the run may have partially over¬
ridden the peripheral heat receptor inputs. The lizard responded with
the appropriate thermoregulatory posture to fairly small T - T
s a
differences (5 C in III, -8C in V). The upright posture was still
maintained at the end of V even though T^ was only 4.5 C above Tg and
3.5 C below T . Significant differences in substrate contact at equal
gut temperatures, as in I vs. V and II vs. IV, were probably primarily
due to differences in peripheral temperature receptor inputs to the brain.
This was especially true in I vs. V, where brain-gut temperature
differences were small due to the nearly equal T^'s in the two experi¬
ments (23 and 27 C).
Figure 55 illustrates some of the ways Cyctura responded to
simultaneous radiant heating and conductive heating or cooling. Sub¬
strate contact increased as T^ increased in all three experiments, as
was observed during the morning heating phase in the field. Substrate
contact at low was greatest in III, the run with the highest substrate
temperature. However, the lizard did not maximally flatten in III at low

Figure 55. Conductive heating and cooling of a single Cyclupa cca“inata,
with and without simultaneous radiative heating with a heat
lamp. Experiments were run at a constant air temperature
(19 C) on three consecutive days for a 985 g adult female;
I; Ts = 20 C, II; Tg = 31, III; Ts = 38. The lizard was
either heating (solid line) or cooling (dashed lines). The
heat lamp was turned off at the points marked by the arrows.
The lizard's mean regulating phase Tj, and range in the
thermal gradient are indicated by the vertical and horizontal
lines, respectively.

SODY TEMPERATURE
PERCENT MAXIMUM POSSIBLE SUBSTRATE CONTACT
_-W

210
T-, . Rather, it balanced heat intake from conductive and radiant sources.
b
Further increases in substrate contact would have decreased radiant
input, due to the considerable angle of the heat lamp. As T^ increased
from I to III, the lizard reached increasingly higher T^'s before moving
away from the heat lamp. Core temperature at radiant heat escape rose
5.3 C, from 34.6 to 39.9 C, while substrate temperature increased 18 C,
from 20 to 38 C. Thus, the T-, - T difference decreased from 14.6 to
only 1.9 C! The lizard was probably regulating skin surface temperature
rather than gut temperature. When surface temperature reached a narrow
escape range, the iguana departed, at a low T ^ on cold substrate, at
a higher S’ ^ on warmer substrates. When the heat lamp was turned off
in I, the lizard soon reduced contact nearly maximally due to the low
T . However, maximum elevation was only briefly maintained. The posture,
s
utilizing both fore and hindlegs in a walking stance, was probably too
strenuous to maintain for long periods. The lizard cooling in II reached
nearly maximum substrate contact at a ^ of 2.5 C above TThe
substrate was still a heat sink. This demonstrates the mistake of
assuming thigmothermia just because the animal relaxes on the substrate.
Maximum contact was a response to low T and the small T- - T
^ a os
difference. In the field, the conductive heat loss while maintaining
a maximally prostrate posture on a substrate slightly cooler than the
body core may be less than the forced convective heat loss if the
animal assumed an upright posture.
Figure 56 illustrates the changes in substrate contact of a
lizard subjected to increasing, stable, and decreasing substrate tempera¬
tures, with T held constant and no radiant heating with a heat lamp.
CL
Substrate contact increased as increased, as in Figures 54 and 55.

Figure 56. Thigmothermic responses of a single Cyolura cavinata to
periods of increasing, stable, and decreasing substrate
temperature. Experiments were run on three consecutive
days on an adult male weighing 1288 g. Body and substrate
temperatures vs. percent substrate contact have been
plotted (dashed and solid lines, respectively). Minutes
since t , the start of the experiment, are indicated by
Arabic numbers. The lizard was in thermal equilibrium
with air and aluminum substrate temperatures at Ta of
Experiments I and II and with Ta and the cool plywood
substrate temperature at TQ in Experiment III, at which
time the plywood was removed.

212

213
The greatest rate of increase of substrate contact occurred at T.^
ranges of 33.3-34.6 (I), 31.0-32.7 (II), and 36.0-37.4 C (III) (T -
s
Th = 9.4-10.5, 7.9-9.8, and 6.7-9.7 C, respectively). Smaller T^ -
21 's were coupled with lesser substrate contacts. Due to the low T
the lizard relaxed nearly maximally at a T as high as 50.1 C in III,
O
a T approximately 3.9 C above the mean critical thermal maximum of the
s
body core. Substrate contact was abruptly reduced when T^ moved into
the upper voluntary tolerance range. A contact cycle was produced in
II by first increasing, then decreasing substrate temperature.
Low Body Temperature Behavior
Behavioral responses were lost during cooling in order of decreas¬
ing energy demand and neural pathway complexity, and regained in the
reverse order during heating, at approximately the same intestinal
temperatures as during cooling (Figure 57). Crawling, the most
energetically demanding behavior, was lost first, and opening the eyes,
the least demanding, was lost last. Running and walking abilities were
lost sequentially at higher I^'s, but the actual temperatures were not
determined. Bloating was the next to the last response lost before
lizards became completely immobilized. Lizards utilized the hyoid
musculature to maximally inflate the lungs, greatly increasing the
circumference of the trunk. Thus, a nearly helpless animal could still
wedge itself securely in its burrow or, perhaps, intimidate a predator
by bloating.
Lizard critical thermal minima (Spellerberg 1972a) and maxima
(Lowe and Vance 1955, Larson 1961) increase, within limits, as
acclimation temperature increases. The mean critical thermal minimum

214
COOLING REWARMING
Figure 57. Behavioral responses of Cyclura carinata at low body temperatures.
The temperature of the large intestine was recorded when a lizard
lost and regained each of seven behavioral responses during cooling
and reheating, respectively: 1. crawling, 2. turning over when
placed on the back (critical thermal minimuni, 3. maintaining the
head in a horizontal orientation with respect to the ground when
the body long axis was elevated and depressed at the tail, 4. main¬
taining a constant field of view by rotating the head or eyes when
the body was rotated laterally, 5. moving the legs when pinched
while placed on the back, 6. bloating, 7. closing and opening the
eyes. The sample contained six adults. Horizontal lines are
means, vertical lines; ranges, and rectangles; plus and minus two
standard errors of the mean. The dotted line marks the lowest
temperature recorded during the study (exposed substrate tempera¬
ture; Iverson 1977).

215
for summer lizards in the Caicos was probably one to two degrees higher
than 12.8 C, which was calculated for animals acclimatized to late
May environmental temperatures in Gainesville. In support of this
hypothesis, all lizards acclimatized to the Caicos summer were completely
anesthetized at 10 C when cooled for transmitter implantation.
A lizard must meet several locomotory requirements, one of which
is to move successfully from place to place at the lowest T. it is ever
likely to endure. It isn't surprising, then, that Cyclura's CTMin was
well correlated with the lowest environmental temperature recorded
during the study, 12.5 C. The correlation holds for other lizards
examined thus far (Spellerberg 1972b). Cyclura probably experienced a
of 12.5 C on only rare occasions, unless sequestered for the night
in a surface level retreat, since cold fronts did not last long enough
for deep soil temperatures to fall this low.
The CTMin varies significantly for lizards living at different
latitudes. Liolaemus multiformis, ranging high into the Andes of
southern Peru, is able to move about slowly at a of 1.5 C (Pearson
1954). The 29 species of lizards from southeastern Australia examined
by Spellerberg (1972a) all had CTMin's lower than Cyclura's. Expectedly,
minimum environmental temperatures fell lower there than in the Caicos.
Both Anolis carolinensis and Sceloporus occidentalis have similar CTMin's
to Cyclura's (Kour and Hutchison 1970).
The CTMin of some lizards could be lowered by acclimation to
below -0.52 C, the freezing point of the whole body (Spellerberg 1972a).
That is, the lizards could still turn over when supercooled. This
probably would not be true for Cyclura, since none of Spellerberg's
acclimation ranges approached the necessary 13 C. The acclimation

216
range appears to be greater for the CTMin than for the CTMax (Kour
and Hutchison 1970). This is logical since lizards can always retreat
from high environmental temperatures, but not from minimums passively
experienced while in a nocturnal retreat. The correlate of this is
Spellerberg's finding (1972a) that the CTMin was better correlated than
the CTMax with the geographic distribution of lizards.
When placed on its back, held firmly for a few seconds with legs
adpressed to the trunk and tail, and then gently released, Cyatura
remained quiescent for short periods before righting itself. This so-
called tonic immobility has been induced in other lizard species
(Prestrude and Crawford 1970, Hennig and Dunlap 1978). The righting
response could be induced prematurely by gentle poking of the body.
On the contrary, it could not be induced, even by strong prodding, once
the lizard had reached the CTMax or CTMin. Thus, tonic immobility did
not obscure the CTM determinations.
The lethal minimum body temperature was not determined. However,
two summer acclimatized lizards cooled to 0 C in the Caicos were
partially paralyzed and subsequently died after rewarming.
High Body Temperature Behavior
Panting in Cyotuva was a proportionally controlled behavior,
probably with various effector outputs directly proportional to the sum
of positive deviations of many body temperatures from a reference set-
point temperature, presumably located in the hypothalamus. Effector
outputs consisted of the gaping time per minute, the degree of gape, the
extent of elevation and blood engourgement of the tongue, and probably,
the respiratory and salivation rates. The panting threshold temperature,

217
at which the lizard initially gaped, was estimated by gut and head
temperatures. Unfortunately, the respiratory rate was not determined
during gaping of restrained lizards heated in the field. It is
probable, however, that the respiratory rate increased as T^ rose
during gaping. Thus, Cyctura is a true panting lizard rather than
a thermal gaper, a lizard with reduced ventilation rate during gaping
(Heatwole et at. 1973).
Radiantly heated, restrained lizards in the field initially panted
for only a few seconds. As T^ continued to rise, tongue elevation and
the degree of gaping increased and the mouth was closed less often,
eventually only long enough to recoat the surface of the tongue and
walls of the buccal cavity with saliva (Figure 44, Illustrations 13
and 14). The saliva secretion rate appeared to increase with T^.
Well hydrated lizards, fed white grapes in the Gainesville pen,
salivated sufficiently to overflow the mouth and form a droplet on the
lower jaw. On the other hand, three lizards heated in the field in
June did not drool at all, probably due to the lower water content of
their food. Restrained lizards alternated between panting bouts and
struggling to free themselves from their tethers. Panting ceased
during struggling. It could also be stopped briefly by scaring the
animal.
The mean intestinal heating rate in three restrained, morning
sun heated lizards dropped from 0.111 C/min just before panting commenced
to 0.069 C/min during panting, even though heat input was increasing
with time. It is likely that the reduction in head heating rate was
even greater (Crawford 1972).

218
Eye bulging, which usually became visually apparent soon after
or before panting commenced and continued at least until the CTM was
reached, gave the lizard a perpetual stare. It probably resulted from
contraction of the jugularis muscle surrounding each internal jugular
vein, increasing the blood pressure in the orbital venous sinuses
between and around the eyes and diverting venous flow to the external
jugular and vertebral veins (Heath 1966). Eye bulging probably serves
the same two functions it does in Phrynosona. The skin immediately
around the eyes was commonly the first head skin shed during the single
annual molt in Cyclura. Eye bulging may aid in this process. During
two successive heating runs of a single large male in the photothigmc-
tron, the head heating rate decreased very soon after eye bulging and
considerably before panting commenced. Head temperature was 3.4 and
3.9 C above gut temperature. Head heating decreased from 0.59 and
0.40 C/min to 0.09 and 0.07 C/min, respectively, a mean reduction of
84 percent. Gut temperature continued to rise steadily after bulging
in the first run, at a rate of 0.10 C/min. However, in the second run,
it increased from 0.04 to 0.08 C/min, as expected if the body core was
functioning as a heat sink for the head. Thus, rerouting venous blood
away from the internal jugular vein-internal carotid artery counter-
current heat exchangers during eye bulging probably lowers the positive
^heacT^body <^-’-^^erence at high temperature. Eye bulging may serve a
head cooling function both in emergency in conjunction with panting
and alone in less urgent situations below the panting threshold 2^.
The gut temperature range for the panting threshold was at least
11.0 C, from 32.8 C for a jig restrained lizard during heating in the

219
controlled temperature room in August, to greater than 43.8 C, the T^
maximum for free-ranging lizards basking without panting in the late
afternoon. The threshold declined as heat input increased at midday
and again increased in the afternoon as heat input waned (Figure 58).
This is further evidence for a peripheral heat receptor input influenc¬
ing an effector output.
The log of the total panting time after shading in the roof top
arena was well correlated with the sum of the gut and head temperature
lag times, and less well correlated with each separate time (Figure 59).
The multiple linear regression was: Log Total Panting Time = -1.421 +
0.097 Gut Lag Time + 0.115 Head Lag Time. Thus, gut and head tempera¬
tures appear to be about equally important in determining the total
panting time after shading. As the sum of the gut and head temperature
lag times increased linearly, the panting time increased exponentially.
The reason for this was that the longest panting times occurred during
the hottest part of the day, when the highest shaded air and substrate
temperatures occurred, as well as the greatest radiant input from the
aluminum walls of the arena. Although head temperature fell rather
rapidly after shading and panting commenced, gut and body surface
temperatures did not.
The intestinal temperature at the critical thermal maximum averaged
46.2 C (46.0-46.3) for three freshly caught Cyclura heated in the sun
starting at 0700 on July 2 (body weights = 150, 536, and 800 g). The
head-intestine temperature difference was probably +1-2 C at the CTM,
even though the mean intestinal heating rate after panting commenced was
only 0.07 C/min. In the turtle Chelodina longicollis, the head-body
temperature difference at the CTM increased with body weight, from

HEAD AND GUT TEMPERATURES (°C)
AT START OF PANTING
o
43
42
41
40
-I L I I I I I I I I | | I
10 II 12 13 14 15 16
TIME
Figure 58. Daily adjustment of the panting setpoint to changing heat input. A 1790 g male was repeatedly
heated to the panting setpoint (gut temperature; solid circles and line, middle ear tempera¬
ture; open circles and dotted line) while contained for two days in July in a rooftop arena
in Gainesville. Curves were fitted by eye.
220

Figure 59. Correlation of total panting time with head and gut thermal lag times. A
single 1790 g male was repeatedly heated to panting during two days in the
rooftop arena and immediately shaded when panting commenced. The length of
time that head temperature (open circles), gut temperature (open triangles),
and the sura of both (solid circles) continued to rise after shading (lag
times) have been plotted against the total minutes of each panting bout.
The correlation coefficients for the three linear regressions are indicated
at the top of each line.

TOTAL PANTING TIME (MIN)

223
approximately 0.8 C at 100 g to 3.5 C at 1200 g (Webb and Witten 1973).
The head temperature at the CTM was independent of body weight, but the
posterior body temperature at the CTM decreased with increasing body
weight. Thus, head temperature as well as gut temperature should have
been measured during CTM determination in Cuoliwa.
When both panting and struggling ceased at 48.0 C (47.9-48.1), two
lizards were rapidly cooled. Since neither recovered, 48.0 or lower
must equal the lethal maximum (LM). The three lizards simultaneously
reached the LM at 1025, even though body weights varied by a factor of
five.
The minimum thermal safety margin (Heatwole 1970), defined as the
difference between the maximum voluntary tolerance and the CTM, approxi¬
mated 4.8 C at midmorning (X MVT = 41.4 C between 1000 and 1100 for eight
hyperthermic summer lizards). The three lizards used in the CTM
determination passed through the MTSM in a mean time of 68 minutes,
sufficient to avoid fatal overheating during practically any high T^
encounter. Of course, the MTSM time decreased considerably as midday
approached. For example, a 375 g male took only 15 minutes to heat from
41.4 C to its gut measured CTM of 44.9 C at 1310 on July 3. This value
is low in comparison to the three lizards measured on July 2, probably
due to an increased thermal lag between head and body temperatures as
heating rate increased, rather than a lowering of the CTM as measured
by head temperature. In fact, the CTM peaked around midday in two
turtle species, Chryserrrys picta (Kosh and Hutchison 1968) and Chelodina
longiaotlis (Webb and Witten 1973).

224
Head-3ody Temperature Differences
Gut and orbital venous sinus temperatures were monitored in a
single 1600 g male Cyclura in the photothigmotron. Head-body tempera¬
ture differences of up to +13.7 C were recorded (T â–  =40 C, T , , , =
^ avr ’ substrate
20, heat lamp 30 cm from the lizard's head). The maximum head-body
temperature difference was only -2.3 C when T and T were reversed
a s
and the heat lamp was directed at the lizard's trunk. The 11.4 C
difference in the absolute values was primarily due to the greater
surface to mass ratio of the head. In still air, is more important
than T in control of both head and body temperatures. For example, with
CL
a T of 21 C and a T of 40 C, the steady state head and gut temperatures
were 32.6 and 34.7 C, respectively, 4.2 and 8.4 C closer to substrate
than air temperatures. Lizards basking after a long burrow submergence
on cool, rain soaked sand in the midday sun should have the largest
transient head-body temperature differences. Individuals resting in
the midafternoon shade, characterized by nearly stable gut and head
temperatures and a small Tn - T , should have the smallest head-body
temperature differences.
The head and body temperatures of a single iguana were monitored
for two days in the Gainesville rooftop arena (Figure 60). When the
lizard was in the sun, the head always heated faster than the gut (X
head/gut heating rate ratio = 2.4, range = 0.9-6.0, N = 7) and head
temperature usually exceeded gut temperature (X head-gut temperature =
1.0 C, range = -0.1-1.6 C). Thus, daily head temperature maxima for
free-ranging lizards were probably higher than the coelomic and gut
maxima listed in Table 1. In the shade, the lizard's head consistently

Figure 60. Head-body temperature differences during heating and cooling of a 1790 g male
Cyoluva carinata in the Gainesville rooftop arena. Head temperature was
measured just inside the tympanum with the probe of a near-field transmitter
(dashed line). Core temperature was measured in the gut with another near¬
field transmitter (solid line). The lizard was manually shaded (horizontal
lines) immediately after panting commenced (vertical lines and rectangles).

226

227
cooled faster than its gut. The mean head-gut difference was -0.3 C
(-1.5-0.4). It appears that the head-body temperature difference is
greater during the heating phase than during the cooling phase. Head
and body temperatures approached equality as the time in the shade
increased.
The Integument: Metachromatism and Reflectivity
Head, trunk, and leg integuments were dark gray to charcoal black
at low body temperatures during nocturnal inactivity, the first minutes
of the morning basking phase, and long periods of overcast. Iguanas
were usually darkest immediately after emergence. The ventral pectoral
region was usually the darkest area on a lizard in its dark phase
(Figure 61, #7). The proximal ventral tail was always the most highly
reflective area (#4). Dorsal dark phase reflectivity was well matched
to the fused coralline sand substrate, which was darkened by a thin
encrustation of two species of blue-green algae (#2 vs. #3). Extensive
tracts of this sparsely vegetated substrate lay along the windward side
of Pine and Water Cays, interspersed with patches of cream colored dunes
sand. Low lying karst ridges, as in Rocky Coppice, were also encrusted
with the same algal growth. However, it was not determined whether
lizards in their dark phase preferentially basked on dark substrates.
The high Cyatura density associated with karst ridges in Rocky Coppice
may be due in part to the advantages of dark hillsides to basking
lizards: 1. good color matching at low 2. fairly large unvegetated
areas, 3. protection from the wind, and 4. high net heat input on sun¬
facing slopes. The greater abundance of burrows and greater diversity

PERCENT REFLECTIVITY
Figure 61. Integumental and substrate reflectivities. Skin patch 1 was excised from an adult Water Cay
lizard in its light phase, patches 2, 4, 5, 6, and 7 from another Water Cay adult in its
dark phase. Patches were excised from the 1) dorsal abdomen, 2) dorsal abdomen, 4) ventral
tail immediately posterior to the anus, 5) ventral abdomen, 6) dorsal tail immediately
posterior to the anus, and 7) ventral pectoral region. Reflectivity curve 3) is for a piece
of a common Caicos substrate, fused coralline sand encrusted with two species of algae
(Phylum Cyanophyta). Numbers in parentheses are mean percent absorbances.
228

229
of food plants in Rocky Coppice (Auffenberg in prep.) may also account
in part for the high lizard density.
Live adult iguanas primarily absorbed rather than reflected near
infrared light, as indicated by photographing them with Kodak Infrared
Ektachrome film. The animals, appearing blue colored while in the
regulating phase range, stood out clearly from the surrounding red
colored photosynthetic vegetation, which reflected near infrared light.
Lizards were well concealed in the scrub due to excellent color
matching, countershading, and cryptic behavior. The integument lightened
as body temperatures increased, a change well correlated with the
animal's movement into the brush during the first foray. To varying
degrees, the head, trunk and legs became pale yellow-green and light
gray on Water Cay (Figure 61, #1) and Pine Cay lizards, respectively.
Tail color change with increasing T, was slight. Countershading made
it difficult to distinguish the ventrolateral side of the lizard from
the sand substrate. The ventral abdomen was grayish cream in the dark
phase and cream colored in the light phase and was reflectively less
labile than the ventral pectoral region and the dorsal body surface.
Cryptic behavior enhanced concealment of lizards in the bush. The
animal sat motionless in an upright posture until an approaching potential
predator, in this case myself, was seen. The lizard soon walked about
five to ten meters out of sight and sat motionless again. The behavior
was repeated when I followed. On the contrary, when not habituated to
humans, located in open areas, or recently emerged, lizards commonly
ran to the nearest burrow without making any initial eye contact.
Color change probably has a thermoregulatory as well as a conceal¬
ment function in Cyclura. The equation of Hutchison and Larimer (1960)

230
was used to estimate the mean heat gain of dorsal skin patches exposed
to direct solar radiation:
d ~ i?("ioo/)
(6)
where H is the mean heat gain in langleys (cal'cm ‘''min--*-), R is the
energy content of solar radiation between 400 and 1100 nm equal to 1.07
langleys for an air mass of one (high transmission), and A is the mean
percent absorption of the skin patch over the same spectral range. The
mean heat gains for dorsal abdominal skin patches from lizards in their
light and dark phases were 0.763 and 0.919 langleys, respectively. The
light phase value fell in the range found by Hutchison and Larimer for
four desert lizard species (X = 0.747, range = 0.685-0.318). Two of
these lizards were iguanines, Dipsosaurus dorsalis and Sauromalus obesus
The dark phase value fell in the range of their six forest species (X =
0.948, range = 0.898-0.990). The rates of heat gain were estimated to
be 0.78 and 0.92 C/min for lizards in their light and dark phases,
respectively, a difference of 15.3 percent. These data are for a
2
964 g animal in a walking posture (surface area = 1034 cm , assumed to
be the same as a Sauromalus obesus of the same weight; T^ = 36 C) on a
level, light sand substrate (T^ = 40 C) in still air (T = 30 C) on a
clear day, calculated using Norris' technique (1967, pp. 209-210).
Melanin was deposited sparingly between some muscle blocks and
more abundantly in the epidermis and the dermis just peripheral to
the muscle layer. Peritoneal melanization increased as the thickness
of the overlying musculature decreased: 1. the peritoneum was lightest
dorsally, where the epaxial musculature was thickest, darker ventrally,
and darkest in a clearly defined lateral band, 2. the peritoneum was
lighter in the pleural than the abdominal cavity, 3. juveniles had

