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198 IGUANA  •  VOLUME 13, NUMBER 3  •  SEPTEMBER 2006 MUENSCH, LEININGER, WERTH, FAWKS, AND THOMAS

World War II-era spotlights testify to the military role of Coconut Island. Today, they remain among decorative plantings utilized exten-
sively by Cuban Brown Anoles (Anolis sagrei).

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Originally, Moku O Lo‘e, owned by the Bishop family
estate, was used by shepherds and local fishermen. During

the 1930s, Christian Holmes, owner of Hawaiian Tuna Packers
(now Coral Tuna) and heir to the Fleischmann yeast fortune,
purchased the island for his tuna-packing factory. At that time,
the island was 12 acres in size and had several coconut trees,
which is how it got its popular name, Coconut Island. Holmes,
unhappy with the small size of the island, had it expanded to 28
acres and also created many fishponds. Holmes imported hun-
dreds of exotic plants to the island, constructed a large saltwater
swimming pool equipped with a slide and a diving board, built
outdoor bars at various points around the island, introduced a
bowling alley, and reconstructed a shooting gallery that he had
bought at an amusement park in San Francisco. Coconut Island
even housed a small zoo for a short time. Animal residents
included donkeys, a giraffe, monkeys, and a baby elephant. After
Holmes’s death, these animals became the foundation for the
Honolulu Zoo.

During the war, the military used the island as an R&R
post for officers. In 1947, a group of five wealthy oilmen bought

The Anoles of Coconut Island, 
Kane‘ohe Bay, O‘ahu, Hawai‘i

Alexander J. Muensch, Penny D. Leininger, Dusty E. Werth, Angela M. Fawks, and Sydney M. Thomas

Department of Biology, Avila University, Kansas City, MO 64145

IGUANA  •  VOLUME 13, NUMBER 3  •  SEPTEMBER 2006 199ANOLES OF COCONUT ISLAND

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Coconut Island in Kane‘ohe Bay. Photograph courtesy of the Hawaii
Institute of Marine Biology.

Lagoon on Coconut Island. Plantings in the foreground were occupied by Anolis sagrei.

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the island. Eventually, one of them, Edwin Pauley, bought out
the interests of the other four and became sole owner of the
island, where his family spent their summers. Many famous peo-
ple visited Coconut Island as Pauley’s guests. Included among
them were Harry Truman, Lyndon B. Johnson, John Wayne,
Red Skelton, Richard Nixon, and Ronald Reagan. In 1951,
Pauley helped establish the Hawai‘i Marine Lab. The name was
changed in 1965 to the Hawai‘i Institute of Marine Biology,
now affiliated with the University of Hawai‘i.

Amphibians and Reptiles of Coconut Island
No terrestrial amphibians or reptiles are native to Hawai‘i, but
many species have become established, introduced primarily as
escaped or released pets. Due to frequent movement of people
and materials back-and-forth from O‘ahu to Coconut Island,
many of the species established in the Kane‘ohe area now occur
on Coconut Island as well. In 2001, documented species
included the Cane Toad (Bufo marinus), Stump-toed Gecko
(Gehyra mutilata), Mourning Gecko (Lepidodactylus lugubris),
Metallic Skink (Lampropholis delicata), and Island Blind Snake
(Ramphotyphlops braminus). In 2004, two species of Anolis, 
A. carolinensis and A. sagrei, were found on the island. Both had
highly restricted distributions, suggesting that both introduc-
tions had occurred shortly before their discovery.

Coconut Island Anoles
Anolis carolinensis, the Green Anole, is moderately sized (SVL of
Hawaiian animals = 51–76 mm, total length = 125–230 mm).
These lizards are native to the southeastern United States.
Populations also have become established in Europe (Spain),
Japan (Bonin and Ryukyu islands), the West Indies (Grand
Bahama Island, Anguilla), and the Pacific (all of the major
Hawaiian islands; Guam; Tinian and Saipan in the Northern
Mariana Islands; Yap; Koror and Malakal islands in the Republic
of Palau (or Belau). An introduction on Sand Island, Midway
Atoll, failed.  Green anoles were first reported on O‘ahu in 1950,
when they were initially misidentified as A. porcatus (a closely
related Cuban species).

Anolis carolinensis is a “trunk-crown ecomorph” (ecomor-
phologies anatomically and behaviorally influence microhabitat
use as a way of reducing interspecific competition). These lizards
are abundant in Hawai‘i, especially in gardens and resorts, where
introduced plants and insect prey provide suitable conditions.
Although they will forage on the ground on occasion, Green

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Mourning Geckos (Lepidodactylus lugubris), here shedding its skin in
one of the suites on Coconut Island, and Stump-toed Geckos (Gehyra
mutilata) are abundant.

Cane Toads (Bufo marinus) are well-established on Coconut Island.

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IGUANA  •  VOLUME 13, NUMBER 3  •  SEPTEMBER 2006 201ANOLES OF COCONUT ISLAND

Anoles spend nearly all of their time on elevated perches in
bushes, trees, or artificial substrates like fences and buildings.

Anolis (Norops) sagrei, the Cuban Brown Anole, is a mod-
erately sized (SVL of Hawaiian animals = 38–64 mm, total
length = 130–210 mm) lizard native to Cuba, Bahamas, and
Cayman Islands, but which has become established in various
mainland (e.g., southeastern United States, Belize, and southern
México to northern Honduras) and insular (e.g., St. Vincent,
Grenada, Jamaica, Hawai‘i [O’ahu and K’auai], and Taiwan)
locales. Brown Anoles were first reported on O‘ahu in the late
1970s and early 1980s.

Anolis sagrei is a “trunk ground ecomorph,” meaning that
it spends most of its time within two meters of the ground, and
is generally less arboreal than A. carolinensis. Anolis sagrei is most
commonly found in shrubs, on tree trunks, the ground, and
rock piles, but readily exploits human dwellings, planters, and
fences. It feeds on small invertebrates (mostly insects), but will
occasionally take small lizards. Foraging strategies typically
exploit an ambush mode, often involving a rapid descent from
a vertical, head-down, sit-and-wait posture. This species can also
be found in a vertical, heads-up position while basking, espe-
cially in the morning.

Both species are habitat generalists, a trait that appears to
have served them well as colonists of areas far from their native
ranges. Like most anoles, both are diurnal, largely arboreal, pri-
marily insectivorous, and intensely territorial. Several females
and juveniles may occupy the territory of a single male. Adult

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Male Anolis carolinensis from Coconut Island (2004).

Female Anolis carolinensis from O‘ahu (2006). Although apparently extirpated on Coconut Island, Green Anoles remain abundant elsewhere in the
Hawaiian Islands.

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Female Anolis sagrei from Coconut Island (2006). 

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Male Anolis sagrei from Coconut Island (2006). 

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IGUANA  •  VOLUME 13, NUMBER 3  •  SEPTEMBER 2006 203ANOLES OF COCONUT ISLAND

males frequently exploit different microhabitats, such as higher
perches, than females or juveniles.

Somewhat more robust than the more gracile Green Anole
and apparently more aggressive, introduced A. sagrei has largely
displaced the native Green Anole, especially in disturbed habi-
tats, where the two species are sympatric in peninsular Florida
and elsewhere in the southeastern United States.

Methods
During a visit to Coconut Island in March 2006, we sought to
establish the ranges of both species, which we expected to have
increased since the discovery of the two populations in 2004.
We also examined microhabitat use by anoles in different size
classes