231
darker peritoneums than adults. Porter (1967) demonstrated that the
black peritoneum helped absorb ultraviolet light (spectral range = 290-
400 nm) before it could penetrate and possibly mutagenically damage the
viscera.
The Integument: Sensory Spots
Specialized "tactile" or "sensory" spots have been described for
the integument of a wide variety of reptiles (Mauer 1895, Jaburek 1926).
In general, the epidermal cells are more columnar than in the surrounding
unspecialized areas, and the overlying inner alpha- and outer beta-
keratin layers are variously modified (Miller and Kasahara 1967). I
found spots in all representatives of 21 iguanid genera examined, most
numerously in the nine iguanine genera. Spots in Cyclvra aav'Lnata
numbered from zero to +500 per scale, located on different areas of
scales from different parts of the body. They were circular, peglike,
flat to slightly domed superficially, and 0.1 to 0.2 mm in diameter.
Each spot consisted of a dermal papilla covered with a stratified
columnar epithelium and extended above the living cells of the rest of
the epithelial surface. The surface of the peg was covered with a thin
layer of alpha-keratin, slightly thinner than over the rest of the
epithelium. The outer beta-keratin layer only extended up to the side
of each peg, not forming a "lens-like" cap on top as reported for
Varanus bengalensis (Miller and Kasahara). The outer flat or slightly
domed alpha-keratin-covered peg surface was practically continuous
with the surface of the surrounding beta-keratin. In the living animal,
the peg may be slightly elevated above the rest of the integumental
surface. Faint traces of nerve fibers were detected in the

232
hematoxylin-eosin stained preparations, extending from the dermal
papilla into the sensory epithelium.
As in other iguanids, spots were concentrated on the head. They
9
were especially numerous on the rostral (X = 2100 spots/cm“, N = 8,
SD = 143, range = 1950-2400), mental, and labial scales (Figure 62).
Spots were absent from the palmar and plantar surfaces of the feet,
perhaps due to the likelihood of abrasive damage by hard substrates.
Spot densities were generally higher on the ventral sides of the
trunk and legs, the reverse on the tail.
Sensory spots most likely serve as touch or pressure receptors
(mechano-receptors) in Cyolura, since they are concentrated at common
contact points with external objects: the snout with objects during
locomotion and rest, the lips with leaves, fruits, and other iguanas
during male-male fighting, the ventral anterior trunk with objects
during basking and with three trunks during climbing, and the ventral
forelimbs with branches during climbing and with other iguanas during
mating. It is also possible that sensory spots serve a thermal sensory
function. They have been suggested to be radiant heat receptors in the
Varanidae (Miller and Kasahara) and the Rhynchocephalian, Sphenodon
punctatus (Maderson 1968). Berk and Heath (1975b) observed jaw rubbing
in Dipsosaurus dorsalis similar to that seen in Cyolura in the same
context; lizards were in the minimum Th range of the regulating phase
and just about to initiate a shuttling movement to the hot end of the
thermal gradient. They suggested that jaw rubbing either served as a
displacement activity or for substrate temperature sensing. The higher
ventral trunk and leg concentration of sensory spots may be related to

Figure 62. Density of sensory spots on different parts of the lizard body surface. Spot
densities were determined at the following locations: Head 1) dorsally on and
around the parietal eye, 2) laterally just anterior to the tympanum, 3) upper
lip, 4) lower lip, 5) ventrally between the mandibular rami; Trunk 6) anterior
dorsal, 7) anterior lateral, 8) anterior ventral, 9) posterior dorsal, 10)
posterior lateral, 11) posterior ventral; Forelegs 12) dorsal upper arm,
13) ventral upper arm, 14) dorsal forearm, 15) ventral forearm, 16) dorsal
forefoot, 17) palm; Hindlegs 18) dorsal thigh, 19) ventral thigh, 20) dorsal
calf, 21) ventral calf, 22) dorsal hindfoot, 23) plantar surface; Tail 24)
dorsal proximal, 25) ventral proximal, 26) dorsal distal, 27) ventral distal.
The sample consisted of eight adult lizards. Horizontal lines are means,
vertical lines are ranges, and rectangles are plus and minus two standard
errors of the mean.

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LOCATION OF SPOT COUNT

235
two other possible thermal sensory behaviors, sand swimming and hard
substrate rubbing with the ventral body surface.

PHYSIOLOGICAL ASPECTS OF THE THERMAL BIOLOGY OF
CYCLURA CARINATA
Heating and Cooling Rates
With constant air temperature and wind speed, low relative
humidity, and substrate and wall temperatures equaling air tempera¬
ture, Newton's law of cooling adequately describes lizard body
temperature change:
<^b
dt
-A(T,
T )
a
(6)
Integration and rearrangement yields:
¿n (T, - T ) = -at + k
b a
(7)
where t is time in minutes, a is a cooling constant in min \ and k is an
integration constant. Plotting £n(T^ - T ) vs. t yields a straight line
with the cooling constant as the slope. The apparent thermal conductance
3 -1 -1 -1
(cm 02'g hr *°C ) is then estimated from the equation:
C =
a -Hs -60
E
(8)
where H is the approximate specific heat of lizard tissue (0.82
s
cal'g ^*C 1) and E is the caloric equivalent of a cubic centimeter of
_3
oxygen (4.8 cal*cm O2) assuming a respiratory quotient of 0.85. C is
an apparent conductance since the effect of metabolic heat production
is not accounted for, A corrected C can be calculated by using the
following equation (Bartholomew and Tucker 1963):
M
C = C ± ¿--J-y- (9)
b a
236

237
3 -1 -1
where M is the metabolic rate (cm 0?*g *hr ) during a cooling (+M/ (T^ -
Tq)) or heating ((T^ - T^)) run. There is some question whether this
later technique is correct (Strunk 1971).
Iguanas heated faster than they cooled both in the cage and on the
jig (Figure 63 and Table 20). The low slope of the line during the
initial twenty minutes of heating on the jig may have been caused by a
fear-induced bradycardia. This has been reported in Alligator mississip-
piensis (Smith 1976). Lizards almost always heated and cooled more slowly
in the cage, undoubtedly due in part to the greater leg and ventral trunk
contact with the substrate than when resting on the narrow wooden strip
of the jig. However, the rates were not significantly lower in the cage
(p > 0.05, student t-test). The mean cooling/heating conductance ratio
was less in the cage than on the jig, but again the difference was not
significant. Apparently, some lizards thermoregulated behaviorally in
the cages, increasing their heating rates by minimizing substrate contact
and decreasing their cooling rates by maximizing contact. The mean C/H
ratio might have been significantly less in the cages had the lizards
been habituated to their surroundings for a longer time.
The uncorrected C/H ratio at 30 C for a single Amblyrhynchus
cristatus (0.52, wind speed = 113 cm/sec, Bartholomew and Lasiewski 1965)
is the only lizard value reported in the literature that is less than the
lowest jig value for Cyclura carinaba, 0.62. The ratios for two
Airrphibolurus barbatus in still air, 0.64 and 0.71 (Bartholomew and
Tucker 1963) are similar to Cyclura cage and jig means, 0.66 and 0.72,
respectively. Dipsosaurus dorsalis in still air (X = 0.88, Weathers
1970), four varanid species at wind speeds of 230-300 cm/sec (X = 0.88,
Bartholomew and Tucker 1964), and TiliqvLa scincoides at 136-156 cm/sec

Figure 63. Heating and cooling rates of a 1435 g male Cyolura earinata in a constant
temperature chamber. The lizard (Number 4 in Table 20) was heated (solid
circles) and cooled (open circles) on a jig and heated (solid triangles)
and cooled (open triangles) in a cage located in a constant temperature
chamber set at 40 and 20 C, respectively. Regression lines were fitted
to the points by the method of least squares. The low slope of the initial
jig heating line may be due to a fear-induced bradycardia (B).

239

o -1 -1 -1
Table 20. Estimated thermal conductance (cm Ü2*g 'hr °C ) during
heating and cooling of lizards in still air.
A. UNCORRECTED FOR METABOLISM
ON JIG IN CAGE
Lizard
Weight (g)
Heating
Cooling
C/H
Heating
Cooling
C/H
1
810
0.229
0.176
0.768
0.236
0.159
0.674
2
812
0.221
0.164
0.742
0.208
0.136
0.654
3
895
0.215
0.140
0.651
0.183
0.112
0.612
4
1435
0.195
0.121
0.621
0.164
0.095
0.579
5
1472
0.191
0.132
0.691
0.135
0.098
0.726
6
1510
0.154
0.131
0.851
0.144
0.101
0. 701
X
1156
0.201
0.144
0. 721
0.178
0.117
0.658
SD
349
0.027
0.021
0.084
0.039
0.025
0.055
240

Table 20. Continued
B. CORRECTED FOR METABOLISM
Body Temperature
25 C
30 C
35 C
Metabolic Rate
Correction
I. RESTING METABOLIC RATE1
C/H (X and Range)
Jig: 0.827(0.719-0.981) 0.812(0.699-0.963) 0.900(0.767-1.078)
Cage: 0.779(0.698-0.872) 0.759(0.671-0.848) 0.857(0.751-0.919)
II. CONDUCTANCE DURING HEATING = CONDUCTANCE DURING COOLING
Required M/Resting M and Required 77/Maximum M (X and Range)
Jig: 2.54(1.13-3.58) & 0.61(0.24-0.80)
Cage: 2.73(1.81-3.17) & 0.66(0.39-0.82)
III. MAXIMUM METABOLIC RATE2
2.84(1.34-4.16) & 0.62(0.25-0.82)
3.04(2.10-3.88) & 0.67(0.41-0.84)
C/H (X and Range)
1.46(0.68-2.14) 6 0.38(0.15-0.49)
1.56(1.09-1.74) & 0.41(0.24-0.51)
Jig: 1.219(1.094-1.534) 1.250(1.100-1.639)
Cage: 1.240(1.075-1.500) 1.278(1.079-1.618)
2.318(1.762-4.225)
3.235(1.615-6.476)
3 — 1 -1
The equations relating resting metabolism (cm 02’g *hr x) to body weight (N = 5 lizards, mean weight= 1252 g,
range = 834-1722 g) were W25C = ~ 0.0000217 W + 0.109 (r = -0.450, not significant at p < 0.05), M30C =
-0.0000395 W + 0.146 (r = -0.926, significant at p < 0.025), and M35C = -0.0000574 W + 07212 (r = -0.821,
significant at p < 0.05).
No correlation was found between maximum oxygen consumption and body weight in the five lizards above (Footnote
1). Therefore, only one maximal metabolic rate corrected was used at each body temperature, calculated from
the equation, Log Mmax = 1.425 Log T- 2.446 (N = 5 lizards, Tjy = 13-43 C).
241

242
(0.85 and 0.90, Bartholomew et al. 1964) all have C/H ratios closer to
unity than Cyolura. Weathers (1970) demonstrated that the ratio was
independent of body weight in Dipsosaurus, partially justifying the above
comparisons. However, he also .found that the ratio increased slightly
as wind speed increased.
Weathers (1972) found that increasing water vapor pressure in the
controlled temperature room had no effect on the cooling rate of Dipso-
saurus but markedly increased its heating rate. He hypothesized that
water condensed on the lizard's skin at body temperatures below the dew
point, adding the latent heat of fusion to the conductive and con¬
vective heating of the air. Water evaporating from the skin surface
above the dew point would reduce the heating rate. This change in heat
input was evident in the nonlinear heating rate of these small lizards.
Water condensed on the skin when relative humidity was above 31.8 per¬
cent, a value undoubtedly exceeded in my experiments. Unfortunately, no
attempt was made to control nor even measure relative humidity during
heating and cooling of Cyolura. Neither were any lizards killed for use
as heating and cooling controls. However, jig heating curves were all
practically linear. Also, any water vapor effect should be less in
Cyolura than the smaller Dipsosaurus.
No investigator has concurrently measured oxygen consumption and
heating vs. cooling rates in lizards. Bartholomew and Tucker (1963,
1964) and Bartholomew et al. (1965) measured heart rates during heating
and cooling and during oxygen consumption at several body temperatures,
then assumed the metabolic rate from the appropriate heart rate. Since
neither heart rate nor oxygen consumption was measured during heating
and cooling in the present study, three metabolic corrections have been

243
calculated: I. resting metabolic rate, II. the fraction of resting and
maximum rates which equalized heating and cooling rates, and III.
maximum metabolic rate (Table 19). In no case did the resting metabolic
rate correction totally account for the low C/E ratios. However, lizards
were completely quiet during only 33 percent of the runs. Struggling on
the jig and climbing the walls of the cage occurred during ten and three
of twelve runs, respectively. Lizards were also disturbed during chamber
switching. Thus, it is unlikely that any lizard metabolized at its
resting rate during heating or cooling. The maximum metabolic rate
correction more than corrected for the faster heating rates. However,
lizards could not have sustained a near maximum rate for the duration
of the run, an average of 148 minutes during heating and 156 minutes
during cooling. The question is, at a T^ of 30 C, for example, whether
lizards averaged more or less than approximately three times resting or
two-thirds maximum oxygen consumption. An unrealistically high metabolic
correction, 82 percent of maximum on the jig and 84 percent of maximum
in the cage, would be required to equalize the rates for lizards with
the smallest C/E ratios. Thus, Cyclura probably has some physiological
control of its heating and cooling rates.
In the jig, the C/E ratio (Y) was directly related to the ratio of
struggling bouts during cooling vs. heating (X), with the equation:
Y = 0.04809 X + 0.6671 (r = 0.800, p < 0.025, N = 7). This was expected,
since the more struggling that occurred during cooling relative to
heating, the greater the relative difference in metabolic heat pro¬
duction at any one TThe C/E ratio was not correlated with the ratio
of escape attempts or escape attempts plus walking during cooling vs.
heating in the cage. Perhaps, behavioral thermoregulation overrode brief

244
behaviors which increased the cooling rate. However, in one small
iguana, data not included in Table 20, the C/H ratio was 1.496 in the
cage, clearly due to escape attempts during cooling.
Climbing and walking in the cage and struggling on the jig occurred
at significantly higher temperatures during heating than during cooling
(Table 21). This was partly due to the greater amount of time the
lizards were at high T^'s during heating than during cooling. Also,
lizards were actively trying to avoid different unfavorable thermal
conditions during heating and cooling, near the maximum and minimum
voluntary tolerances, respectively. The first defecation during heating
occurred at 5.0 C above the last defecation during cooling, following
the pattern for climbing (11.6 C), walking (1.7 C), and struggling
(5.6). Lizards relaxed on the substrate to the level of the pectoral
girdle at a higher during cooling than during heating, although the
difference was not significant. Head contact with the substrate occurred
for brief periods only during cooling. This increased substrate contact
undoubtedly decreased the cooling rate. Guiar flutter ceased an average
of 9.7 C lower during cooling on the jig than in the cage. Participation
of the intercostal muscles in inhalation was probably reduced by the
jig restraining strap, located immediately posterior to the forelegs,
to the point that supplementary guiar flutter was necessary for adequate
respiration at low 2% 's.
Only two of seven restrained lizards and none of the caged lizards
panted during heating. One animal panted at the end of the run at a
T^ of 39.0 C. The other, Number 3 in Table 20, started at 32.8 C and
continued intermittently during 30.9 percent of the remainder of the
run. The lizard also struggled six times.

Table 21.
Correlations of lizard behaviors with body temperature during heating
and cooling in the cage and on the jig.
BEHAVIOR
HEATING
OR
COOLING
CAGE
OR JIG
x tb
RANGE
SD
ST)cool/Wheat
N/7
LIZARDS
SIGNIFICANCE
LEVELl
Climbing
Heating
Cage
37.0
34.5-38.3
1.524
8
o nos
Climbing
Cooling
Cage
31.6
22.9-37.5
3.800
19
Walking
Heating
Cage
36.4
24.6-38.3
2.952
1 45
25
0 005
Walking
Cooling
Cage
31.2
22.9-38.2
4.276
45
Defecation
Heating
Cage
37.2
34.1-38.7
1.763
9 09
5
mc
Defecation
Cooling
Cage
33.8
29.1-37.6
3.682
6
Struggling
Heating
Jig
35.4
29.3-38.3
2.625
1 74
22
0 005
Struggling
Cooling
Jig
31.4
23.6-38.6
4.564
24
Pectoral Girdle
Heating
Cage
33.9
28.8-37.2
7
NIC
Contacts Substrate
Cooling
Cage
36. 3
31.9-38.5
7
Head Contacts
Substrate
Cooling
Cage
31.7
27.0-36.8
4
Starts Guiar
Heating
Cage
35.0
32.1-37.6
7
MQ
Flutter
Heating
Jig
32.4
24.6-37.9
7
Stops Guiar
Cooling
Cage
33.4
32.1-34.6
7
0 005
Flutter
Cooling
Jig
23.7
23.0-25.7
7
Student t-test for difference in X 2^'s.
245

246
Breathing rates were significantly higher during heating than
during cooling in both the cage and on the jig (Figure 64). Only one
caged lizard had about equal breathing rates. The difference may be
due to thermal hysteresis. Head temperature probably increased faster
than intestinal temperature during heating and decreased faster than
intestinal temperature during cooling. Thus, for any one intestinal
temperature, brain temperature may be considerably higher during heating.
Since the respiratory center is located in the hypothalamus and direct
heating of the area, at least in cats (Nakayama et at. 1963) leads to
an increased respiratory rate, thermal hysteresis may account for the
breathing rate difference. A direct temperature effect on the firing
rate of lung stretch receptors may also be involved. Reduced tidal
volume due to restraint may have caused the higher mean breathing rate
during cooling in the jig than in the cage. Mean breathing rate ratios
of individual lizards during heating and cooling were uncorrelated with
their respective conductance ratios. Thus, unless the convective heat
gain through the lungs during heating and the lower convective heat loss
during cooling were offset by higher and lower evaporative cooling,
respectively, the difference in respiratory rates did not contribute
significantly to the higher lizard heating rates.
The breathing rate during heating in the cage was highly temperature
dependent between 22 and 30 C (Qgg = 4*32) and became nearly temperature
dependent between 30 and 38 C (Q-jg = 1.18), almost encompassing the
preferred T, range. The rate was moderately temperature dependent over
the full cooling range in the cage (Qjg = 2.57 from 38 to 30 C, 2.12 from
30 to 22 C). Lizards were occasionally apneustic (6 of 14 runs) when

247
UJ
!—
ID
CO
50
45
40
35
30
25
2 20
cc
CD
15
!0
5
OH
«»•
20 22 24 26 2 8 30 32 3 4 3 6 3 8 40
BODY TEMPERATURE
Figure 64. Breathing rates of lizards during heating and cooling in a
constant temperature chamber. The solid circles and solid
line refer to the rate during heating, open circles and dashed
line to the rate during cooling in the cage. The dot and
dash and dotted lines refer, respectively, to heating and
cooling respiratory rates on the jig. Lines are running
means for six animals.

248
first placed in the 40 C chamber but rarely so at any other time (1 of
28 runs).
Aerobic Metabolism
A semilog plot of resting metabolism (cm^O^/g'hr)
to a straight line (Figure 65):
Log Mp = 0.0322 Tb - 2.003
vs. T-, was fitted
b
(10)
(N = 30, 13.4 < Tb <_ 43.0 C, r = 0.954)
Resting oxygen consumption in Cyolura was slightly greater than in
Sauromalus hispi-dus and Varanus gouldii, weighing about half as much, on
the average (Bennett 1972). The Cyolura line was also above points plotted
for a 1252 g lizard using equations for body weight vs. resting metabolism
calculated for many lizard species (Bennett and Dawson 1976). However,
the positive 95 percent confidence limits for their 30 and 37 C equations
do exceed the Cyolura line. The iguanas may not have fully recovered
from exercise before resting rates were taken, may have been acclimatized
to lower environmental temperatures (late autumn in Gainesville), or
been stressed by the face mask. Hutchison et aL (1977) reported lower
resting oxygen consumption in Ambystoma trigrdnum when tested in an all
body chamber rather than with a face mask.
The log of a maximum oxygen consumption vs. 2^ was best fitted to
a second degree polynominal, using the statistical technique of Mills
(1955):
Log M = -0.00076 21 2 + 0.0662 T, - 1.6128
s a b b
(N = 30, 13.4 <_ Tb < 43.0 C, r = 0.965)
(ID
Maximum oxygen consumption occurred at 43-44 C, a 1 C extrapolation above
the data. Sauromalus and Cyclura, relatively sluggish herbivores, had
similar curves lying below the linear plot of Varanus, an active carnivore.

Figure 65. Correlation of resting and maximum oxygen consumption
with body temperature. Solid lines and circles refer to
Cyclura aarinata (X weight = 1252 g, N = 5, each animal
run at six 2V' s). Regression lines were constructed using
equations (1Ü) and (11). Values for Sauromatus hisp'idus
(dotted lines,_X weight = 574 g) and Vccranus gouldii
(dashed line, X weight = 674 g) were included for com¬
parison (Bennett 1972). The resting metabolic rates (open
circles) and plus and minus the 95 percent confidence
limit (vertical lines) were calculated for a 1252 g lizard
from equations based on many lizard species (Bennett and
Dawson 1976).

OXYGEN CONSUMPTION-CM/ (G HR)
250

251
Due to metabolic heat production, Cyclura T^'s measured immediately
after seven minutes of activity were nearly always above the air tempera¬
ture of the chamber (mean T^ - Tdifference = 0.8 C, range = -0.6 to
1.9 C, T = 0.987 T, - 0.398).
a d
The Qjo for resting metabolism was constant at 2.10, lacking an
adjustment toward temperature independence at the preferred T\ range
(Figure 66). This is a fairly common characteristic of lizards (Bennett
and Dawson 1976). Unfortunately, M was not measured at 5 and 10 C. A
significant increase in Qln would be expected in this T-^ range in a heat
tolerant species (Dawson 1967). TheQ^0for maximum metabolism declined
from 2.49 between 15 and 20 C to 1.07 between 40 and 43 C. This shift
from temperature dependence below the preferred range to temperature inde¬
pendence at and above the preferred range is characteristic of all
lizards studied thus far, except for several varanids (Bennett and
Dawson 1976). Over most of the T^ range, the thermal sensitivity of
the maximum metabolic rare was less than that of the resting metabolic
rate.
The M[y'Mp ratio or factorial scope nearly reached six between
T-^ s of 20 and 25 C, then declined steadily above 25 C (Figure 66).
The maximum ratio occurred slightly below body temperature of morning
emergence. Increments of five to six-fold at 30 C and three to six¬
fold at preferred body temperature were typical for lizards (Bennett
and Dawson 1976).
The aerobic scope for activity, determined by subtracting the
resting from the maximal metabolic rate at a particular T-^, is a useful
index of the capacity of a species to support a bout of activity with
energy derived from oxygen-dependent pathways. Cycluva's maximum

BODY TEMPERATURE
0.6
or
0.5 x
ó
0.4
E 03
0.2
5
o
i
LlJ
CL
O
O
CO
0.1
0
LU
CD
>-
X
o
Figure 66. Analysis of aerobic metabolism. Oxygen consumption Qjo's for resting (dot and dash line)
and maximally active lizards (dotted line), the ratio of maximum to resting metabolism
(dashed line), and aerobic scope (solid line) are displayed. Horizontal lines and numbers
refer to the ranges of: 1) emergence Tfo's, 2) mean preferred Tu, and mean regulating
phase T¡j's in the field.
ho
Ln
N)

253
aerobic scope occurred between 37 and 39 C [maximum at a Th of 38.1 C;
2
0.47 cm /(g'hr)], corresponding well to its preferred Tof 37 C
(Figure 66). This correspondence has been demonstrated for several
other lizards (Wilson 1974, Bennett and Dawson 1976), but not for
varanids (Bennett 1972) nor two turtles (Gatten 1974a). Aerobic scope
declined to 28 percent of maximum at 15 C, 2.3 C above the critical
thermal minimum, and extrapolated to 22 percent of maximum at the CTMin.
At T^'s of 40 C, scopes for Sauromalus hispidus and Cyclurc oarinata
were about half that of the more active Varanus gouldii [0.47, 0.47, and
3
0.89 cm /(g'hr), respecively; Bennett 1972]. Nonetheless, the predicted
scope for reptiles at a of 30 C and at preferred T^ (Gatten 1978, based
on 20 species) were only 86 and 69 percent of the actual values for
3
Cyctura [predicted scopes of 0.340 and 0.324 cm Ü2/(g*hr) at 30 C and
37 C, respectively; actual values were 0.397 and 0.471; mean body weight
was 1252 g]. Thus, the rock iguana can hardly be called a sluggish
reptile, even though it is a herbivore.
Heart Rate
The points relating the log of resting heart rate to body tempera¬
ture were best fitted to two straight lines meeting at a body temperature
of 25.0 C (Figure 67):
Log HRv= 0.0562 2^-0.0122 (N=15, 13.4£2V_< 26.2 C,r= 0.903) (12)
Log HRr= 0.0370 2^ + 0.4461 (N= 20, 24.9 £ Tb< 43.0 C, v = 0.955) (13)
Body temperature vs. maximum heart rate was fitted best by a second
degree polynomial:
Log = -0. 00069 Tj^ + 0.0782 2% +0.0440 (N = 3a 13.4 < 27^ < 43.0 C, r « 0.987) (14)

200
150
100
80
~60
)540
<
ÃœJ
LÜ
h-
<
£T
(T
<
LÜ
X
4
15
_j i | L_
20 25 30 35
BODY TEMPERATURE
40
45
Figure 67. Heart rate as a function of body temperature. Maximum (open
circles) and resting (solid circles) heart rates are plotted
Regression lines were constructed using equations (12)-(14).

255
The greatest resting and maximum heart rates were recorded at the highes
body temperatures tested.
Resting heart rate Q^ was 3.63 between 15 and 25 C and 2.35
between 25 and 40 C (Figure 68). The change at 25 C fell in the
range for emergence body temperatures and at an expectedly high T^
relative to more cold resistant lizards (Templeton 1970). The 3.63
value was slightly higher than in other lizard species monitored between
body temperatures of 20 and 25 C (Q-^q = 2.0-2.9: Templeton 1970, Wilson
and Lee 1970, Burns 1971, Bennett 1972). The 2.35 value fell in the
range found for other lizards between 25 and 40 C (§ = 1.8-2.5).
Maximum heart rate increase declined steadily between 25 and 40 C but
never reached the high level of body temperature independence achieved
for oxygen consumption. The ratio of active to resting heart rate
remained fairly constant at 1.56-1.69 between 13.4 and 35.0 C, then
dropped to 1.24 at 43.0 C. The heart rate increment (HR -HR ) was
m v
greatest at a T^ of 38.5 C, slightly above the mean preferred body
temperature and within the range of mean field body temperatures.
Licht (1965a) found a similar pattern in four lizard species.
According to the Fick equation, oxygen consumption under steady
state conditions is equal to the product of heart rate, stroke volume,
and the amount of oxygen extracted by the tissues (A-V difference).
Oxygen pulse, defined as oxygen consumption divided by heart rate, is
equal to the product of the stroke volume and the A-V difference.
Resting oxygen pulse in Cycluva declined from 8.3 x 10 ^ to 3.7 x 10 ^
3
cm 09/(g*beat) between 13 and 43 C (Figure 69). The oxygen pulse during
maximum activity declined rather precipitously from 24.9 x 10 ^ to 8.3 x
10 3 between 13 and 43 C. Thus, Cycluva must be quite dependent on

cr
X
s
£
cu
x
JO
40
30
20
10
Figure 68. Heart rate analyses. Heart rate Qjo's for resting (dot and dash line) and maximally
active lizards (dotted line), the ratio of maximum to resting heart rate (dashed line),
and the heart rate increment (solid line) are displayed. Horizontal solid lines refer
to the ranges of: 1) emergence Tfo, 2) mean preferred y¿>, and 3) mean regulating phase
Tfo's in the field.
NJ
ON
HR INCREMENT

LÜ
h-
<
en
h"
en
<
UJ
X
U-
o
o
H-
Z>
m
en
h-
2
O
O
2
UJ
O
en
UJ
o_
UJ
CL
o
o
en
UJ
CD
>-
x
O
o
\—
H
2
UJ
2
UJ
en
o
2
Figure 69. Relationship of oxygen consumption to heart rate at different body temperatures. Resting (dot
and dash line) and maximally active (dotted line) lizard oxygen pulse, oxygen pulse increment
(solid line), and the percent contribution of increased heart rate to increased oxygen trans¬
port during activity (dashed line) are plotted. Curves are constructed using equations (10)-(15).
N)
Ui

258
increased heart rate to provide the increased oxygen demand of the
tissues at higher T^'s, especially during activity. Since active oxygen
pulse declined more rapidly than resting pulse as T^ increased, the
oxygen pulse increment (active pulse - resting pulse) also declined.
At 40 C, active pulse was only 2.5 times greater than resting pulse.
Gatten (1974b) proposed a correction to Bartholomew and Tucker's
(1963) equation for the percent contribution of increased heart rate
to increased oxygen transport during activity:
Percent Contribution =
L
(HR - HR )
a r_
HR _
(M - M )
a r'
M
x 100 (15)
where HR , HR , M , and M are active and resting heart rates and active
a v a r 6
and resting metabolic rates, respectively. The curve calculated for
Cyclura is plotted in Figure 69. Increased heart rate contributed the
greater share vs. oxygen pulse for meeting increased oxygen demand
during activity at 7^'s ^-3 and 37 and the smaller share at 24 C.
The significance of the bimodal character of the curve, if any, is not
known. The contribution of increased heart rate to satisfy the increased
oxygen demand during activity is rather small in comparison to the con¬
tributions of stroke volume and A-V difference at any one T-^. This is
in contrast to the importance of heart rate increase in satisfying
increased oxygen demand as T^ increased oxygen transport during activity
(15-40 at 5 C intervals, N = 6) using the Bartholomew and Tucker
equation were 54.5 (49-65), 21.4 (17.2-24.4), and 14.8 (10-20) for
Sauromalus h-ispidus, Cyclura carinata, and Varanus aould-ii (Bennett
1972). Thus, the Cyolura values are considerably closer to Varanus
than to Sauromatus. The two main reasons for this are the higher resting
heart rate and higher maximum oxygen pulse of Cyalura than Sauromalus.

259
Recoverv from Maximum Exercise
— - d ■ ■ ■ ■■ -
Struggling to escape was spontaneous during the first 1.5-2.0
minutes in the environmental chamber, maintained simply by holding the
animal off the ground. The most numerous and powerful escape attempts,
consisting of clawing, lateral thrashing of the trunk, and rotating or
thrashing of the tail, occurred during the initial 30 minutes (Figure
70). My hand hold had to be changed frequently during the second through
fourth minutes to maintain struggling. Movement became discontinuous
and considerably less forceful during minutes four through nine. Iguanas
were practically helpless during inactive periods. Total activity peaked
at 25 C (sum of movements per 9 minutes = 189, 324, 405, 326, 298, and
236 in Figure 70). This seems to contradict the data indicating a
steadily increasing oxygen debt as 2^ increased. The experiment should
be repeated with a large sample, ideally with each individual exercised
only once at only one 2% . Subjectively, based on the hand exercising
of six iguanas, power output as well as the quickness of individual
movements appeared to increase from 15 C into the preferred range.
Iguanas commonly became briefly apneustic and bloated during
exercise, especially after the initial minute at T-,' s of 25 C and
below. Apnea occurred during the initial seconds of oxygen monitoring
in four of thirty runs. The first peak in the oxygen consumption record
after apnea ceased was disregarded as inaccurately high, and the second
lower peak was taken as the maximum instantaneous rate during the
recovery period. Unfortunately, individuals may have a still greater
metabolic rate during the exercise period, when measurements were not
taken. Bennett and Gleeson (1976) demonstrated that oxygen consumption
in Saeloporus ocaidsntalis maximized during the five minute exercise

260
Figure 70. Temporal pattern of body movements while struggling to escape.
The records are for a single 450 g male tested on six con¬
secutive days. Each tail rotation or thrash, trunk thrash,
and walking movement of opposing fore and rear legs was
counted as one activity unit. Activity occurring each half
minute was converted to a percentage of the total activity
per nine minutes.

261
period at T-, ' s of 25 to 40 C, rather than during the subsequent recovery
period.
The procedure of Bennett (1972) was used to analyze the recovery
records. A smooth curve was drawn through each recovery record,
necessitated by the small volume of the mask. The times required to
reach 75, 60, 50, 40, 30, 25, 20, and 10 percent of each lizard's
aerobic scope were taken from each of the thirty records. Sample means
were calculated for each percent recovery at each T~ . The resulting
recovery patterns at the six T-, ' s were plotted in Figure 71. Recovery
was completed most rapidly at 2^'s of 15-25 C, more slowly at 31-37 C,
and most slowly at 42 C. Return to the resting rate of oxygen consumption
obviously required a long time, an average of 89, 86, 82, 102, 99, and
118 minutes at 2^'s of 15, 20, 25, 31, 37, and 42 C, respectively.
The return of whole body lactate to the resting level probably required
even longer. Sauromalus hispidus had somewhat similar recovery patterns
(Bennett 1972), recovering most rapidly at 25-30 C and least rapidly at
15 and 40 C. The main difference between the results for the two species
was Cyclura's slower recovery at 's of 31 C and higher, especially in
the later minutes. The standard deviation of the mean partial recovery
time at each T-^ increased linearly as recovery time increased. This
was probably due to individual differences in recovery efficiency as
well as unequal work outputs during the exercise period. The sample
size was too small to test for possible sex and size related differences
in recovery rates.
The rather abrupt change in the slopes of the lines in Figure 71
indicates that two or more processes with different equilibration rates
were operating during recovery from exercise (Bennett 1972). The

Figure 71. Effect of body temperature on the rate of recovery from seven
minutes of maximum activity. Lines were fitted by eye to mean
values for five adult Cyolura carinata. Ty is given above
each recovery curve.

PERCENT AEROBIC SCOPE
263

264
elimination of the alactacid debt, the more rapid process, has been
equated with the replacement of oxygen stores in the blood and tissues
and utilization of oxygen during the resynthesis of creatinine phosphate.
The elimination of the lactacid debt corresponds to the conversion of
lactic acid (Margaria et at. 1933). This is probably an oversimplifi¬
cation of the biochemistry involved (Stainsby and Barclay 1970). Never¬
theless, it is still worthwhile to assume a two process recovery when
analyzing the data.
The curves in Figure 69 were approximated by the equation:
- t t
T1 T2
Percent Aerobic Scope = K^e + K.2& “ (16)
where T^ and T0 are the time constants in minutes to 63 percent recovery
of the fast (alactacid) and slow (lactacid) components, respectively, K^
and K9 are the intercepts of the components, t is the recovery time in
cj
minutes, and s is the base of natural logarithms (see Atkins 1969, pp.
101-106, for the method of calculation). The total oxygen debt to 95
percent recovery was estimated by summation of the areas under the curves
3 -1 -1
of the log of the aerobic scope (cm 0„'kg *min ) vs. time for the two
components. The results of these calculations are given in Table 22.
The alactacid debt was much smaller than the lactacid debt at all 2V's.
b
The alactacid debt was practically independent of while the lactacid
debt and, thus, the total debt were highly temperature dependent.
Similar results were found for Sauromatus hispidas (Bennett 1972). How¬
ever, total recovery oxygen was greater in Cyctura at 31 C and above.
This difference is not due to the weight differences of the lizards in
the two samples (X weight = 574 and 1252 g for Sauromatus and Cyclura,
respectively).

Table 22
Recovery oxygen for CyeluPa oaririata after seven minutes of maximum
exercise.
ALACTACID DEBT
LACTACID DEBT
TOTAL OXYGEN DEBT
OR RECOVERY OXYGEN
Body
Temperature
K1
(% Scope)
T1
(Min)
Debt
(cm302/Kg)
k2
(% Scope)
t2
(min)
Debt
(cm^02/Kg)
(cm302/Kg)
15
35.8
1.7
2.5
64.2
25.3
32.9
35.4
20
61.6
5.5
12.7
38.4
37.9
50.1
62.8
25
57.9
3.6
11.0
42.1
30.5
74.0
85.0
31
44.5
2.1
5.8
55.5
35.1
122.5
128. 3
37
43.7
1.8
5.6
56. 3
33.1
170.4
176.0
42
42.8
2.2
6.6
57.2
46.3
215.7
222.3

266
The actual mean recovery times at various 's can be compared
to calculated recovery times based on known mean rates of oxygen con¬
sumption (Table 23). When the iguana is assumed to be metabolizing at
its maximum rate during recovery and using all its oxygen consumption
in the recovery process, an obviously impossible approach, recovery is
achieved an average of 6.2 times faster than actual over the T^ range
of 15 to 42 C. The multiple is not much smaller (X = 4.85) when the
animal is assumed to be metabolizing maximally and reserving its rest¬
ing metabolism for processes other than recovery; that is, using its
full oxygen scope for recovery. The portion of oxygen consumption devoted
to recovery is closer to the resting rate over most of the Tu range, a
mean of 1.2 times the mean resting rate from 15 to 37 C. Thus, assuming
the iguana was totally quiescent during recovery in the dark, a reason¬
able estimate, approximately the overall equivalent of the resting
metabolic rate was devoted to recovery.
Total recovery oxygen to 95 percent of the standard rate was also
calculated directly by measuring the area under the curves of the
3
recorder readouts. A plot of total recovery oxygen (cm 0?/kg) vs.
was best fitted by a second degree polynomial:
:
Log Recovery Oxygen = -0.00046 T^ + 0.0553 T^ + 0.8449 (17)
(N = 27, 13.4 <_ T, < 42.0, 2’ = 0.946)
The for total recovery oxygen decreased from 2.58 between 15 and 16 C
3
to 1.49 between 41 and 42 C. The mean rate of recovery [cm C?/(g"hr)/
min] during the first 5 minutes vs. T^ was best fitted by the
equation:
Log Mean Recovery Rate^ =-0.00127 + 0.0949 T^ - 3.1627 (18)
(N = 29, 13.4 £ Tb < 42.0, r = 0.787)

267
Table 23. Comparison of actual mean recovery time after maximum
exercise with recovery times at three hypothetical
rates of oxygen utilization.
Body Actual Mean Recovery Calculated Recovery Time
Temperature Time (Min) (Actual t Calculated)
Oxygen
Consumption
Rate: Maximum
Scope
Resting
15
89
13.1(6.8)
16.1(5.5)
70.3(1.3)
20
86
14.8(5.8)
17.8(4.8)
86.1(1.0)
25
82
13.8(5.9)
16.7(4.9)
80.5(1.0)
31
102
15.0(6.8)
18.6(5.5)
77.8(1.3)
37
99
16.9(5.9)
22.4(4.4)
68.4(1.4)
42
118
19.9(5.9)
29.8(4.0)
59.7(2.0)

268
The rate became practically independent of T^ near the preferred range
of the lizard (Figure 72). The mean recovery rate during the latter part
of the recovery period, from minute 15 to the return to the resting rate
of oxygen consumption, vs. was best fitted by the equation:
Log Mean Recovery Rate.-, = 0.0228 T^ - 3.4725 (19)
(N = 29, 13.4 <_ Tb < 42.0, r> = 0.805)
Thus, the mean rate of late recovery increased linearly with increasing

MEAN RATE OF RECOVERY (CM02/(GHR))/MIN
269
0.1
0.08
•
0.06
•
•
• •
•
•
i
0.04
« * ^
0.02
i
\ *
• \
•
•
• •
0.01
0.008
• /
• /
' 4 . *
0.006
0.004
% *
0.002
.. . •
• • •
•
0.001
0.C008
i 1
\ •
\ *
•\
» \
•A
• • • \
8
0.0006
-
0.0004
•
0.0002
i i i i i 1 1—
10 !5 20 25 30 35 40 45
BODY TEMPERATURE
Figure 72,
Early and late phase recovery from maximum exercise in CyeZuva
car-inata at different body temperatures. The mean rate of
change in oxygen consumption from the cessation of exercise
to the end of the fifth minute of recovery (upper points) and
from the fifteenth minute to the resting metabolic rate (lower
points) are plotted against Tfc. Curves were drawn using
equations (18) and (19), respectively.

DISCUSSION
Body Temperature Relations
Comparison of Cyclura. carinata's Mean Body Temperature with Other
Diurnal Lizards
In order to rank CycZura carinata according to its relative degree
of heliothermy, field body temperatures of diurnal lizards from five
families have been taken from the literature and catalogued in Table 24.
Unfortunately, interspecies 2^ comparisons are not completely reliable,
since the data were collected under a variety of conditions. 's were
measured in different seasons. All occupied habitats may not have been
sampled adequately. Some lizards may not have reached regulating phase
T^’s when they were captured. Some samples may be biased by collecting
too many lizards displaying a particular behavior; for example, mainly
perched individuals (Licht et al. 1966, Bradshaw and Main 1968). None¬
theless, as the number of species for which data is available increases,
the likelihood of making an incorrect statement about the relative degree
of heliothermy of any one species decreases.
Cyolura has a high T^ in comparison to most other diurnally active
lizards. Of the skinks, 29 of 30 species fall below Cyolura's mean T^
range of 38.0-39.7 C, only one species is in the range, and none are
above it (29, 1, 0). Cyolura's relative standing in the other families
are: iguanidae (47, 6, 1), varanidae (2, 1, 0), agamidae (5, 3, 1),
teiidae (5, 1, 8), overall (89, 11, 11, or 80.2, 9.9, 9.9 percent. Only
in the teiidae do the majority of the 21,.' s lie above Cyolura's.
270

Table 24. Field body temperatures of diurnally active lizards. Species in each
family are arranged according to increasing mean body temperature.
Lizards which are herbivorous as adults are marked with an asterisk.
SPECIES
x tb
SE
RANGE
SOURCE
ACAMIDAE
Molo oh horridus
34.0
27.0-40.0
Pianka and Pianka 1970
Arnphibolurus minimus
34.2
0. 31
32.6-37.4
Bradshaw and Main 1967
Arnphibolurus barbatus
34.8
0.37
30.0-39.7
Lee and Badham 1963
Arnphibolurus omatus
36.6
0.23
33.0-41.8
Bradshaw and Main 1967
Arnphibolurus retioulatus
37.0
0.90
35.0-40.6
Licht et al. 1966
Arnphibolurus isolepis
38.8
0. 3
37.8-39.3
Pianka 1971b
Arnphibolurus oaudioincbus
39.0
0.48
34.8-41.0
Licht et al. 1966
Arnphibolurus inermis
36.1
Pianka 1971a
36. 7
0.53
34.0-43.8
Bradshaw and Main 1967
39.3
Licht et al. 1966
Diporophora. bilineata
44.3
0. 33
42.3-46.0
Bradshaw and Main 1967
(Lizards arboreally active at noon)
IGUANIDAE
Anolis ooulatus
27.4
21.7-32.0
Ruibal and Philobosian 1970
28.2
0.17
23.2-33.0
Brooks 1968
Anolis limifrons
28.5
0.59
25.6-31.0
Ballinger et al. 1969
Anolis allogus
29.2
0.06
26.2-33.5
Ruibal 1961
Anolis luoius
29. 3
0.10
24.8-32.4
Ruibal 1961
Anolis aoutus
30.3
25.6-34.8
McManus and Nellis 1973
* Basilisous plumifrons
31.7
0.50
28.0-35.5
Mirth 1964
Anolis homoleohis
31.8
0.14
26.2-35.0
Ruibal 1961
* Conolophus suboristatus
32.0
25.7-37.5
Carpenter 1969
(Taken during intermittent overcast)
Anolis allisoni
33.0
0.21
28.2-36.6
Ruibal 1961
Anolis sagrei
33.1
0.22
27.4-36.1
Ruibal 1961
K>

Table 24. Continued
SPECIES X Tb SE RANCE SOURCE
IGUANIDAE (Continued)
Saeloporus occidentalis 33.4
Saeloporus merriami 33.6
Saeloporus gracilis 33.6
Saeloporus poinsetti 34.2
Phrynosoma solare 34.7
Basilisaus vittatus 34.7
Uta thalassina 34.8
Saeloporus magister 34.8
Saeloporus undulatus 34.8
Phrynosoma coronation 34.2
34.9
Saeloporus jarrovi 35.0
Urosaurus nigrioaudus 35.0
* Conolophus pallidus 35.1
Saeloporus graciosus 35.2
Saeloporus squamosus 35.3
* Cyclura pinguis 35.3
Urosaurus omatus 35.5
Sator grandaevus 35.5
Holbrookia maculata 35.7
Saeloporus orautti 35.9
Saeloporus olivaaea 36.0
Urosaurus graaiosus 36.1
* Iguaria iguana 36.1
36.1
Uta stansburiana 35.0
36.2
Saeloporus woodi 36.2
Urosaurus auriculatus 86.3
Urosaurus clarionensis 36.4
* Cyclura oornuta stejnegeri 36.8
0.30
15.2-38.8
Cunningham 1966
29.6-37.4
Brattstrom 1965
30.3-39.1
Brattstrom 1965
30.8-38.4
Brattstrom 1965
Baharav 1971
0.07
32.6-36.5
Hirth 1964
0.40
32.6-38.5
Soule 1963
0.17
Parker and Pianka 1973
25.0-38.9
Brattstrom 1965
0.69
20.8-40.6
Cunningham 1966
28.0-43.0
Heath 1965
32.2-37.0
Brattstrom 1965
0.51
33.8-39.5
Soule 1963
31.6-37.1
Carpenter 1969
0.24
22.7-39.2
Cunningham 1966
32,5-38.0
Brattstrom 1965
27.1-39.8
Carey 1975
26.8-39.5
Brattstrom 1965
0.42
30.2-39.5
Soule 1963
40.8
Clark 1965
0.32
32.6-38.5
Soule 1963
Brattstrom 1965
0.26
Pianka and Pianka 1970
34.0-37.2
Hirth 1963
42.4
McGinnis and Brown 1966
0.29
20.9-40.6
Cunningham 1966
0.24
32,1-39.8
Soule 1963
0.25
Bogert 1949a
32.3-39.0
Brattstrom 1965
29.6-39.0
Brattstrom 1965
35.8-37.8
Wiewandt 1977
272

Table 24. Continued.
SPECIES
X Tb se
RANGE
SOURCE
IGUANIDAE (Con
tinued)
Phrynosorna platyrhinos
36.8
29.0-43.0
Heath 1965
Sceloporus varibilis
36.9
33.6-40.0
Brattstrom 1965
* Ctenosaura hemilop}ia
37.1 0.53
32.0-39.6
Soule 1963
* Ctenosaura similis
37.1
34.5-40.1
Fitch 1973
Crotaphytus oollaris
37.2
20.7-43.3
Brattstrom 1965
IJma sooparia
37.3 0.40
Pianka and Pianka 1970
* Sauromalus obesus
35.3
41.9
Johnson 1965
37.7
42.0
Cowles and Bogert 1944
46.0
Case 1976
* Amblyrhynchus cristatus
32.5 0.57
26.7-37.5
Bartholomew 1966
(Lizards in the prostrate basking
pos ture)
37.9 0.33
36.2-40,0
Bartholomew 1966
(Lizards in the elevated basking posture)
Holbrookia propinqua
37.9
40.9
Judd 1975
Holbrookia laceraba
38.1
42.5
Clark 1965
Phrynosoma m'calli
38.3
43.0
Heath 1965
* Uromastix acanthinurus
38.0-39.5
21.0-46.0
Grenot 1976
Urna no tata
38.8
35.0-40,5
Cowles and Bogert 1944
* Cyclura carinata
38.0-39.7
33.7-42.5
Present study
(Free-ranging lizards only)
Crotaphytus wislizeni
39.2 0.32
36.5-41.8
Cummingham 1966
Callisaurus draconoides
38.6 0.24
34.6-41.4
Soule 1963
39.4 0.76
36.4-42.4
Cunningham 1966
* Dipsosaurus dorsalis
40.9 0.44
34.0-43.6
Cunningham 1966
42.1
38.0-46.4
Norris 1953
SCTHCIDAE
Eumeces anthracinus
25.0
33.9-33.0
Brattstrom 1965
Eumeces skiltonianus
25.0 1.36
16.8-34.0
Cunningham 1966
Sphenomorphus sabanus
25.8
24.0-28.4
Brattstrom 1965
Eumeces gilberti
26.7
21.5-31.0
Brattstrom 1965
Eumeces brevilineatus
28.0
Erattstrom 1965
273

Table. 24. Continued
SPECIES
X tb
SE
RANGE
SOURCE
SCINCIDAE (Continued)
Lygosoma septentrionalis
28.5
24.4-32.0
Brattstrom 1965
Lygosoma laterale
28.8
22.0-35.5
Brattstrom 1965
Leiolopisma rhomboidalis
28.9
0.58
24.2-35.0
Wilhoft 1961
Eumeoes latioeps
29.3
22.8-33.2
Brattstrom 1965
Sphenomorphus tympanum
29.4
18.2-36.5
Spellerberg 1972c
Sphenomoxrphus quoyi
30.0
17.4-34.2
Spellerberg 1972c
Ctenotus brooksi
30.6
0. 36
Pianka 1969
Leiolopisma fusoum
30.8
0. 32
24.5-34.5
Wilhoft 1961
Sphenomorphus kosoiuskoi
30.9
16.9-36.0
Spellerberg 1972c
Ctenotus dux
32.0
0.35
Pianka 1969
Tiliqua soinaoides
32.6
39.5
Bartholomew et al
Tiliqua rugosa
32.7
27.0-36.5
Licht et al. 1965
Mabuya rudis
32.8
25.4-38.6
Brattstrom 1965
Ctenotus helenae
32.8
0.52
Pianka 1969
Eumeoes fasoiatus
33.0
13.5-37.0
Brattstrom 1965
Mabuya mabouia
33.0
0.43
30.1-37.7
Brooks 1968
Ctenotus pantherinus
33.1
0.38
Pianka 1969
Ctenotus schornburgkii
33. 3
0. 39
Pianka 1969
Eumeoes obsoletus
34.0
17.5-36.3
Brattstrom 1965
Ctenotus granáis
34.2
0.51
Pianka 1969
Ctenotus atlas
34.5
0.83
Pianka 1969
Ctenotus calurus
35.6
0. 36
Pianka 1969
Ctenotus quattuordeoimlineatus
35.8
0.30
Pianka 1969
Mabuya striata
35.8
31.0-39.5
Brattstrom 1965
Ctenotus leae
37.7
0.41
Pianka 1969
Ctenotus leonhardii
38.0
0.20
Pianka 1969
TEIIOAE
Ameiva fusoata
36.1
0. 28
26.0-40.1
Brooks 1968
Atneiva festiva
36.2
0.26
32.0-39.8
Ilirth 1964
Ameiva ameiva
37.6
35.1-39.3
Brattstrom 1965
274

Table 24. Continued
SPECIES
X tb
SE
RANGE
SOURCE
TEIIDAE (Continued)
Arneiva quadrilineata
37.6
0.06
34.6-40.0
Hirth 1964
Ameiva pluvianotata
37.8
0. 34
33.8-40.0
Hirth 1964
Cnemidophorus lemnisoatus
38.8
34.5-42.3
Brattstrom 1965
Cnemidophorus hyperythrus
39.9
0.25
38.3-41.6
Soule 1963
Cnemidophorus exsanguis
39.9
0.16
Schall 1977
Cnemidophorus oeralbensis
40.0
0.45
37.4-42.5
Soule 1963
Cnemidophorus inornatus
40.2
0.10
Schall 1977
Cnemidophorus gularis
40.2
0.16
Schall 1977
Cnemidophorus tigris
39.4
0.38
20.0-44.6
Cunningham 1966
40.4
0.09
Schall 1977
Cnemidophorus sexlineatus
41.0
0.47
38.5-43.0
Bogert 1949
Cnemidophorus tessellatus
40.1
0.13
Schall 1977
41.3
0.24
37.4-43,5
Bogert 1949b
VARANIDAE
Varanus varius
35.5
35.0-36.2
Stebbins and Barwick 1968
Varanus gouldii
37.1
0.76
34.4-36.2
Licht et al. 1966
Varanus komodoensis
38.1
36.0-40.0
McNab and Auffenberg 1976

276
Do herbivorous lizards require higher 2^'s to digest their food?
Janzen (1973) has suggested that the teiids, Ameiva ameiva and Cnemido-
phorus lemniscatus, insectivorous on the mainland of Costa Rica and
primarily fruit and vegetation eaters on the island of Providencia,
have elevated their 2V on the island as a result of the difference in
b
diet. The island lizards "must maintain the internal compost heap at a
high temperature in order to process foliage fast enough to get the
energy needed." Jaiizen made no T^ measurements of mainland and island
forms, however, only stating that island forms basked more frequently.
The data in Table 24 are also suggestive. The mean active for 23
genera of primarily carnivorous lizards from four families is 34.8 C
(X + 2 SE = 36.1 C), lower but not significantly lower than the mean for
nine genera of herbivorous iguanids, 36.8 C (X + 2 SE = 37.3). Apparent
or gross digestive efficiency is probably significantly lower in
herbivorous than in carnivorous lizards, especially when leaf vs.
insect eaters are compared (Shine 1971, Iverson 1977). The digestive
efficiency for fruits and flowers, with their lower fiber content, is
probably higher than for leaves, at least in Cyclura (Auffenberg, in
prep.). Both Cyclura stejnegeri (Wiewandt 1977) and Cyclura carinata
prefer fruits to leaves in the field.
Digestive efficiency improves as the preferred T^ is reached in
Dipsosaurus dorsalis (Harlow et al. 1976). A higher field 2^, as expressed
during a long basking period, may compensate for the lower digestibility
of foliage. However, it is premature to state this absolutely.
Seasonal Variation in Body Temperatures of Active Lizards
As available ambient temperatures and radiant heat input gradually
decline in the fall of the year, diurnal heliothermic lizards must do at

277
least one of three things: 1) spend a greater proportion of their day
basking rather than foraging, etc., 2) lower their T-^ during activity,
or 3) brumate. Heliotherms in the subtropics and tropics do not brumate
while temperate species do, such as Dipsosaurus dorsalis and Sauromalus
obesus. Seasonal mean ranges of active lizards which do not brumate,
3.4 C in Amphibolurus isolepis (tropical Western Australia, Pianka 1971b),
5.5 C in Soeloporus ocoidentalis (Berkeley, Calif., Mayhew 1963), may
usually be greater than for active lizards that do brumate; for example,
2.5 C in Amphibolurus fordi (New South Wales, Australia, Cogger 1974).
The Cyolura caminata range was only 0.5 C at Site II for February to
October. However, December-January MBT's for active lizards probably
fell below 38.9 C. The MBT of Cyolura oychlura figginsi, measured in
January between 1000 and 1400 on Great Exuma in the Bahamas approximately
120 miles north of Water Cay, averaged only 31.4 C (Windrow 1977). How¬
ever, some of Windrow's values may have been obtained from lizards in the
morning heating phase. One clear indication that C. oarinata becomes
more eurythermal in the winter is the significantly lower mean February
Th at the start of foraging. Based on the thermal gradient results,
the seasonal lowering of the MBT is probably not due to a decline in the
preferred Tcaused by thermal acclimatization or shorter photoperiod.
However, a reduced winter food supply, resulting in a lowered intake,
may be partially responsible for any MBT decline.
Stability of the Mean Preferred Body Temperature
The stability of the MPT varies in different lizard species.
However, insufficient data are available to determine whether temperate
species have less stable MPT's than tropical species.

278
To test for seasonal acclimatization stability of the MPT, lizards
are collected from the field at regular intervals and tested immediately
in a photothermal gradient. McGinnis (1966) found no change in the MPT
of Scelcporus occidentalis using this technique. March and September
results for Anolis carolinensis were not significantly different (Lic'nt
1968). In the present study, winter and summer samples of Cyclura
caminata had about the same MPT’s. On the contrary, the MPT in Sauro-
malus obesus varied by more than 2.5 C during a 2 year period, with no
logical pattern to the variation (Case 1976). Lacerta vivípara MPT's,
measured in southern England, varied by 5.1 C between April and October
(Patterson and Davies 1978).
The commonly used technique to determine acclimation stability of
the MPT is to collect a single sample, divide it into several groups,
keep each group at a different acclimation temperature in a controlled
environmental chamber for at least 2 weeks, and then test individuals
in a photothermal gradient. Results from different species are inconsist¬
ent, as are the seasonal acclimatization results. Eumeces laticeps had
equal MPT’s after acclimation at 14, 22, and 28 C (Pentecost 1974).
Other workers have reported inverse acclimations; Wilhoft and Anderson
(Scelopcrus occidentalis 1960) and Mueller (5. occidentalis and the
California sample of S. graciosus 1970) found a lowering of the MPT in
their 35 C groups. The responses may have been pathological, since
35 C is above the mean of the field caught lizards. Licht (1967)
found a slightly lower MPT in Anolis acclimated at 32 C than at 20 C.
The difference was significant only during the second of three days in
the gradient. Mueller (1970) reported a higher MPT in the 15 C

279
acclimation group than in the 25 and 35 C groups of Wyoming caught
Soeloporus graoiosus.
Even if the MPT is fairly stable over a wide range of acclimation
temperatures, thermoregulatory precision is commonly greater after low
temperature acclimation. McGinnis's winter Soeloporus sample had a
narrower T^ range than his summer sample (31.7-37.3 vs. 28.6-38.7 C,
respectively). The same was demonstrated for Cyclura in the present
study. Pentecost's Evmeoes acclimated to 14 C did not reach 's as
high as lizards in the 22 and 28 C groups (38.3 vs. 42.8 and 40.7 C,
respectively). In accord with these results, low temperature acclimation
decreases the critical thermal maximum in lizards (Lowe and Vance 1955;
Ballinger and Schrank 1970, Corn 1971).
Late Afternoon Maximum Body Temperature
Adult male and female Cyolura spent considerable time in the
vicinity of the nocturnal retreat after completion of the final foray.
The post-foray period averaged 97 minutes in February (SD = 61, Range =
17-191, N = 11 days) and 86 minutes in October (SD = 41, Range = 12-140,
N = 10 days) and started at a mean time of 1449 (Range = 1336-1548) and
1416 (Range = 1312-1530), respectively. Much of this time was spent
basking in one spot and often led to the highest T- recorded during the
day, especially during the hot summer months. The highest maximum
voluntary tolerance, 43.8 C, was recorded during this time, a temperature
above the panting thresholds determined earlier in the day, indicating
an increased late afternoon high temperature tolerance. Lizards commonly
selected the same wind protected location night after night and always
assumed the prostrate basking posture. Dorsal cutaneous vasodilation and

280
ventral cutaneous vasoconstriction were likely during this time (White
1973).
The morning and afternoon 's of many small lizard species are
not significantly different: Anolis oristatellus (Huey 1974), Amphibolurus
barbatus (Lee and Badham 1963), Amphibolurus inermis (Heatwole 1970),
Cnemidophoms hyperythrus (Bostic 1966), Holbrookia pvopinqua (Judd
1975), and Soeloporus orcutti (Mayhew 1963). All five tropical Anolis
species examined by Ruibal (1961) had significantly higher afternoon
2^'s. This may have been due to higher afternoon air temperatures, as
he suggested, or to including morning heating phase records in the morning
sample.
Reptiles reported to maximize late in the afternoon are all
large: Conolophus suboristatus (Werner, D., personal communication),
Geochelone elephantopus (Mackay 1964), Sauromalus obesus (Case 1976),
Varanus komodoensis (McNab and Auffenberg 1976), and Varanus varius
(Stebbins and Barwick 1968). These reports are all based on small sample
sizes, however. Late afternoon T^ maxima are probably more common in
large reptiles due to the physics of heat exchange. Of course, the
lizard must also voluntarily tolerate high afternoon 2^'s. Daily late
afternoon T^ maxima have also been reported in a number of large mammals;
camel (Schmidt-Nielsen et al. 1957), camel, giraffe, w^ater buffalo, eland,
oryx, and Ankole steers (Bligh and Harthoorn 1965), and euro, gray and
red kangaroos (Brown and Dawson 1977).
The late afternoon maximum T^ may be an example of temperature
hardening, defined by Precht (1973) as "a quick, usually transitory,
adaptation to high and low temperatures." Earlier exposure to high
temperatures during the day may have resulted in a short term increase

281
in heat tolerance, lost during the subsequent night. Heat hardening
(see Hutchison and Maness 1978 for a review) is not the same as
acclimatization, a longer lasting response to temperatures in the
normal range. Strictly speaking, the iguanas had not been heat hardened,
since Tjj was always below the CTMax. However, Hutchison and Maness'
definition may be found to be too restrictive when the biochemical
basis of the phenomenon is finally determined. Of course, a circadian
cycle of temperature tolerance may also be involved. Hovever, this
possibility was not investigated in the present study.
Thermal Safety Margins
Heatwole (1970) defined six thermal safety margins: 1) PTSM-CTM,
a physiological thermal safety margin; the difference between the
critical thermal maximum and the mean T^ in a thermal gradient, 2)
PTSM-LM, a physiological thermal safety margin; the difference between
the lethal maximum and the mean in a thermal gradient, 3) ETSM-CTM,
an ecological thermal safety margin; the difference between the CTM
and the mean regulating phase, field 2L,, 4) ETSM-LM, another ecological
thermal safety margin; the difference between the LM and the mean
regulating phase, field Th, 5) MTSM-CTM, a minimum thermal safety
margin; the difference between the CTM and the maximum field T, and
6) MTSM-LM, another minimum thermal safety margin; the difference
between the LM and the maximum field T. The values for Cyatura are:
PTSM-CTM, 9.2 C; PTSM-LM, 11.0; ETSM-CTM, 6.3 and 8.1 (June and July);
ETSM-LM, 8.3 and 9.9; MTSM-CTM, 3.7 and 5.5; MTSM-LM, 5.5 and 7.3.
These values are slightly smaller than for the majority of the 39 species
of lizards listed by Heatwole; that is, 61 margins are larger, 38 are in

282
range, and only 14 are smaller. This is further evidence that Cyolura
maximizes its field Tn's in the summer. Heatwole suggested that this
is possible in tropical species due to the predictable nonvarying nature
of the upper limit of environmental temperatures.
Panting Threshold
The factors influencing the level of the panting threshold have been
studied intensively by Heatwole and his coworkers (Chong et at. 1973,
Heatwole et at. 1973, Heatwole et at. 1975). They found the threshold
to be quite plastic. It has a diel cycle corresponding to the daily
rhythm of environmental temperature. Seasonal variation is expressed
as higher summer values than at other times of the year, related both
to higher summer acclimatization temperature and longer photoperiod. The
immediate past thermal history of the lizard affects the threshold, that
is, repeated heatings to panting depressed the value. Lastly, they
found that random day to day individual variation, independent of environ¬
mental temperatures, is quite important.
The T£ range for the panting threshold in Cyolura oarinata was
larger than any listed by Heatwole et at. (Table 1, 1973). The only
value approaching 11 C was that for Arrrphibolurus barbatus (31.5-42.4 or
10.7 C). One probable reason for this difference is the common use of
a low heating rate for the determination of the panting threshold,
which prevents unequal heat distribution in the reptile's body.
Templeton (1971) increased the panting threshold of Dipsosaurus dorsalis
from 39.0 to 43.5 C by decreasing the heating rate. However, rapid heat¬
ing of Amphibolurus muricatus did not depress the panting threshold,
though it did increase the variability of the results (Heatwole et at.

233
1973). This probably was due to heat damage, since eight of ten
lizards in their sample failed to recover. The A. muricatus threshold
decreased as much as 1.3 C during ten consecutive heatings of a single
individual over a period of 3 hours. The same phenomenon may have
occurred in the Cyclura carinata results illustrated in Figure 57,
although the threshold would presumably not have risen again in the
afternoon.
The most likely explanation for the midday depression of head and
body panting threshold temperatures is the importance of a skin, tempera¬
ture sensory input, as demonstrated in Sauromalus obesas by Crawford
and Barber (1974). The stomach temperature at the panting threshold
in Varanus alblgularls ranged from 35.4 to 38.6 C, whereas skin tempera¬
ture remained near 39 C (Bligh et at. 1976), indicating that the latter
was being regulated. Since the skin-core temperature difference is
greatest during midday heating, the core depression in Cyclura would
be explained if skin temperature is also of major importance in the
control of the panting threshold.
It should be standard procedure to find the maximum panting
threshold range for each lizard species, as Spellerberg (1972a) suggested
for the CTMin, since some interesting ecological correlates would be
expected. This demands acclimation of lizards to a wide range of
ambient temperatures prior to testing, as well as immediate testing of
acclimatized lizards. A desert dwelling lizard might have a wider
panting threshold range, an adaptation to both large temperature extremes
and low water availability. Amphibolurus barbatus, occupying both arid
and mesic habitats, did elevate its panting threshold in response to
dehydration, adaptive as a wTater conserving measure in hot dry weather.

284
On the other hand, A. muriaatus, a dweller of more mesic regions, did
not alter its threshold T-^ with increasing dehydration (Parmenter and
Heatwole 1975). Secondly, maximum panting threshold ranges of helio¬
therms may be greater than for thigmotherms and nocturnal lizard species,
which are subjected to less rapid heating and cooling rates. Thirdly,
the threshold range might be expected to rise ontogeneticaliy in
species with large adult size, due to increasing temperature differences
between the skin and the body core with increasing body weight.
Body Temperature and Reproductive State
Reproductively active, ovoviviparous, female snakes appear to have
higher preferred field 's during activity than males and reproductively
quiescent females. Mean 's of Crotalus uiridis females were 2.1 and
2.0 C above males in the spring and summer, respectively (Hirth and
King 1969). The 's of two oviparous species, Mastioophis taeniatus
and Coluber* oonstrdatov, did not differ significantly with sex. Pregnant
Tharrmophis sirtalis and Tharmophis ovdinoides both had higher mean TVs
chan males (2.3 and 2.2 C, respectively) and nonpregnant females (0.6
and 1.0 C) in a thermal gradient (Stewart 1965). Pregnant field caught
female Agkistrodon contovtrix had higher activity T^'s than males
(Fitch 1960). An oviparous female Python moluvus bivittatus elevated its
T->d an average of 6.2 C (Range 4.1-7.4) above substrate temperature during
brooding (Van Mierop and Barnard 1976). Also, T^ was higher above
substrate temperature in the gravid snake (X = 3.1, Range 1.5-5.5)
than in the same snake when postbrooding (X = 1.1 C, Range 0.1-1.7).
The pattern may be reversed in ovoviviparous and at least some
oviparous lizards. Males and preovulatory and postparturient females of

285
Sceloporus cyanogenys and ovoviviparous species, all had higher 's
than pregnant females in a thermal gradient (Garrick 1974). Pregnant
ovoviviparous female Lacerta vivípara had lowered preferred 21/ s in a
thermal gradient than males (Patterson and Davies 1975). Gravid females
of five species of oviparous Cnemidophorus all had lower mean 2// s than
males and nongravid females, though the differences were not significant
(Schall 1977). Gravid Cyclura carinata appear to have depressed regulat¬
ing phase 21/s. Mayhew (1963), Bostic (1966), Brooks (1968), and Heatwcle
(1970) found no significant difference in mean 2^/ s of males and females
of six oviparous lizard species. However, none of these workers
examined the reproductive state of the females in their samples. This
is necessary since all mature females may not reproduce in a given year,
as Nagy (1973) found for Sauromalus obesus, and the period of gravid
female hypothermia may be of short duration.
In the snakes, the elevated embryo temperature may prevent ab¬
normal development (Fox et at. 1961) and certainly increases the rate
of normal development. In the lizards, oviducal embryos are presumably
more sensitive to high temperatures than the tissues of the parents.
Ethological Considerations
Lizard Activity
Cyclura is closer to a Type I than a Type II lizard (Heatwole et
at. 1969); strictly diurnal, emergent only when T. rises above burrow
temperature, and rather stenothermal when weather permits. Cyclura
tailors its activity period to the yearly change in pliotoperiod, emerging,
on the average, 3 hours after sunrise and submerging an hour and a

286
half before sunset, regardless of season. Few forced submergences
resulted from overcast, due to the usual short duration of the overcast
periods and the low cooling rate of lizards due to large body size.
Forced submergences due to high environmental temperatures were also
rare in well vegetated areas.
Many small, burrow utilizing, diurnal heliotherms, presumably as
a result of their rapid heating rates or greater eurythermality, emerge
earlier than Cyclura during warm weather. Preemergence summer basking
of Amphibolurus fordi in Triodia grass tussocks began very soon after
sunrise and postemergent basking outside the tussocks started around
0715 (Cogger 1974). Uma sp. emerged as early as 0500 in July (Mayhew
1964), and Amphibolurus inermis started emerging around 0700 in summer
(Bradshaw and Main 1967). Dipsosaurus dorsalis may be a unique case;
it becomes active in its burrow up to two hours before emergence (DeWitt
1963, Gelderloos 1976), moving to the shallow portion of the burrow
where it heats to the minimum voluntary tolerance of the regulating
phase (McGinnis and Dickson 1967). It emerges late in the day compared
to other desert lizards (Norris 1953). Type II lizards, such as many
Anolis species, remain above ground all night and become active almost
at sunrise. For example, many Amblyrhynchus oristatus had already
finished basking and were ready to enter the sea by 0730-0800 (Carpenter
1966). However, not all diurnal lizards emerge early in the morning.
As size increased in thirteen species of Ctenotus, some of which were
sympatric, activity patterns changed in a distinct fashion (Pianka 1969).
The smallest lizard group, consisting of four species, had short,
midday-encompassing activity periods. The next largest group, also

287
consisting of four species, had longer bimodal activity distributions.
Only the five largest species were active all day.
Diurnal lizards usually have a single activity peak. Unfortunately,
the quantitative measure of activity is usually a tally of how many
lizards are seen or captured per hour, rather than of individual lizard
movement. The activity peak most commonly occurs sometimes during the
morning hours during warm weather (Heath 1965, Johnson 1965, Tanner and
Krogh 1974, Gelderloos 1976, present study). A morning activity peak
seems adaptive on several grounds: 1) it occurs before the surface
environment becomes too hot, 2) feeding is completed as soon as possible,
leaving the rest of the high T-^ time for digestion, 3) territorial
encounters, if any, would be energetically wasteful to continue through¬
out the day. High environmental temperatures commonly result in a
bimodal activity distribution. The daily maximum activity peak some¬
times occurs in the late afternoon rather than during the morning under
these conditions {Uta stansburiana, Irwin 1965; Sauromalus obesus,
Nagy 1973 and Case 1976; Gopherus berlandieri, Rose and Judd 1975).
Cyctura remains emergent during the full time environmental
temperatures are suitable for maintaining preferred even during the
dry season. Dipsosaurus dorsalis, on the contrary, does not reemerge
during summer afternoons, even though environmental temperatures are
suitable for it to do so. Norris (1953) and Porter et at. (1973) have
suggested that this lack of afternoon activity is a water conserving
measure. Water loss is undoubtedly less of a problem for Cyolura than
for Dipsosaurus. Rain may not fall for more than 15 months in parts
of the range of Dipsosaurus, whereas rainless months in the Caicos are
rare. Also, the dry season in the Caicos is cool to warm, rather than

288
hot as in North American deserts. Thirdly, Cyclura probably has the
same passive means of reducing respiratory water loss as Dipsosaurus
(Murrish and Schmidt-Nielsen 1970) ; a portion of the water vapor added
to the inspiratory air by evaporation is recovered by condensation on
the cool nasal passages during exhalation. Fourthly, since it possesses
a nasal salt gland, Cyclura probably also decreases water loss during
the dry season by cloacal water resorption from the urine (Schmidt-
Nielsen et al. 1963).
Eurythermy vs. Stenothermy
Eurythermic lizards spend little or no time thermoregulating.
Since T^ and shaded air temperature are very similar, eurytherms are
sometimes referred to as thermoconformers or as thermal generalists.
Examples are most commonly found in heavily shaded, tropical forest
regions: Spenomorphus sabanas (Inger 1959), Anolis oculatus (Ruibal
and Philobosian 1970), Anolis tvopidolepis (Fitch 1972), and Anolis
polylepis (Hertz 1974). Other eurythermic lizards are slightly less air
and substrate temperature dependent, occurring in more open areas:
Gerrhonotus multicarinatus (Cunningham 1966) and Phrynosoma platyrhinos
(Pianka and Parker 1975). Stenothermic lizards thermoregulate, main¬
taining T^ in a narrow range during the day in the face of changing
environmental temperatures. These thermal specialists constitute the
bulk of the diurnal lizard fauna in Australian, North American, and
southern African deserts.
Large body size and the nature of the Caicos Island environment
probably restrict Cyclura to a semiarboreal herbivorous niche. Due to
its large body size and the low prey density, it is unlikely that adult

289
Cyclura could survive on a diet of insects. The three small, diurnally
active lizard species present are all insectivorous; Mabuya mabouya,
occupying the shaded leaf litter habitat; Ano Its scrip tus, occupying
the shaded arboreal habitat, and Liocephalus psammodrommus, living on the
ground and preferring more open areas than Mabuya. Liocephalus has only
a slightly lower activity temperature than Cycluva (X = 36.4 C,
range = 35.1-38.0, N = 15 September captures). The nearly totally
herbivorous diet of even hatchling Cyclura (Iverson 1979) may be due to
competitive exclusion by the three insectivorous species. The height
and density of the Caicos vegetation are too low in most areas for
Cyclvo'a to be as arboreal as Iguana iguana. Due to its dependence on
anaerobic metabolism during maximum activity and the resulting rapid
onset of fatigue, it is doubtful that Cycluva could subdue a mammal
much larger than a rat. But mammals have been absent from the Caicos
until very recently. Although several species of small birds are
present, it is questionable how effective iguanas would be in an arboreal
pursuit. Although certainly effective for feeding on leaves and fruit,
their locomotion through bushes is generally slow and rather halting.
It is surprising that Cyclura does not feed on the smaller lizard
species, especially since infrequent cannabalism has been documented
(Iverson 1979). Clearly, the largest source of easily obtainable food
consists of the leaves and fruits lying on the ground. Apparently, this
source is insufficient or lacking in some way, as iguanas frequently
secure arboreally collected fruits and leaves as well. It should be noted
here that all of the largest agamids, iguanids, and scincids are
herbivorous. The argument given above is essentially the same as that
of Pough (1973) to explain the general phenomenon. Large teiids and

290
varanids may all be carnivorous due to the greater ability of these
family members to sustain length]/ strenuous activity without fatigue.
A semiarboreal herbivorous habit in a xeric habitat necessitates
a high regulating phase T^. Climbing over the tops of bushes during
feeding probably results in occasional Tmaxima higher than would be
encountered during exclusive ground feeding. Although large herbivores
have smaller home ranges than similarly sized carnivores, a rule that
probably applies to lizards as well as mammals (McNab 1963, Harestad
and Bunnell 1979) and birds (Schoener 1968) , Cyoluva still usually covers
considerable ground during feeding. This inevitably requires crossing
open areas and stopping in high temperature shade, which in turn leads
to an elevated TAlso, Cyclura usually tolerates high midday tempera¬
tures rather than cooling in its burrow.
Why is Cyolura stenothermic? First, since the maximum upper
limit is less subject to change than the lower limit of the regulating
phase, a high T, lizard is forced toward stenothermia. Secondly,
thermoregulation is energetically cheap in the "low cost habitats"
(Huey and Slatkin 1976) occupied, fairly open vegetatively and commonly
reaching high environmental temperatures. Iguanas can maintain high
Tb's during mid and late summer afternoons without shuttling at all.
Thirdly, food is abundantly available in sunny as well as deeply shaded
areas. Lizards do not have to forage exclusively in the shade, late
in the afternoon, or, as the ant eating Phvynosoma platyrhinos (Pianka
and Parker 1975), after sunset. Since less time is devoted to feeding,
more time is available for thermoregulation. Fourthly, Cyclura is
highly territorial. Males must be able to chase other males away from
their territories and females must be able to chase intruding males and

291
females away from their nests during the entire activity period. Keep¬
ing T^ in a narrow range as much as possible during that time is
adaptive, since the lizard is physiologically more efficient over this
narrow Tâ– , range. Since enzymes function optimally within a narrow T^
range, it is energetically less expensive to function optimally only in
this range, since operating optimally over a wide Th range would require a
battery of different isozymes, many of which would be inactive at any one
time (Heinrich 1977).
Cycluva carinata may have some capacity for invading more densely
vegetated areas, where it would be forced to maintain lower activity
T-< ' s or become a thermoconformer. In the Gainesville pen, when the
diurnal shade time increased to the point of encroaching on the minimum
activity time, lizards emerged in complete shade; that is, they became
more eurythermal. However, iguanas were not abundant in the Mixed
Woodland habitat, an area of considerable shade. This may have been due
to a lack of food or difficulty in digging burrows, rather than the high
preferred Tu of the lizard. Regal (1968) maintained Klaubevina riversiana
for over a year at Th's well below preferred with no deleterious effects
on feeding or health.
Although Cycluva is stenothermic, it still functions adequately
over the full range of s normally experienced. Figure 73 illustrates
most of the body temperature means and ranges measured for behaviors and
physiological parameters during this study. The mínimums of the T,
ranges for behaviors were more variable than the máximums (SD of X for
minimums = 2.94, SD for máximums = 1.57, N = 15). This has also been
noted by Hardy (1962) for Cnemidophovus sexlineatus and is probably common
in high T>:, diurnal heliotherms. Practically the widest range was

Figure 73. Summary of body temperatures of selected behaviors and
physiological maxima of Cyoluva oarinata. Numbers on the
abscissa refer to the following 's: 1) lethal maximum,
2) critical thermal maximum, 3) peak maximum oxygen con¬
sumption after strenuous exercise, 4) maximum voluntary
tolerance (Feb-Oct), 5) panting threshold, 6) peak resting
oxygen consumption, 7) late afternoon burrow submergence
Tfo in the field (Feb-Oct), 8) lizard escape attempts from
June field cages, 9) preferred Tfo in the thermal gradient
(summer and winter samples) , 10) Tj~, during forays (Feb-Oct),
11) clearing of nasal salt gland secretion from the nasal
passages, 12) drinking (June caged field lizards), 13)
defecation, 14) ground feeding (Feb, Oct), 15) arboreal
feeding (Feb, Oct), 16) burrow excavation (Feb, Oct), 17)
start of the first foray (Feb, Oct), 18) maximum mean
recovery rate ([CCO2/(g-h)]/min) during the first five
minutes after strenuous exercise, 19) maximum aerobic scope,
20) mean Tfo of lizards in the regulating phase (Feb-Oct),
21) maximum heart rate increment, 22) minimum daily Tjj of
the regulating phase (Feb-Oct), 23) at emergence in the
field (Feb-Oct), 24) minimum observed feeding Tfo in the
Gainesville pen, 27) critical thermal minimum, 28) loss of
eye opening ability, 29) approximate lethal minimum. Hori¬
zontal lines are means, vertical lines are ranges, and arrows
indicate probable extension of the T^ range beyond the
observed value.

293

294
28.3 C for basking, extending from the minimum T^ for emergence, 15.0 C,
to the maximum late afternoon 2^, 43.8 C. Thus, Cyolura sometimes basked
at T,'s as high or higher than the ranges of many other activities, such
as: drinking, defecating, foray walking, feeding on the ground and in
trees, excavating burrows, and so on. This phenomenon has also been
noted for Scelopcrus oyanogenys (Greenberg 1976), presumably in individuals
which were not hyperthermic during testing. Some behaviors had T-,
frequency distributions which were skewed to the left: the panting
threshold, escape attempts from the June cages, drinking, defecation,
arboreal feeding, and emergence in the field. Skew to the right was
uncommon. The likelihood of submergence into the burrow when scared,
oviposition, and the minimum daily regulating phase Twere examples.
It is adaptive for Cyolura carinata to maximize food intake in the
preferred T^ range. Firstly, lizards were probably better equipped
sensorily to find food. Campbell (1969) and Werner (1972) demonstrated
fairly conclusively that hearing improved in the preferred T\ ranges of
many lizard species. Cnemidophorus sexlineatus was more likely to lose
a pursued insect that stopped moving when at a T\ of 32 C than at 40 C,
presumably due to poorer sensory perception at the lower temperature
(Hardy 1962). Iverson (1977) and I repeatedly observed iguanas running
toward a thrown object, in an "investigative charge," a response which
could be illicited by sound alone. Sites of abundant fruit drop were
probably commonly found by a combination of vision, audition, and olfaction.
Secondly, lizards may have remembered the location of rich food sources
better when they were found while the animals were in the preferred T^
range. Lizards commonly returned to these food sources for several days
Dipsosaurus dorsalis could learn a Y maze at 2\ 's of 27 and
in a row.

295
32 C, but not at 22 C (Krekorian et at. 1968). Unfortunately, due to
using a heat source as a reinforcement, no testing was done in the
lizard's preferred range. Thirdly, reptiles commonly are more efficient
locomotorily when in their preferred T^ ranges. Cruising rate was
independent of Tin the preferred range in Thanrnophis sirtatis (Heckrotte
1967) and Cyelura. The same was true for gross activity in Amphibolurus
barbatus (Lee and Badham 1963). Arboreal feeding was undoubtedly more
energetically demanding than ground feeding, which may be the reason
for the significantly higher Tj_ mean for the former in Cyelura. It may
be especially important for large iguanas to be well heated. Taylor
et at. (1972) calculated that a mouse could run vertically with a much
smaller increase in oxygen consumption over level running (23.5 percent
increase) than could a chimpanzee (189 percent increase) or a horse (630
percent increase). Fourthly, both digestive efficiency (Harlow et at.
1976) and peristalsis (Mackay 1968) are greater in the preferred
range, at least in Dipsosaurus dorsalis and Ctenosaura pectinate,
respectively.
Cyelura did feed at 's below the regulating phase range. The
amount consumed during this anticipatory feeding was usually less than
at higher 2^'s. It was clearly adaptive for a lizard to sample food near
its burrow before or during morning basking or extended overcast while on
a foray, before other foraging lizards found the source. Had a deterior¬
ation of the weather prevented the lizard from reaching or regaining the
preferred Tu range after anticipatory feeding, the likelihood of an
alimentary blockage was minimal since lizards could defecate at T-n's
below the lowest field T^ recorded for ground feeding. As expected,
Cyelura did not feed at 2^’s below the lowest T^ recorded for emerging

296
lizards in the field. The same was true for AnoZis tvopddoZepis (Fitch
1972) and Sceloporus oyanogenys (Greenberg 1976). Feeding T^ can be
depressed in at least one captive lizard; AmphiboZurus fordi fed at 12 C
below minimum field T^ after several weeks in the laboratory (Cogger
1974).
The Tat the initiation of foraging was depressed significantly
during cooler weather. This may have been due to greater hunger during
this time, rather than to lower environmental temperatures. Hunger
depressed the T^ for emergence and the start of activity in Cnemddophorus
sexldneatus (Hardy 1962). Although CyaZura fed all year, it shifted
from a diet of flowers and fruits during warmer weather to leaves in
cool, dry months like February (Auffenberg, in prep.). Guts were no
less full than at other times of the year. However, passage times were
not determined. Also digestive efficiency for leaves may have been
lower than for flowers and fruits.
Comparative Thermoregulatory Behavior of Heliothermic Lizards
Taxonomic classification of lacertids based on differences in
thermally related behavior agrees fairly well with classification based
on morphology. For example, of 25 species of gekkonids examined, only
two were diurnal and two diurno-nocturnal; the rest were strictly
nocturnal (Werner 1969). Obviously, nocturnal activity adds another
dimension to thermally related behavior not found in families of strictly
diurnal lizards. Secondly, a few lizard families (Anelytropsidae,
Anniellidae, Dibamidae, Feylinidae, and Pygopodidae) consist primarily
of fossorial members, which must thermoregulate thigmothermically, if
at all. The majority of lizards show at least some degree of

297
heliothermality (Porter 1972). Thirdly, members of the Lacertidae,
Scincidae, Teiidae, and Varanidae are unable to change their integumental
coloration, whereas most agamids, chamaeleonids and iguanids can (Norris
1967). Fourthly, panting, guiar flutter, guiar pumping, and gaping
behaviors appear to be restricted to specific families or groups of
families (Heatwole et aZ. 1973). Agamids, iguanids, and gekkonids pant;
that is, they open their mouths at high 1s and increase their respiratory
rate, resulting in head cooling by evaporation of water. Only gekkonids
engage in high frequency guiar flutter just prior to panting. Varanids
don't pant. Instead, they engage in a high amplitude, low frequency
guiar pumping (Johnson 1972). Panting is also poorly developed or
absent in the skinks. Egemia hosmeri gapes, opening its mouth without
increasing respiratory or heart rate (Webb et at. 1972). TiZiqua
rugosa and T. scincoides "gulp," opening their mouths only during
inhalation (Heatwole et aZ. 1973). Eumeces obsoZetus and Mabuya striata
do not open their mouths at all at high 1s (Richards 1970).
It would be unwise to define species and genus level phylogenetic
homologies based on thermally related behavior. Normally, as the
similarity of an innate behavioral pattern in a group of species increases,
the probability of common origin also increases. However, since the
laws of heat transfer are constants, similar thermally related behaviors
can just as easily evolve independently in distantly related species,
when they are of similar size and shape and occupy similar habitat types.
Many differences between thermally related behaviors in lizard
species are due to differences in adult body size. Firstly, large size
intrinsically limits what the lizard can do. Dipsosaurus dorsatis
elevated its an average of 7.9 C while in its burrow prior to morning

293
emergence (McGinnis and Dickson 1967). The time of emergence of
Podarais muralis correlated directly with the timing of sun warming of
the lizard's retreat (Avery 1978). These two iguanids may or may not
bask covertly in the same manner as the gekkonid Coteonyx brevis, which
assumes an elevated posture in its rock shelter and makes contact with
the sun-heated roof (Dial 1978). Larger lizards such as Cyctura
cccrinata were not found to bask covertly in sand burrows, probably due
to the thickness of the burrow roof near the mouth and to the common
placement of the burrow mouth under vegetative cover. It would be a
mistake to assume that larger lizards never bask covertly, however.
Sauromatus obesus didn't emerge until the late afternoon during hot days
on the Mojave desert. Individuals had reached 's of 43 to 46 C inside
their rock crevice retreats and were emerging to cool rather than to
bask (Case 1976)! Secondly, lizards which spend part of their lives
buried in direct contact with sand; for example, members of the genera
Uma and Phrynosoma, are all small. Obviously, large lizards would find
it more difficult to bury themselves. Instead, they retreat to predug
burrows, natural rock crevices, or hollow trees. Thirdly, arboreality
is more common among small lizard species. Not only is climbing relatively
more energetically expensive than level walking as body weight increases
(Taylor et at. 1972), but also the variety of vegetation capable of
supporting the lizard's weight decreases. Although Cyctura aarinata is
clearly a capable bush and tree climber, the larger Cyclura stegnegeri
(Wiewandt 1977) and Conotophus suboristatus (Werner, D., personal communi¬
cation) rarely climb in vegetation as adults. Juvenile C. stegnegeri are
frequent climbers. The large Cyclura pinguis is only an "occasional"
climber in low bushes (Carey 1975).

299
As the adult body weight of a lizard species increases, the surface
to volume ratio decreases. This change imposes at least three changes
in the heating and cooling characteristics of the lizard's body, which
in turn alters the thermoregulatory behavior.
Firstly, the rates of heating and cooling decrease and the thermal
lag time increases as body weight increases. For example, an Uta
stansburiana hatchling (0.3 g) in its dark phase heated at a rate of
13.2 C/min in direct midday sun, while a 50.8 g Dipsosaurus dorsalis
in its dark phase heated 7.3 times more slowly (Norris 1967) and a 645 g
Cyclura carinata heated maximally 37.6 times more slowly. The expected
longer morning basking time in larger lizards doesn't necessarily occur,
however. Adult Iguana iguana omitted the morning basking period in
August, exposing only parts of their bodies to the sun and then only for
short durations (McGinnis and Brown 1966) . Thermal comfort postures are
defined as those associated with high and probably function to relieve
thermal stress on all or part of the lizard's body (Greenberg 1976).
Elevation of the tail, toes, legs, and trunk off of hot substrates as
well as panting and retreat to the shade are included. These behaviors
appear to be more commonly observed in small lizards, again probably due
to their higher heating rates. For example, Sceloporus cyanogenys may
lift all four legs and rest with only its ventrum on the hot substrate
(Greenberg 1977). Cyclura carinata, on the other hand, may briefly
elevate its digits off of hot substrates, but usually walks into the
shade instead. Four of five species of Cnemidophorus studied by Schall
(1977) consistently raised their digits during basking. Cyclura carinata
did not. Panting is an infrequent behavior in heliotherms in general,
but is probably more infrequent in large lizards. It usually occurs only

300
when the animal is forced to stay in the open; for example, during day
long, male-male fights in Conolophus aristatus (Werner, D., personal
communication), nesting in female Cyclura stejnegeri (Wiewandt 1977), and
scaring into the open by man in Iguana 'iguana (McGinnis and Brown 1966).
The length of the daily activity period may increase with body weight.
When the frequency distributions of the time of collection of thirteen
Ctenotus species (Fianka 1969) were arranged according to increasing
snout-vent length, the midday active species were small, the bimodaliy
active species were larger, and the all day active species were largest.
Lastly, the increasing thermal lag time as body weight increases may
result in a larger midday depression of the T-h's for seeking shade and
sun. This hypothesis has not been tested as yet.
Secondly, the effectiveness of forced convective heating and cooling
decreases as body size increases (Muth 1977b, Parry 1951, Porter and
Gates 1969, Spotila et at. 1972). During early morning basking several
small lizards have been reported to assume an elevated posture (Greenberg
1977, Hardy 1962, Heatwole 1970, Muth 1977b). Net only does this posture
reduce heat loss to cool substrates, it also increases convective heating
by warmer air further off the ground. No large lizard has been reported
to bask in the morning in a completely elevated posture. Muth (1977a)
recognized three thermoregulatory postures in Callisaurus draacnoides;
prostrate, tail down, and elevated. In field caught lizards these
postures corresponded to mean T^'s of 33.9, 40.5, and 42.7 C, respectively.
A maximally elevated posture is especially effective in convectively
cooling a small lizard, exposing the trunk to higher wind velocity and
lower air temperature. Convective heat loss more than balances the
increased radiant heat gain in this posture (Muth 1977b). Again, the

301
maximally elevated posture is uncommonly seen in large immobile lizards
with high Arboreal behavior should also be more effective as a
morning heating technique (Bradshaw and Main 1967, Cogger 1974, Heatwole
1970) and midday cooling technique (Axtel 1960, Bradshaw and Main 1967,
Judd 1975, Norris 1953) in small lizards. Since arboreality is better
correlated with high surface temperatures in small lizards, the behavior
is more clearly restricted to narrow time bands during the summer day.
Lastly, high winds should be more frequently a cause of premature sub¬
mergence in small lizards (Bradshaw and Main 1967, Bostic 1966, Mayhew
1963).
Thirdly and lastly, large lizards are more tightly coupled to the
solar and thermal radiant environment than small lizards (Norris 1967,
Porter and Gates 1969, Spotila et al. 1972). Large lizards can sustain
a larger change in air temperature and a smaller change in absorbed
radiation while maintaining a constant body temperature. Thus, large
lizards reach higher equilibrium temperatures in the sun (Parry 1951,
Porter and Gates 1969), partially explaining why most of the reptiles
with high late afternoon maximum T^'s are large. Of course, the greater
late afternoon, high tolerance than earlier in the day contributes to
the response. The stronger radiant coupling in large lizards also results
in a more sustained utilization of sun orientation throughout the day;
that is, less random orientation during the activity period than in
small lizards. Another result of high radiant coupling and low con¬
vective coupling may be Norris's finding (1967) that large lizards,
defined as having a surface to mass ratio greater than 3.5 and a body
weight greater than 40 g, are more reflective in both the ultraviolet
and infrared radiation ranges than small lizards.

302
The environmental temperatures faced by a lizard, controlled by
habitat variations in vegetational cover, elevation, and latitude, have
a considerable influence over interspecies differences in thermoregulatory
behavior. The number of times a lizard basks in the sun each hour, called
the basking frequency, as well as the mean length of basking and shade
sitting time periods, show distinct species specific differences corre¬
lated with the openness of the species' habitat (Spellerberg 1972d).
Sphenomorphus quoyi had a high shuttling frequency and low mean basking
and non-basking intervals. The species lives in open rocky areas on
river banks, where exposed areas become quite hot and rocky crevice
retreats remain quite cool. A high basking frequency permitted main-
tainance of a stable regulating phase T^ in a habitat where high heating
and cooling rates could not be avoided. Sphenomorphus tympanum (warm
temperature) had about the same mean field 2^, yet had a longer mean non¬
basking interval, since shade temperatures in its open forest habitat
were fairly high.
Integumental emissivity may vary in a consistent way with the ease
of heating possible in a lizard's habitat. A low long wave emissivity
could lead to overheating in a hot environment. Thus, a 0.965 value for
Sceloporus oecidentalis may be adaptive (Bartlett and Gates 1967). The
0.69 value for Liolaemus multiformis may also be adaptive, since it
permits greater heat retention in a cold environment, the mountains of
Peru (Fearson 1977).
The tropics are characterized by greater thermal and photoperiod
constancy than temperate regions. One would expect this to result in
some basic differences in the thermal biology of animals living in the
two areas. For example, acclimation ability should not be selectively

303
advantageous in the tropics. The CTMax of temperate zone fiddler crabs
could be markedly reduced by lowering the acclimation temperature,
whereas tropical forms did not respond (Vernberg and Tashian 1959). Two
temperate zone salamanders had decreasing standard rates of metabolism
with increasing acclimation temperature while two tropical zone species
were unaffected (Feder 1978). Feder (1976) also found that tropical
salamanders consumed less oxygen while at rest than temperate zone
species of similar body weight. In contrast to fiddler crabs, tropical
amphibians did not maintain unaltered CTMax's when acclimation temperature
was changed. Although the CTM increased as the species latitude of
distribution decreased, acclimation ability did not decrease (Brattstrom
1968). Comparative studies of acclimation ability in temperate vs.
tropical lizards are rare. However, Clark and Knoll (1974) found a
direct correlation between mean and latitude in 28 Anolis species.
The majority of the correlation could be accounted for, through multiple
regression, by the lower light level in the tropical habitats. However,
a significant portion of the correlation was dependent on latitude alone.
The authors hypothesized that raising the MBT at higher latitudes was
adaptive, permitting growth, reproduction, and other activities to
proceed at faster rates in the shorter time frame.
What are some of the interspecies differences in thermoregulatory
behavior of heliothermic lizards which cannot be explained by differing
body sizes, environmental temperatures, and habitats? First, lizards
orient to the sun in two different ways during morning basking. One
group, consisting of several Amphibclupus species (Bradshaw and Main
1967, Cogger 1974, Heatwole 1970), Holbrookia propinqua (Judd 1975),
and Scelopovus cyanogenys (Greenberg 1976), bask in a positive parallel

304
orientation. The other group, including Amphibolurus barbatus (Lee
and Badham 1963), Conolophus subcristatus (Werner, D., personal
communication), Cyclura carinata, and C. stejnegeri (Wievandt 1977),
Leiolopisma. fuscurn and L. vhomboiddlis (Wilhoft 1961), and several
species of Phrynosoma (Heath 1965), bask positive normal as well as
positive parallel. The seasonal orientation shifts in Cyclura carínate
and Phrynosoma are approximately the reverse of one another; that is,
Cyclura basks positive parallel more frequently during low winter
temperatures whereas Phrynosoma only basks positive parallel above
air and substrate temperatures of 20 C. Amphibolurus barbatus orients
positive normal on the ground and positive parallel on logs. When
observed over a wide range of environmental temperatures and under
varied habitat topographies, other heliothermic lizards may also
demonstrate this orientation flexibility. Presumably, the orientation
assumed usually results in maximization of the heating rate.
Secondly, most heliothermic lizards, if they rest on the ground
at all, are variably prostrate during morning heating (Bartholomew
1966, Bostic 1966, Bradshaw and Main 1967, Carey 1975, Cogger 1964,
Hardy 1962, Lee and Badham 1963, Judd 1975, Muth 1977a, Norris 1953,
Werner, personal communication, Wiewandt 1977). Knowing that the
degree of substrate contact varies with disturbance due to the observer,
substrate temperature, and wind velocity, and without interspecies com¬
parisons under identical field conditions, it is unwise to hypothesize
a genetic basis for the minor species differences evident from a search
of the above literature.
Thirdly, several thermal comfort postures are not universally
present in heliothermic lizards. Toe reflexion occurs in at least seven

305
species of Cnemidophorus (Bostic 1966, Kennedy 1968, Schall 1977). It
is present in Cyclura carinata, Dipsosaurus dorsalis (Norris 1953), and
Sceloporus cyanogenys (Greenberg 1976), but is absent in Amphibolurns
inermis (Heatwole 1970). Shovelling behavior occurs in at least
Dipsosaurus dorsalis (Norris 1953) and lima sp. (Stebbins 1943) and very
rarely in Cyclura carinata, but is absent from all Amphibolurus species
studied by Bradshaw and Main (1967).
Complexity of Social Behavior in Cyclura carinata as a Function
of Latitude
Avery (1976) believes that the complexity of diurnal lizard social
behavior or, at least, the potential for increased social complexity,
increases as the species' latitude of distribution decreases. Lacerta
vivípara ranges further north than any other lizard, yet retains a preferred
Tb nearly as great as its southern relatives. Thus, the activity period
is shorter, requiring an increased percentage of active time devoted to
thermoregulation and foraging at the expense of social interaction. This
may have mandated the evolution of its simple social system, characterized
by a lack of territoriality and little interaction between individuals.
As hypothesized, lacertid territorial systems and territorial and court¬
ship display signals are more elaborate in more southerly lizard species.
Cyclura carinata's social behavior is moderately complex, as
expected by Avery's hypothesis. Dominant males are territorial year
round, and territories are fairly rigid, at least where food is abundant.
Local males form a dominance hierarchy based primarily on body size.
Females, however, are only territorial during the nesting season. Specific
male-male and male-female interactions are long term in this long-lived

306
lizard. Yet, the social repertoire appears no more complex than in
other tropical iguanids. Cyclura carinata spends an average of 26.5
percent of the total yearly time above ground. This fits in nicely
between the values for Podarais muralis (19 percent at 44° N) and Agama
agccma (31 percent at 11° N) (Avery 1976, Figure 6, p. 254). Cyclura's
social system, more complex than Podrereis's and less complex than Agama's
agrees with expectation.
Nocturnal Lizard Aggregations
The cooling rate of a reptile decreases as intrasurface contact
increases and intersurface contact with cooler air and substrate decreases.
For example, the carpet snake, Morelia spilotes, reduces heat loss gained
during basking by tightly coiling its body (Cogger and Holmes 1960) and
brooding Python molurus bivittatus reduces metabolically produced heat
loss by coiling around its eggs (Hutchison et al. 1966, Van Mierop and
Barnard 1976). Survival of snakes in temperate region hibernacula is
probably enhanced by the thermal protection of the site, rather than by
any thermal benefit derived from direct contact of individuals (Aleksiuk
1976). Tills does not always appear to be true in the tropics; for
example, Boa constrictor daily basks outside its hibernaculum (Myres and
Eels 1968). Sauromalus varius and S. hispidus aggregate in single
species, mixed sex groups of up to seven individuals (Case 1978).
Digestive efficiency during the initial nighttime hours must be greater
in these groups than in lizards which retreat singly into burrows and
rock crevices. Amphyrhynchus cristatus, living on exposed lava flows
in the Galapagos, forms what must be purely thermoregulatory nocturnal
aggregations, consisting of piles of up to 50 individuals, eight animals

307
in depth (White 1973). The large size of these unique aggregations is
undoubtedly related to the high localized diurnal density in this species
coupled with the small size of the breeding territories. Nighttime
aggregations of more than two Cycluva cccv'inata were not observed in the
field. The group always consisted of a male and a female. The same
pattern has been reported for ;5a.uroma.lus obesus, another highly
territorial iguanine (Case 1978). Long term burrow pairings in Cycluva
cavinata, lasting at least 17 days, were only observed during my
February trip, but may extend into May. Male interest in females during
the day increased in late March through April, culminating in mating in
late April and May (Iverson 1979). Thus, temporary burrow sharing may
serve a reproductive function, such as enhancement of egg development
or female receptivity, rather than a thermoregulatory function. Burrow
sharing may have been more frequent in areas where preferred retreats
were more limited and sufficient food was available to support a high
lizard density. All seven adults in the Gainesville pen squeezed into
a short burrow recently dug by an adult female, rather than using the
previously occupied plywood boxes. Short term pairings, as when two
individuals retreated into the nearest burrow at the start of a heavy
rain or upon my approach, were observed in all seasons.
Physiological Considerations
Aerobic vs. Anaerobic Metabolism
In many lizards more energy is derived anaerobically than aero¬
bically during maximum exercise. Between 60 and 80 percent of the
energy utilized during a 2 minute activity period by Dipsosauvus

308
dorsalis between 's of 25 and 45 C was anaerobically produced (Bennett
and Dawson 1972). This range is conservative since some aerobiosis is
applied to the oxygen debt. Similar results were obtained for Soelopovus
oooidentaiis (Bennett and Gleeson 1976). Whole body lactate was not
determined for Cyclura during exercise and, thus, an estimate of
anaerobic energy production could not be made. Nonetheless, the large
recovery oxygen volume does indicate a heavy reliance on anaerobiosis.
Assuming that the rate of oxygen consumption measured immediately at the
end of the exercise period was a good estimate of the consumption rate
during exercise, recovery oxygen was equal to 70, 71, 71, 73, 76, and
81 percent (2^ = 15, 20, 25, 31, 37, and 42 C, respectively) of the total
oxygen consumption above the resting level during and after exercise.
The anaerobic contribution to energy production in lizards varies
with the length of the exercise period (Bennett and Licht 1972). More
than 50 percent of the total lactate produced during exercise to exhaustion
occurred during the first 30 seconds in several species. This initial
maximum lactate production rate corresponded to the highest activity rate.
An initial burst of activity is typical of iguanids and probably corre¬
sponds to high speed territorial chases and escapes in free-ranging
animals.
The aerobic contribution to activity should be greatest in the
preferred T^ range, where oxygen scope is often greatest (Bennett and
Dawson 1976, Dawson 1975). For example, anaerobic energy production
was approximately 60 percent of the total at 20 C, but only 36 percent
at 37 C over 5 minutes of activity in Uta stansburiana and Cnemidophorus
tigvis (Bennett and Licht 1972). The tapering off of maximum oxygen
consumption at high T^ in Cyolura was probably due more to a maximization

309
of oxygen affinity and capacity of the blood due to high temperature
than to an increasing effect of decreasing pH on oxygen transport, due
to an increasing blood lactate level (Bennett 1973). Both resting and
maximum activity levels of whole body lactate were essentially independent
of over a wide range in all lizards examined except in Dipsosaurus
(Bennett and Licht 1972) and Sceloporus occidentalis (Bennett and Gleeson
1976), with peak maximum production at 's of 40 and 30 C, respectively.
Clear evidence of a high correlation between exhaustibility and
life style has been found for a number of amphibians and reptiles.
Scaphiopus hammondii and Bufo cognatus both failed to fatigue after long
exercise periods, whereas Rana pipiens and Rana catesbiana fatigued
rapidly (Seymour 1973). The maximum oxygen scopes for these anurans
3
were, respectively, 1.22, 1.36, 0.32, and 0.16 cm /(g*hr). The latter
two species rely on a quick jump to water to escape predation. The high
scopes of the former two may be related to their greater digging pro¬
clivities. Bennett and Licht (1974) found similar results for Rana
pipiens and Bufo bóreas. The former had a high anaerobic capacity which
it used for rapid flight from predators. The latter had a low anaerobic
capacity and did not exhaust during its slow movements, relying on
noxious skin secretions and bloating to foil predators. Thamnophis
sirtalis adults took as long as 20 to 27 minutes to reach exhaustion,
presumably because they must subdue slowly exhausting mammalian prey
(Pough 1977). Coluber constrictor and Masticophis flagellum both were
still active after 5 minutes of exercise, whereas Crotalus viridis
was completely exhausted. Lichanura roseofusca intermittently coiled
into a defensive ball after only 1-2 minutes (Ruben 1976). These
differences were clearly reflected in both higher aerobic and higher

310
anaerobic capacities for the more active snakes. Varanus gouldii was
found to be much less susceptible to fatigue than Sauromalus hispidus
(Bennett 1972). The former escaped exhaustion through a combination of
better blood buffering, more efficient lung structure, and higher levels
of muscle myoglobin than the latter (Bennett 1973).
All behavioral motions of Cyclura were of relatively short duration,
alternating with, periods of quiescence. In the Gainesville pen 77.1
percent of lizard movements during the time-motion study lasted less
than 6 seconds. Motions lasting 30 to 60 seconds constituted only
0.7 percent of the total.
Walking, the most commonly observed behavior, was probably a
primarily aerobic activity. Aggressive tracking of habituated adults,
by repeated approaches to distances resulting in lizard withdrawals,
could be continued practically indefinitely. Lizards usually moved
only a few meters before stopping. Amblyrhynchus cristatus adults
swam at an average speed of 27.2 m/min, the "cruise velocity," without
greatly raising blood lactate level (Bartholomew et at. 1976). Lizards
swimming at "burst velocity" averaging 44.6 m/min or 1.64 times as fast,
did exhaust. These two swimming speeds are probably analogous to
walking and fast running speeds in Cyclura.
More strenuous forms of behavior in Cyclura carinaba were either
of short duration and usually not immediately repeated (chases) or
discontinuous, with pauses (climbing, digging, and fighting). Territorial
chases never lasted more than a few seconds. Iverson (1979) found that
only 7.5 percent of male-male encounters resulted in fights and these
were also of short duration (N = 4, 30-70 sec). Sand kicking from
burrows was a decidedly intermittent behavior, similar to that described

311
for Soaphiopus hamnondii (Seymour 1973). These behaviors may well have
resulted in substantial rises in lactate production. Yet, since Cyolura
was fully capable of escape behavior just after completing any of these
more strenuous activities, some anaerobic capacity must always be held
in reserve. Any build-up of lactate could be at least partially removed
during the pauses, especially if the iguana had reached preferred Tu.
Bennett and Dawson (1972) and Bartholomew et at. (1976) have suggested
that lizards are aware of their reserve capacity, through neural sensory
inputs involving blood pH, [H+]/[OH ] or Pq9 and reduce activity to an
aerobically sustainable level long before an incapacitating debt is
incurred. Nonetheless, a daily buildup of lactate may result in
decreased or abbreviated activity. Protracted copulations, fighting,
chases, and nesting in field cages or the Gainesville pen did result in
decreased activity for the rest of the day. The lowest daily lactate
concentration probably occurred just before morning preemergence burrow
movement. The timing of maximum activity in midmorning to midday may
in part be due to this low lactate level. Liver removal of high blood
lactate levels commonly takes a long time in reptiles. Blood and liver
lactate in Natrix rhombdfera required 7 hours to return to resting
levels after only 10 minutes of maximum activity at 35 C (Gratz and
Hutchison 1977). Five hours after a 1 hour dive, Chelonia mydas had
removed only one-half the lactate produced (Berkson 1966).
Why do lizards commonly rely on anaerobic energy production, when
the incapacitation resulting from a high lactate level obviously increases
the animal's vulnerability? Bennett and Licht (1972) argue on energetic
grounds. Lizards operate over a wide range of body temperatures. To
provide a rapid responsiveness aerobically over the full T^ range would

312
require a high basal rate of metabolism. Anaerobic readiness, on the
other hand, does not require a high resting metabolic rate, yet is avail¬
able at all times. Its effectiveness is also essentially temperature
independent.
Cyclura oarinata fled from predators rather than relying on a
static defense, as do some amphibians and reptiles with lower rates of
maximum oxygen consumption (Bennett and Licht 1974, Ruben 1976). The
animal was familiar with the location of burrows outside its usual home
range, to which it could escape when caught far from one of its home
burrows (Iverson 1977). Thus, the location of infrequently visited
burrows must be remembered for long periods. Nonetheless, dogs and
cats, introduced onto Pine and Water Cays during the present study, ran
down adult iguanas before they reached their burrows. When halted
short of a burrow, partially exhausted iguanas may have resorted co a
defensive bloat or a prostrate posture, as observed in the laboratory.
Reliance on a quick escape provided primarily by anaerobiosis clearly
proved ineffective against large carnivores since the Cycluvo. popu¬
lation plummeted to zero within 2 years after the cat and dog
introduction (Iverson 1979). Direct killing and habitat destruction
by man were also important contributions to the population decline.
Neural Basis for Shuttling Behavior in Lizards
The problem of locating the neurons which control lizard thermo¬
regulatory behavior has been approached in two ways; that is, either by
monitoring various characteristics of neuronal action potentials during
heating and cooling of preparations or by monitoring thermoregulatory
behavior in intact animals during or after various manipulations. Taking

313
the first approach, Cabanac et at. (1967) recorded the firing rates of
neurons in the putative thermoregulatory brain center while heating and
cooling the area with a thermode. Also, recordings of peripheral nerve
modalities during local heating and cooling have indicated the presence
of thermal receptors in the integument: Iggo (two mammals, 1969), Bailey
(lizard, 1969), Kenton et at. (caiman and alligator, 1971), Necker
(caiman, 1974), and Spray (frog, 1974). The simplest approach in the
intact animal studies is to implant one or more thermocouples or
thermistors and observe either shuttling behavior in a thermal gradient
(Berk and Heath 1975a, Barber and Crawford 1977 and 1979) or thermal
operant behavior (Garrick 1979). Technically more sophisticated approaches
involve implanting a thermosensitive region of the brain with a thermode
(Hammel et at. 1967, Myhre and Hammel 1969) or producing a localized
brain lesion (Kluger et at. 1973, Berk and Heath 1975b). Biochemical
approaches have also proven fruitful. Lizards have been injected with
pyrogens (see Kluger 1979 for a review) and putative transmitter sub¬
stances (Bligh et at. 1976).
Hammel et at. (1967) and Cabanac et at. (1967) were the first to
show that the preoptic area of the hypothalamus was involved in the
control of thermoregulation in lizards. Berk and Heath (1975b) sub¬
sequently found evidence for involvement of the medial and periventricular
preoptic areas, nucleus of the anterior hypothalamus, nucleus supra-
chiasmaticus, paraventricular nucleus, medial wall of the telencephalon,
and the ventro-medial telencephalon. The posterior hypothalamus
(Kluger et at. 1973-lizard), the medulla (Lipton 1971-rat), and the
spinal cord (Rautenberg 1969-pigeon, Thauer 1970-rat) may also have
controlling influences on thermoregulatory behavior in lizards. The

314
parietal eye-pineal organ-parapineal organ complex also influences thermo¬
regulatory behavior in lizards (Ralph et at. 1979). Peripheral thermal
receptors in the integument clearly have an effect (Myhre and Hammel
1969, Cabanac and Hammel 1971, Barber and Crawford 1979, present study).
Despite all of this work, however, the identification of neural control
tissues and the pathways between them and the effector tissues remains
far from complete.
Lizard thermoregulatory control has been called "broad band" (Bligh
1966), "coupled on-off" (Heath 1970), "stochastic dual-limit" (Barber and
Crawford 1977), and "duel threshold" (Barber and Crawford 1979). Lizards
have coarse T-h control, which they maintain primarily behaviorally.
However, mammalian fine control mechanisms; that is, nonshivering thermo¬
genesis, piloerection, shivering, insensible perspiration, and sweating,
are lacking in lizards. Only vasomotor control has been demonstrated.
Lizards have two distinct T^ "setpoints" (Heath 1970), a maximum and a
minimum, and a refractory range in between where shuttling behavior, but
not proportional control, is in abeyance. For example, in a two tempera¬
ture, forced convection chamber that was heated or cooled depending on
which end of shuttle board Dtpsosaurus was sitting on, shuttling ceased
when the cold temperature was elevated to 36 C or the hot temperature
was lowered to 43 C, leaving a refractory zone of approximately 7 C
(Barber and Crawford 1979).
The neural basis of lizard thermoregulation was considerably
clarified by Cabanac et at. (1967), who discovered two groups of tempera¬
ture sensitive neurons in the hypothalamus of the skink, Titiqua
sainaoides. One group, called cold-sensitive neurons, increased their
firing rate with decreasing brain temperature. Another group, the

315
warm-sensitive neurons, increased their firing rate with increasing brain
temperature. Most of the neurons recorded fell into a third group, the
temperature insensitive neurons, which did not change their firing rate
with increasing brain temperature. Similar firing rate patterns have
been found in the recordings of mammalian hypothalamic neurons (see
Bligh and Moore 1972 for a review). The problems with Cabanac's study
were the small number of neuronal recordings, the failure to record the
change in firing rate over the full temperature range cf each neuron, and
the failure to consider other neuronal modalities, such as spike amplitude
Also, for some reason, no one has followed up on the study using other
reptiles.
Hamnel (1965), thinking of mammals, was the first to hypothesize
how warm and cold neurons interact to determine the T- setpoints. Heat
producing mechanisms (or shuttling into sunlit areas in the case of
lizards) may be initiated when the firing rates of the cold-sensitive
neurons exceed the rates of the temperature insensitive neurons, which
act as inhibitory references. Heat dissipating mechanisms (or shuttling
into the shade in lizards) may be initiated when the firing rate of the
warm-sensitive neurons exceed those of the insensitive neurons. Since
many neurons in each of the three groups are probably involved, each
with a different Qfor firing rate or a different constant firing
rate level, the setpoints probably result from an integrative averaging
of many different firing rates.
Lizard setpoints are not fixed, but rather shift simultaneously
either up or down in response to numerous stimuli: digestive state
(Regal 1966-snake, Gatten 1974c-turtle), reproductive state (Garrick 1974)
thyroid function (Wilhoft 1966), darkness (Regal 1967, Vernon and Heatwole

316
1970), inter- and intra-individual variation (present study), fever
(Kluger 1979), and parietalectomy (Roth and Ralph 1976 and 1977). Three
hypothalamic neuron changes have been proposed to account for these
changes in setpoint, in accordance with the Hamrnel model. First, the
constant firing rate of the temperature insensitive neurons could increase
or decrease. An increase in firing rate above that of the cold neurons
was proposed by Berk and Heath (1975a) as the mechanism for nightly
voluntary hypothermia in diurnal lizards. However, since this would
also result in the elevation of the upper setpoint, this is an unlikely-
explanation for the simultaneous depressions of upper and lower setpoints.
Secondly, lines representing warm and cold neurons firing rates vs.
ambient temperature could shift simultaneously and in the same direction
along the X-axis without changing slope, that is, sensitivity. Lastly,
the Qjq's °f the temperature sensitive neuron firing rates could change.
This has already been demonstrated for rabbits (Cabanac et at. 1968)
and cats (Eisenman 1969) during pyrogen induced fevers. In both these
cases the slopes of the cold neurons increased and the slopes of the
warm neurons decreased, raising the setpoints. Temperature insensitive
neuron firing rates were generally not effected by pyrogen administration.
Data gathered in the present and other studies point to involvement in
lizards of both X-axis shifts and Qj^ changes, as will be discussed
below.
Thermal gradient studies have revealed that the standard deviation
of the mean minimum T^ is significantly greater than the SD of the mean
maximum T^ for individual lizards of at least two species (Berk and
Heath l9l5a-Divsosaurus dorsalis, Garrick 1919-Soelopovus cyancgenys).
Although not true for Cyclura, the trend is in the same direction and,

317
as shown by the heating and cooling experiments, the SD of mean intestinal
temperature for heat seeking behaviors was significantly greater than for
heat avoidance behaviors (Table 21). As hypothesized by Berk and Heath
(1975a) and Barber and Crawford (1979), the neurological basis for this
consistency in lizards may be the greater firing rate Q^q for warm
neurons than cold neurons found by Cabanac et at. (1967) for Tztiqua.
Mammalian hypothalamic neurons may (Wit and Wang 1968) or may not
(Cabanac et at. 1968, Eisenman 1969) show this characteristic. Qjo's
for peripheral thermosensitive neurons in the cat, monkey, and rat do
(Iggo 1970). A second modality which may partially account for the
observed SD difference in lizards is the range of ambient temperatures
over which the receptors fire. The range was wider for rat scrotal
cold receptors than for warm receptors (Iggo 1969). Unfortunately, the
reported lizard data to date are limited to recordings from only eight
temperature sensitive hypothalamic neurons (Cabanac et at. 1967).
The larger Q~^ for the firing rate of warm neurons than cold neurons
may also be the cause of the negative skew for Dipsosaurus 2C frequency
distributions, first reported by DeWitt (1967) and more recently found
in a number of other poikilotherms by DeWitt and Friedman (1979) as
well as the present study. Inter-species increases in field operant
2^'s (Bogert 1949, Pianka 1969) are coupled to seme extent with higher
negative skew in their T^ frequency distributions. The same may be
true for panting frequency distributions (Heath 1965). If an increase
in mean T^ is the result of a simultaneous increase in the Q^ of the
cold neuron firing rate and a decrease in the Q^(j of the warm neuron
firing rate, theoretically resulting in an increase in the negative
skew of the 2^ frequency distributions for both minimum and maximum

318
Tb's, then the overall T^ frequency distribution should also become
more negatively skewed as mean T-n increases. Indeed, both the skews
for minimum and maximum T^ in Cyolura summer lizards became more negative
as their respective mean temperatures increased (Figure 74). As the
firing rate slope for the cold neurons increased, the standard deviation
of the mean minimum ZV should decrease, as was the case. However, the
standard deviation of mean maximum also decreased, though somewhat
more irregularly, even though it would be expected to increase based on
a decreasing Qjq for the warm neuron firing rate. Perhaps, peripheral
input overrode the hypothalamic warm neuron influence at high T- . On a
lizard by lizard basis, there was no correlation between the standard
deviations of the means and either mean minimum or mean maximum T, . It
b
may be unwise at this time to draw any conclusions concerning the relation¬
ship between temperature sensitive neuron firing rate Qjn and the
variation m mean
T
â– y
For individual Dipsosaurus (Berk and Heath 1975a) and Cyolura, the
SD of the mean for minimum T-^' s was directly and linearly related to the
SD of the mean for maximum T,'s. The distance up the SD vs. SD .
b max mm
line in Cyolura was almost significantly, indirectly correlated with body
weight (N = 12 field lizards, r = -0.414). Thus, increasing thermal lag
with increasing body weight may be the explanation for this SD correlation
rather than having a neurological basis. The smaller linear correlation
for SD vs. SD . for the smaller Dipsosaurus also agrees with this
explanation (r = 0.435 for Dipsosaurus, 0.804 for Cyolura, both in the
thermal gradient). However, the correlation of distance out the line
vs. body weight was not calculated for Dipsosaurus, since body weights
of individual lizards were not reported (range = 27-60 g). The more

0.2
1.4
0.1
0
-0.1
-0.2
ro
o
-0.3
-0.4
-0.5
-0.6
-0.7
*
\
\
\
\
\
\
\
\
\
\
\
\
\
36 37 38 39 40 41 42 36 37 38 39 40 41 42
MEAN OF Tb MINIMA AND MEAN OF Tb MAXIMA
Figure 74. Effect of increasing body temperature on frequency distribution skew and standard deviation
of the mean of Tfo minima and Tj~, maxima. The three samples consisted of the highest, mid-
range, and lowest mean Tfo days of data (maximum of two days per lizard) for eight summer
field lizards. All minimum and maximum 's for each day were used except those resulting
from burrowing and cloud cover. Minimum Tfa's are represented by dashed lines, maximum Tfc's
by solid lines. The asterisks indicate that the skew was significantly negative (p < 0.05
for máximums, p < 0.01 for mínimums)..
319

320
rapid increase of SD . than SD in both Cyolura and Dipsosaurus is
mxn max u r
again suggestive of a higher for the warm neuron firing rate vs.
ambient temperature.
Bligh (1972) has described a neuronal control model for mammalian
thermoregulation which is applicable to lizards as well. The model
consists of two main lines of central nervous system interneurons, one
ultimately synapsing with motor neurons controlling shuttling into the
shade and the other shuttling into the sun. Each line receives inputs
from: 1) either stimulatory hypothalamic warm neurons (to the shade
shuttling line) or cold neurons (to the sun shuttling line), 2) inhibitory
Iwpothalamic temperature insensitive neurons, 3) non-thermal, inhibitory
and stimulatory neurons from higher brain centers, 4) either stimulatory
peripheral warm or cold receptor neurons, and 5) inhibitory neurons from
the other line. One and two have already been discussed. Evidence for
higher brain center inhibitory input in Cycluva is limited to the ob¬
servation that shade seeking behavior can be temporarily overridden by
feeding and social behaviors. Evidence for peripheral warm and cold
receptors in lizards is mostly indirect. For example, high ambient
air temperature in a shuttle box depressed the colonic temperature at
exit (Myhre and Hammel 1969) and low T' elevated it (Cabanac and Hammel
1971) in Tiliqua sc'inco'tdes. Yet, peripheral input may be even more
important than brain inputs in determining the shuttling setpoints, as
indicated in the following three studies. First, direct heating of the
preoptic region of T-it'iqua decreased both the mean maximum skin tempera¬
ture and maximum colonic temperature, while cooling increased the mean
maximum and mean minimum skin temperatures and the colonic temperature
(Myhre and Hammel 1969). Cooling the preoptic region approximately

321
10.7 C below control hypothalamic, shade-seeking temperature raised
skin and colonic temperatures only 1.3 C, on the average (0.13 C increase
per 1 C decrease in hypothalamic temperature). Heating the preoptic
region 4.7 C decreased skin and colonic temperatures by 1.6 C, on the
average (0.34 C decrease per 1 C decrease in hypothalamic temperature).
Thus, considerable change in hypothalamic temperature was required to
change skin and colonic temperatures. Also, skin and colonic maxima
were three times more sensitive to an increase in hypothalamic tempera¬
ture than a decrease. Secondly Bligh et at. (1976) found in Va?anus
atbigutcaris that core temperature at the initiation of "panting" ranged
from 35.4 to 38.6 C, while skin temperature varied hardly at all, in¬
dicating that the latter may be regulated. Thirdly, Barber and
Crawford (1979) found for Dipsosarurus that while colonic temperature at
exit from the hotbox decreased as air temperature increased, maximum
skin temperature remained constant. They hypothesized that skin tempera¬
ture rather than core temperature was being regulated.
Since mean minimum T, increased faster than mean maximum T-, , the
b b’
refractory interval in Cyatura oar-inata decreased as mean T. increased,
both xfithin and between individuals (Figure 22 and Table 10). This was
not just a result of seasonal differences in environmental temperatures,
since summer data spanned the entire range in Figure 22, encompassing
February and October results. Clearly, the relationship has adaptive
significance. A lizard operating at high T\ is more likely to keep
maximum below a dangerous level than maintain a narrow range of
minimum Th's. Nonetheless, the two setpoints are coupled. The
alternative, a narrowly variable maximum range uncorrelated with the
minimum, would be disadvantageous for those individuals in the population

322
with the widest refractory range. In a large lizard like Cyclura
carinata, too much time would be wasted seeking out and sitting in an
appropriately cool spot before activity could begin again. When the
Berk and Heath (1975a) and the Barber and Crawford (1979) data for
Dipsosaurus were graphed in the same way as the field data for Cyclura,
the correlations between mean high and mean low shuttling 2^'s were not
significant (linear correlation coefficients were 0.153 and 0.113,
respectively). The mean maximum did not get above 43-45 C for a wide
range of mean minimum colonic temperatures. This may have been due to
the limitations of laboratory shuttle boxes, which are visually, thermally,
and socially restrictive, as well as to the lizard's smaller size.
Vaughn et al. (1974) did demonstrate that individual Dipsosaurus during
fever have a shrinking refractory interval as mean T~_ increased, with
minimum .7, increasing faster than maximum. Also, mean maximum colonic
and skin 21 were significantly correlated with mean minimum colonic and
D
skin temperatures, respectively, in the larger Tilrqua sc-incoides, with
mean minimum increasing faster than mean máximums (Myhre and Kammel 1969).
Garrick (1979) demonstrated a different pattern for a shrinking
refractory interval in Sceloporus cyanogenys; that is, a more rapid
decrease in mean colonic exit temperature than entrance temperature as
heat lamp intensity increased. The Scelopovus response to high heat
input was probably qualitatively identical to the midday depression in
core T-d maxima and minima for Cyclura in the field. Higher peripheral
input at maximum 21 due to higher peripheral temperature may be integrated
U
with a lowered core input due to a greater surface-core temperature
differential to produce a shade seeking shuttle at a lowered core tempera¬
ture. The depression in minimum 21 after high intensity heating in

323
Cyclura and Scelopovus is harder to explain. In fact, Barber and
Crawford (1979) reported no drop in minimum core or surface Twith
increasing hot chamber temperature in Dzpsosauvus. The best explanation
is that "shaded" air temperature in Barber andCrawford's shuttle box
was kept constant, whereas it was definitely elevated under midday
field conditions and also in the non-circulating air, operant condition¬
ing apparatus used by Garrick. The resultant higher peripheral tempera¬
ture during cooling would not only directly inhibit peripheral cold
receptor firing, but, according to the Bligh model (1972), would also
increase the inhibitory influence of any peripheral warm receptors.
The neurological basis for the narrowing of the refractory interval
as mean T^ increased in Cyolura may be both a rightward unequal shifting
along the X-axis of the firing rate lines for hypothalamic cold and warm
neurons and an unequal change in the slope for the two lines (Figure 75).
As mean increased, the rightward shift in the cold neuron line may
exceed that of the warm neuron line. This alone could account for the
decreasing refractory interval. However, when the a^'s for the frequency
distributions of minimum and maximum T^ ' s for the eight summer field
lizards were plotted against mean minimum and mean maximum T-,, respectively,
the lines were fairly linear, and the slope of the minimum T-^ line was
greater than the maximum (Figure 74). The skew of the distribution of
minimum 's should become more negative as the cold neuron firing rate
slope increases and the skew of the -distribution of máximums should
become more negative as the warm neuron firing rate slope decreases.
These slope changes are also expected based on the firing rate responses
of the hypothalamic warm and cold neurons in mammals during fever (Cabanac

NEURON FIRING RATE
324
Figure 75. Hypothetical neural basis for changes observed during increas¬
ing mean body temperature in Cyclura cari.na.ta. Numbers 1), 2),
and 3) indicate the firing rates of the temperature insensitive,
cold, and warm neurons, respectively. Arrows indicate 4) the
lower and 5) upper setpoints for Tfo during the regulating phase
and 6) the increasing and 7) decreasing firing rate slopes of
the cold and warm neurons, respectively. The figure is
modified from Berk and Heath (1975a).

325
et at. 1968, Eisenman 1969). If the slope changes occur in Cycluva as
T-d increases, they would contribute to the decreasing refractory
interval.

SUMMARY OF RESULTS AND CONCLUSIONS
1. Cyotura, car'inata is a diurnal heliotherm. Its T^ cycle can con¬
veniently be divided into four phases; morning heating, regulating,
evening burrow cooling, and nightly stable Mean field, regulating
phase Tjâ–  ranged from 38.0 to 39.7 C, higher than for many other
iguanids. Although faced with above lethal black body temperatures
during part of the day over much of the year, iguanas were steno-
thermic during the regulating phase, with a mean range of only 3.3 C
for all monitoring days.
2. Iguanas were initially hyperthermic in both the field and the
thermal gradient. The mean time between release and the maximum
hyperthermic response was 2.0 days in the field, with a mean maximum
T^ range of 4.4 C. The hyperthermic responses may have been due to
bacterial infection, transmitter meal, and thermal learning in
coelomically implanted, transmitter force-fed, and thermal gradient
individuals, respectively.
3. The 0.5 C seasonal variation in mean regulating phase at Site II
was close to the annual variation and expectedly low for a tropical
lizard. Mean T^ did not correlated perfectly with seasonal variation
in ambient temperatures; the October mean was 0.4 C higher than the
warmer month of July. Summer mean Th of nonhyperthermic iguanas
was 1.6 C higher in sparsely vegetated Semi-open Scrub than in a
mixture of Dense Scrub and Rocky Coppice. The difference was
significant during hyperthermia as well. Mean T-'s were probably
326

327
even higher in Open Scrub, but iguana density was too low there to
warrant monitoring. Iguanas thermoregulated as well in field cages
with only burrow shade as in cages with burrow shade, vegetational
shade, and elevated perches.
4. Mean preferred did not differ significantly on any day of the first
four days for iguanas acclimatized to summer vs. winter Gainesville
temperatures, more extreme than in the Caicos. Thus, acclimatization
to a temperature range as large as experienced in the field did not
alter the acute (first day) nor final (fourth day) preferendum. However,
winter means were significantly higher during the first two days than
during the second two days in the gradient. The ranges of variation in
both the field and gradient were less in winter, indicating a narrowing
of temperature tolerance. Mean preferred 2^ was lower than mean
operant T^ in both summer and winter, the difference being greatest for
the hottest environment, the least densely vegetated site in June.
5. Mean T^ probably did not differ with sex, with the exception of gravid
females, which, based on one individual, had lower Th's and weaker
hyperthermic responses. On the average, adult females heated and
cooled faster than adult males, simply due to their smaller size.
6. Mean regulating phase Tâ– , decreased as overcast time increased, the
correlation improving 2.8 times as environmental temperatures decreased
from July to February. Continuous overcast had to be lengthy for T^
to fall below the mean minimum. Tâ– , commonly increased initially
during medium to heavy overcast, due to either or both thermal lag
and iguanas seeking out hot substrates and assuming a prostrate
posture when T-^ was rising just prior to overcast and due only to
the latter when T^_ was falling just prior to overcast.

328
7. T£ was nearly independent of shaded air temperature, demonstrating
the iguana's dependence on radiant heat gain. The mean T- -
difference ranged from 5.9 C in July to 11.0 C in February, with a
minimum difference of 1.3 C and a maximum difference of 15.6 C. T^
was usually considerably less than sun exposed sand temperature,
except early and late in the day and during extended overcast, when
T decreased faster than T-, .
s b
Placing transmitter probes dorsal to the back, under the abdomen,
and in the gut of iguanas in the Gainesville pen permitted a crude
analysis of dorsal and ventral heat transfer, dividing the lizard's
day between four heat transfer conditions. Iguanas were thigmothermic
as well as heliothermic during part of the morning heating phase.
During the morning portion of the regulating phase, they alternated
between pure heliothermia (gaining heat dorsally, losing it ventrally)
and losing heat both dorsally and ventrally. Pure thigmothermia
(gaining heat ventrally, losing it dorsally) was rare, constituting
only 2.5 percent of the active time. Thigmothermia, with or without
dorsal heat gain, constituted only 20.7 percent of active time, nearly
half during the short morning heating phase. Iguanas were heliothermic
nearly 70 percent of the time. The number of heat transfer conditions
was directly and linearly related to both mean Tâ– < and to the total
number of iguana movements.
8. Individual mean minimum T^ was highly correlated with mean maximum
T^, as was intraindividual change in minimum daily T^ vs. maximum
daily T-^. The linear regression lines extrapolated to approximately
the maximum voluntary tolerance, with the individual mean minimum T^
and intraindividual change in minimum daily T^ increasing faster than

329
the individual mean maximum T\ and intraindividual change in maximum
daily , respectively. Thus, the regulating phase 2^ interval
decreased as mean increased. As a corollary, the SD of mean Th
also decreased as mean T-^ increased.
A Berk and Heath (1975a) analysis of daily consecutive maximum-minimum
and minimum-maximum pairs in the field and the thermal gradient
revealed a high positive correlation; that is, when the second
maximum was greater or less than the first maximum, the second minimum
was greater or less than the first, respectively. Daily and seasonal
variation in this pattern correlated with daily and seasonal vari¬
ation in ambient temperatures, at least partially explaining the
high positive correlation.
The standard deviations of individual mean minimum ' s were commonly
slightly but not significantly greater than the SD's of maximum T-,'s.
o
Individual SD 's were directly and linearly correlated with their
max
SD . 's, that is, if an individual regulated its maximum T, well, it
mm b
also regulated its minimum Th well. The correlation was in part a
passive result of body weight variation, since the SD's increased non-
significantlv as body weight decreased. SD decreased faster than
SDmin> expected since overheating was a more serious problem than a
wide minimum T^ range.
Mean T~ was not correlated with body weight. Yet, large iguanas
(X weight = 1446 g) were slightly but not significantly better thermo¬
regulators than smaller ones (X weight = 729 g), no matter what their
mean T-. As mean increased, mean SD decreased more rapidly in the
larger individuals, which may be due to their greater danger of
overheating due to thermal lag.

330
Overall frequency distributions of individual minimum and maximum
2^'s were most commonly negatively and positively skewed, respectively,
whereas individual T^ distributions of intermediate 's plus maximum
and minimum 2^'s tended toward negative skewness, both in the field
and the thermal gradient. When individual Tâ– , frequency distributions
were examined on a daily basis vs. mean 2^, negative skew was most
evident in the midrange of mean 2^'s, and a weak, nonsignificant
positive skew occurred at high mean TThe negative skew increased
faster for the minima than for the maxima as their separate mean Th's
increased.
Results from minimum and maximum 2%, analyses were interpreted in
terms of the present limited knowledge of the neurological basis of
lizard thermoregulation.
9. Maximum 2^ most commonly occurred late in the summer day, especially
among large males basking in the few remaining sunlit areas. This
trend continued in February and, to a lesser extent, in October.
Maximum daily Twas also recorded during the final hour in the
winter and summer thermal gradients, shifting to that time after
four days in the gradient. It appears that Cyc Zura' s T^ tolerance
is greatest at the end of the active period. In the field, heat
exchange was usually in a brief equilibrium at this time since
even though exposed substrates were hot, radiant flux was decreasing.
Thus, if body surface temperature was regulated rather than the core,
core T-, would tend to increase during this time. The gradient
results point to a periodicity in temperature tolerance even in the
absence of changing environmental temperature. A literature search
revealed that most if not all lizards with higher afternoon 2^'s are

331
large, which is expected based on their larger surface-core tempera¬
ture difference and tighter coupling to radiant heat exchange at
equilibrium.
Surface rather than core temperature may be regulated in Cyclura,
as indicated by the above circumstantial evidence as well as four
other points: 1. core temperatures were higher in October than July,
2. maximum, minimum, and panting threshold core T-, 's all dropped to
D
a minimum at midday, 3. the maximum T^ recorded during an overcast
period was commonly higher than the preceding maximum, 4. during
radiant heating in the photothigmotron, the T- at escape to the
shade increased as substrate temperature increased.
10. Morning basking was a deliberate attempt to rapidly raise T^ from
near burrow temperature to the of the regulating phase. Almost
all behavior during this phase was thermoregulatory, with minimum
feeding and social behavior. The morning heating rate was, on the
average, 3.3 times greater than the regulating phase heating rate
and 4.0 times greater than the regulating phase cooling rate. Regu¬
lating phase heating, by behavioral, passive, and physiological
means, was 1.15 times greater than cooling. Iguanas also heated
faster than they cooled in a controlled environmental room. All
heating and cooling rates were dependent on body weight, with a
doubling of weight decreasing the morning heating rate by an average
of 54 percent, the regulating phase heating rate by 59 percent, and
the regulating phase cooling rate by 62 percent.
Core nearly equilibrated with burrow temperature after a mean
of only 19 percent of nightly submergence time. Movement was
minimal in the burrow, yet a common decrease in the cooling rate

332
at a mean T, of 34.4 C was probably due to a postural change, an
increased adductive contact of the limbs with the trunk and tail.
Average maximum burrow depth correlated well with the region of
nearly perfect, 24 hour, soil temperature stability. Burrows also
provided an ambient temperature higher than surface temperatures
during the night and during cold fronts. Seasonal variation in
mean, burrow terminus temperature probably did not vary more than
4 C.
The iguana cooling rate in burrows was probably reduced somewhat
during the prebreeding months, when a male and female sometimes
shared the same burrow. Larger nocturnal aggregations did not occur
in the field.
11. Preemergence movement, consisting of a stepwise crawl toward the
burrow mouth, began up to 37 minutes prior to emergence in one
individual. Iguanas usually only emerged when their T^'s would
quickly rise above burrow temperature. Emergence was always delayed
by medium to heavy overcast. However, on rare occasions (12.5
percent of observations) iguanas would emerge and stay out even
though T-d did not rise appreciably above burrow temperature. Summer
free-ranging iguanas, removed from their habitats for up to a day
during transmitter implantation, all demonstrated a phase delay in
emergence time after release, which gradually disappeared over the
next several days. Presumably, their biological clock was re¬
entraining to the natural day length. Emergence was a two step
process; the crawl from deep in the burrow, presumably initiated by
a biological clock, and the actual emergence from near the burrow
mouth after a preemergence pause, probably activated by a range of

333
light intensities reaching the iguana's eyes, an intensity nearly
always corresponding to a high positive net heat transfer.
Once emergent, iguanas usually stayed out the full day. No
adult submerged due to intolerably high surface temperatures, at
least in monitored Semi-open Scrub and more densely vegetated areas.
Submergences due to overcast and rainfall were also uncommon, as
expected in a large, slow cooling ectot’nerm.
Activity time was tightly coupled to photoperiod. The mean
seasonal decrease in activity time , 3 hours and 26 minutes, was
almost identical to the seasonal decrease in photoperiod, 3 hours
and 14 minutes. Activity time was not appreciably reduced
relative to sunrise or sunset by the decreased water availability
in the dry season. Seasonal change in submergence timing was
slightly less well tied to the sunset time than emergence was to
sunrise, since seasonal variation in access to heated sunlit areas
late in the day was greater than access to morning basking platforms
at the primary study site. Several examples of long term, significant
differences in activity times between individuals were linked to
differences in shade timing due to topographical variation between
home ranges. Activity periods of dominant males in the field were
longer than for females and subdominant males, due to their later
submergence times. Activity time in the Gainesville pen was also
directly related to body weight. However, unlike the field, emergence
times were even more tightly coupled to body weight than submergence
times. This difference was probably due to the enormously higher
iguana density in the pen than in the field, and the consequent replace¬
ment of territories with a weight correlated peck order.

334
The range of emergence 1s in the field, 6.1 C, was slightly
less than the submergence range, 8.2 C. 2% independent emergence
was more clearly demonstrated in the Gainesville pen, where iguana's
emerged at Th's as low as 15 C. T^ at submergence was an average of
10.3 C higher than at emergence. No penned individual could be
enticed by an artificially warmed external environment to remain
outside the shelters after light intensity had decreased to a low
level.
12. Activity during the morning heating phase was essentially constant in
rate and, thus, independent of increasing T-, . Iguanas were most
easily scared back into their burrows during this time, the submergence
distance being considerably less than burrow length. An abrupt
increase in activity occurred when the first foray commenced.
Although neither the duration nor the mean heating rate of the morning
heating phase differed from February to October for equal weight
iguanas, the mean f^ at the start of the first foray was significantly
lower in February than October, perhaps an acclimatization to lower
winter temperatures.
The major portion of the regulating phase consisted of a series of
forays through the sun-shade mosaic, usually starting and ending at
the present burrow of residence. Maximum activity occurred during
the first foray. The activity distribution was unimodal during the
cooler months of November and December and became steadily more
bimodal toward midsummer, with the midday lull corresponding to
maximum surface temperatures. Burrow excavation was usually restricted
to the midday hours of 1000 to 1400, with short excavation periods
alternating with short basking periods. High radiant input during

335
the basking periods may be the reason for the midday excavation
timing, since excessive heat loss was prevented and the procedure
could be completed quickly.
Arboreal activity probably served two thermoregulatory functions
as well as the securing of leaves and fruits; the avoidance of high
surface temperatures at midday in midsummer and the seeking out of
warmer basking sites in the winter morning. On the average, rose
during arboreal feeding in October.
The effectiveness of cloud cover in reducing activity was greatest
during midafternoon, with a lesser peak in effectiveness at the
start of the first foray. It was least effective during the hour
of morning maximum activity.
13. The mean distance traveled on clear or nearly clear days did not
differ significantly between first, second, and third forays. The
rate of travel declined on each successive foray, but again less
than significantly. Foray length was not depressed by overcast
time equal to or less than 45 percent of foray time. Iguanas simply
stopped if the overcast period was of long duration, assuming a
prostrate posture until the overcast cleared, then resumed travel.
As a result of the delay, the mean rate of travel during forays was
inversely correlated with overcast time. Forays only occurred over
a 5.5 C mean T^ range, from 35.0 to 40.5 C. Foray length and mean
rate of travel were independent of T^ over this range, except when
overcast time was 30 percent or less. The mean rate of travel was
then inversely related to since iguanas spent more time sitting
in the shade avoiding overheating.

336
Iguanas were more eurythermal during the first than subsequent
forays. during second and subsequent forays usually both increased
and decreased relative to the starting T^, staying near the middle
of the regulating phase range. Iguanas actively maintained upper
and lower thermal buffer zones, entered only during infrequent long
term activity in sun-exposed and deeply shaded areas.
14. The log of mean distance traveled per day was directly and linearly
related to the log of the confinement area. Active time averaged
only 1.8 percent of the emergent time in the summer Gainesville
pen and was estimated to equal 7.5 percent of emergent time in the
field, a reasonable estimate based on tracking observation. In a
limited time-motion study in the pen, thermoregulatory movements
took up an average of 43 percent of total active time. Movement was
defined as body displacement in the horizontal plane and thus did not
include lifting limbs off hot substrates and proportional changes in
substrate contact and trunk symmetry relative to the sun. Individual
thermoregulatory movements took considerably less time than feeding
and social movements. The total number of thermoregulatory movements
per day was probably inversely related to body size.
15. Ground feeding during February through October occurred at a signif¬
icantly lower T^ than arboreal feeding, since iguanas did not climb
bushes before the start of the first foray. Since Iverson (1977)
observed morning basking without- feeding in bushes in winter, the
absence of arboreal feeding below a of 35.7 G was not due to an
inability to climb. Although iguanas fed on the ground at 's as
low as 23.6 C in the Gainesville pen and 30.0 C in the field, both
feeding rates and meal size were lower than at higher T^'s. Probably

337
arboreal feeding at low T^ was rare or absent due to disinclinations
to feed and to stop morning basking, usually performed on the ground.
Defecation was possible over a wide range of 2^'s, ^rom 25.0 to
probably 46.2 C. However, the range for undisturbed iguanas was
smaller, 33.6-40.4 C. Defecation usually occurred once a day during
the first foray.
Water was normally obtained from food. Nonetheless, iguanas could
drink proficiently from free-standing sources. Drinking in field
captives was restricted to high mean T38.2, probably due to a
reluctance to break morning basking.
Expelling nasal salt gland secretion via sneezing occurred at a
fairly uniform rate during the day and was primarily restricted to
the regulating phase (X = 37.7 C) .
16. The thermoregulatory movements and postures of Cyclura aca'inata were
illustrated and described. Major thermoregulatory behaviors were
quantified in relation to changing under different ambient thermal
environments.
The most important behavioral means of T^ control was moving
between sunlit and shaded areas. Shuttling resulted in a rapid
reversal of the direction of 2V change and was most in evidence on
the T£ records at the minimum and maximum Tj_' s of the regulating
phase. Shuttling rate varied significantly between individuals in
field cages and was directly and linearly related to T^ stability.
3asking occurred over the full range of body temperatures, all the
way to the maximum voluntary tolerance, 43.8 C; that is, throughout
the regulating phase. Only once was observed to rise above this
temperature, to 44.0 C in a small male in obvious heat stress unable

338
to reach shade due to a large male blocking its path in the field
cage. Heat gain postures were particularly common early and late
in the day, when solar flux was low. Only 18.9 percent of daily
direct sunlight time in the summer cages was distributed between 1000
and 1500. During maximum midday heat input, iguanas were quite
selective in choosing shade with a low air temperature and a high
wind velocity, a predictive process clearly tied to learning the
thermal habitat, possible in a territorial animal which stayed for
long periods in one area.
17. Iguanas in the photothigmotron maximized radiant heat input below the
minimum voluntary tolerance and either abruptly or gradually decreased
the input as the preferred T-, range was approached. Orientation of
the long axis relative to the rays of the sun was coupled with trunk
tilting. At low substrate temperature below the minimum voluntary
tolerance, iguanas maximized the trunk exposure area on the side
facing the sun by tilting the body away. This response was lost at
higher substrate temperatures. Blindfolding eliminated sun orienta¬
tion behavior, apparently mediated visually, but did not alter
thigmothermic behavior.
Sun orientation changed daily and seasonally. Consistently
predictable patterns of daily orientation were most in evidence at
T\ and ambient environmental temperature extremes: positive parallel
and positive perpendicular orientations during warming toward the
preferred Trange and negative parallel and perpendicular orienta¬
tions during approach toward the maximum voluntary tolerance. The
main heat gain orientations were positive perpendicular and positive
parallel in summer and winter, respectively. The latter may have

339
been more effective in winter, when the sun declination was high,
since foreleg elevation of the forebody not only reduced contact
with the cold substrate but also brought the long axis of the body
closer to a perpendicular angle with the incoming radiation. Object
utilization was also more frequent during winter morning and evening
basking. Negative parallel orientation, minimizing radiant heat gain
from the sun, was more commonly seen during the summer than the
winter.
Iguanas may preferentially heat their proximal colon, located on
the left side of the anterior abdomen, by orienting that side toward
the sun.
Patterns of travel during forays changed in a regular fashion
during the February and October day. Iguanas most commonly moved
clockwise during the morning (peak at 1000 to 1100), linearly at
midday, and counterclockwise in the afternoon (maximum plateau
between 1100 and 1500), probably visually keying the change to the
movement of the sun across the sky.
Hie rate of heating of dried skin patches from various parts of
the iguana's body increased faster posterior to the imbricate edges
of the scales than anterior as the angle of incidence of the heat
lamp radiation increased, especially for the more highly imbricated
patches. Since the dorsal scale rows ran obliquely toward the mid-
sagittal crest at an anterior angle of approximately 60°, iguanas
should orient at greater than 90° to maximize heat input. This did
appear to be the case, though more work is needed to be sure.
18. Iguanas deliberately changed their substrate contact by using neck, tho¬
racic, abdominal, foreleg, and hindleg muscles to increase heat input

340
from the substrate or to avoid excessively hot substrates. Maximum
substrate contact varied from approximately 13.4 to 16.0 percent of
total surface area. Head-neck contact, foreleg participation, hind¬
leg participation, and lateral compression accounted for maxima of
1.5, 48, 30, and 16 percent of maximum possible substrate contact,
respectively.
T . T j and T-, all had an effect on the amount of substrate contact
aJ sJ b
in a low, radiant heat exchange rate environment. A T greater than
Th and a T^ equally less than T^ resulted in low substrate contact
and decreasing T-^. Conversely, a T^ less than T^ and Tq equally
greater than T-^ resulted in a high substrate contact and increasing
T, . Once in the preferred range in a cool air environment, substrate
contact decreased even though T remained constant and above T-, . Thus,
the iguana increased cool air contact to reduce further T-^ rise.
Iguanas were quite sensitive to small Tg - T^ and T^ - T^ differences.
During radiant heating in the photothigmotron, substrate contact
only gradually increased as T^ increased, even when was consider¬
ably higher than 2^. Since the same gradual relaxation was consistently
observed in the field, especially during the morning heating phase,
it may be a means of either maximizing radiant input while minimiz¬
ing the normal heat loss to a cooler substrate, or of achieving a
balance between radiant and thigmothermic heating. In fact, substrate
contact in the photothigmotron was greater at higher substrate tempera¬
tures. However, the initial low body contact may also be due to
greater wariness at low T-^, noted in the field. Holding the head
higher off the ground increased the visual field, decreasing the
chance of being caught unprepared for escape at a where recovery
from exercise occurred at a low rate.

341
Measurement of substrate temperature beneath maximally prostrate
iguanas commonly revealed a small positive - T__ difference rather
than the expected thigmothermia. Nonetheless, the rate of heat loss
in this posture could easily be less than in a more upright posture
due to decreased convective heat loss and probable localized ventral
vasoconstriction. Also, the energy expenditure to maintain the
prostrate posture was minimal.
19. Behavioral responses during cooling were lost in the order of decreas¬
ing neural pathway complexity. Lizards could still bloat below the
CTMin, adaptive in that a nearly helpless iguana could wedge itself
securely in its burrow or, perhaps, intimidate a predator. As
expected, the CTMin, 12.5 C, was about equal to the lowest environ¬
mental temperature recorded during the study.
20. Cyotura caminata, was a true panting lizard. At least five effector
outputs probably increased proportionally above a panting threshold
temperature; that is, gaping time per minute, degree of gape, extent
of elevation and blood engourgement of the tongue, respiratory rate,
and salivation rate. The function of panting is likely the cooling
of the brain at high head temperature.
Eye bulging started soon before or soon after panting and was
correlated with a reduction in the heating rate of the head at
high head temperature. It probably also served directly as a means
of removing ecdysing skin from around the eyes.
Head temperature was consistently higher than gut temperature dur¬
ing basking (X head-gut temperature = 1.0 C, range = -0.1 to 1.6 C),
the head heating up to six times faster than the gut during one short,
two-day monitoring period. Rapid head heating presents the

342
heliothermic, high T^, ectotherm with an additional selective
advantage for evolving a precise panting response and jugularis
constriction.
The gut temperature at the panting threshold decreased from 32.8 C
to above 43.8 C as the rate of heating increased. This is further
circumstantial evidence for a panting regulator input from skin
surface, temperature receptors. The panting time after shading was
well correlated with the sum of gut and head temperature lag times,
indicating that both temperatures may be important in sustaining
panting.
The CTMax was approximately 46.2 C and the LM approximately 48.0
C. The minimum thermal safety margin was 4.8 C at midmorning, more
than adequate for slow heating adults to avoid fatally overheating
by finding- shade.
21. Integumental reflectivity varied considerably over the body surface,
being least labile and highest on the ventral tail and abdomen and
lowest and quite labile on the dorsal surface. The change in
reflectivity and coloration with increasing probably served both
concealment and thermoregulatory functions. The low reflectivity,
dark phase characteristic of morning basking was well matched
spectrally to the algal encrusted calcium carbonate substrates on
which iguanas commonly basked. The initiation of the light phase,
with its tinges of green and yellow evident in Water Cay iguanas,
correlated well with the T^ range of the first foray into the sun-
vegetational shade mosaic. The rate of heat gain in the light phase
was estimated to be 15.3 percent less than in the dark phase under
one set of defined thermal conditions.

343
The histology and distribution of integumental sensory spots
suggests that they serve as mechanoreceptors. However, since jaw
rubbing, sand swimming, and hard substrate rubbing with the ventral
body surface all occurred in contexts suggesting substrate tempera¬
ture detection, sensory spots may also be thermal receptors.
22. The mean, uncorrected conductance cooling/heating ratios in the cage
and on the jig were 0.658 and 0.721, respectively. Iguanas always
heated faster than they cooled, with a slightly but not significantly
lower rate in the cage than on the jig. The jig ratio was at the
low end of values reported for other lizards. Mean metabolic
corrections of 62 and 67 percent of the maximum metabolic rate at
30 C were required to increase the C/H conductance ratios to one
for iguanas on the jig and in the cage, respectively. Since this
seems unrealistically high, Cyctura does appear to have some physio¬
logical thermoregulatory control, presumably via locally regulated,
peripheral vasomotor control.
Climbing and walking in the cage and struggling on the jig occurred
at significantly higher temperatures during heating than during
cooling, corresponding to active avoidance of high and low 's,
respectively. The same was true for defecation.
The respiratory rate was significantly greater during heating,
perhaps due to thermal hysteresis, either between gut and brain
temperatures or between gut and lung temperatures.
23. The log of the resting metabolic rate was directly and linearly
related to T^ and was consistently above expectation, probably
due to incomplete recovery from forced maximum exercise at the time
of measurement. The log of maximum oxygen consumption vs. T^ was

344
best fitted to a second degree polynomial. The maximum rate occurred
at 43-44 C, near the maximum voluntary tolerance. Maximum rates
were similar to another fairly sluggish herbivore, Sauromalus
hispidus, and lower than rates for the active carnivore, Varanus
gould'i't. The maximum rate of oxygen consumption was less thermally
sensitive than the resting rate above 22.5 C, slightly below the
Tb of emergence, reaching a one-half as large as 42 C. The
factorial scope was also largest at slightly below emergence 2^,
decreasing from about a maximum of six to about four in the mean
preferred T\ range. As expected, maximum oxygen scope occurred at
38.1 C, slightly above preferred T^.
24. The log of resting heart rate vs. Th was fitted best by two straight
lines meeting at 25.0 C. This commonly observed break occurred at a
higher T^ than in more cold resistant lizard species. The log of
maximum heart rate was fitted best to a second degree polynomial.
The resting and maximum heart rate lines extrapolated to equality
at 47.4 C, near the lethal maximum. Maximum heart rate was never
more than 1.69 times the resting rate between 13.4 and 43.0 C. As
expected, the heart rate increment maximized at 38.5 C, near the
preferred T^. Both resting and maximum exercise oxygen pulse
declined between 13.4 and 43.0 C, by factors of 2.3 and 3.0,
respectively. Thus, iguanas depended on increased heart rate
rather than increased stroke volume and A-V difference to deliver
the increased oxygen demand to the tissues as T^ increased. This
contrasts to the low contribution of heart rate increase at any one
T-, (X = 14.7 percent between 15 and 40 C using Gatten's 1974b equa¬
tion) to the satisfaction of the increased oxygen demand during
activity.

345
The high similarity between preferred T^, 37.0 C, and 2^'s for
maximum heart rate increment, 38.5 C, maximum oxygen scope, 38.1 C,
and maximum early rate of recovery from maximum exercise, 37.3 C,
may in part be due to the high stenothermality of Cyolura ccurtnata.
25. Maximum burst activity could be maintained for only the first thirty
seconds during hand manipulation. Iguanas were practically helpless
between feeble struggling bouts during the fourth to ninth minutes.
They depended heavily on anaerobic metabolism to attain maximum
activity, as indicated by the large oxygen debt. Iguanas recovered
slowly from maximum exercise, requiring a mean of 82 to 118 minutes
to reach the resting metabolic rate. Recovery occurred in the
shortest time at a T^ of 25 C, followed by 15 and 20 C, 31 and 37 C,
and finally, 42 C. The recovery rate increased as T-, increased,
with the log of the overall recovery rate and the log of the rate
during the first five minutes vs. T-^ fitted best by second degree
polynomials, and the log of the rate from minute 15 to full recovery
fitted best by a straight line. The maximum rate of recovery during
the first five minutes was well correlated with preferred TThis
may be adaptive, since small oxygen debts were undoubtedly more
common than large debts in the field. The alactacid debt averaged
only 8.5 percent of the total debt (3.0-20.2 percent) and was
practically T^ independent, whereas the lactacid debt averaged 91.5
percent of the total debt and increased curvilinearly with 21 . Over¬
all, approximately the equivalent of the resting rate of metabolism
was devoted to recovery from the oxygen debt, over and above resting
metabolism devoted to other processes.

346
Strenuous activities were usually of short duration in the field,
probably because recovery from extended maximum exercise was such a
slow process. Walking was probably primarily an aerobic activity,
based on its discontinuous nature and comparison of rates of travel
with other lizards for which blood lactate levels were measured.
More strenuous activities were also either discontinuous (climbing,
digging) or of short duration and not soon repeated (copulations,
escapes to burrows, male-male territorial chases). Two infrequent
activities, nesting and male-male fights, probably did result in an
oxygen debt. Protracted copulation, fighting, chases, and nesting
in field cages and the Gainesville pen were followed by depressed
activity for the rest of the day. The inability of iguanas to
sustain burst activity for periods longer than 30 seconds, coupled
with the ineffectuality of their static defense, were probably the
main causes of their extirpation from Water Cay by dogs and feral
cats.

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BIOGRAPHICAL SKETCH
David Leslie Auth was born on July 7, 1945, in Plainfield, New
Jersey. His family moved almost immediately to Wilmette, Illinois,
where he spent practically all of the next 22 years. He graduated in
1963 from New Trier Township High School and from Northwestern University
in 1967. He entered the University of Florida in the fall of 1967 and
received the Master of Science degree in zoology in June of 1969.
Drafted during the war in Vietnam, he spent the next four years in the
United States Navy, first as a cryptologic technician with the Naval
Security Group in Puerto Rico, then as a drug counselor at the Naval
Drug Rehabilitation Center in Jacksonville, Florida. He returned to
the University of Florida in January of 1974 and has continued work
toward the Doctor of Philosophy to the present time.
He is a member of the American Association for the Advancement of
Science, the American Institute of Biological Sciences, the American
Society for Ichthyologists and Herpetologists, the Ecological Society
of America, and Phi Kappa Phi.
367

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Archie Carr
Graduate Research Professor of
Zoo 1ogy
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
0 fj.
Kaufmann7
Jc^hi^ H.
Processor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
Professor of Zoology

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree of
Doctor of Philosophy.
This dissertation was submitted to the Graduate Faculty of the Department
of Zoology in the College of Liberal Arts and Sciences and to the Graduate
Council, and was accepted as partial fulfillment of the requirements for
the degree of Doctor of Philosophy.
April, 1980
Dean, Graduate School

UNIVERSITY OF FLORIDA
1262 08667 042 8





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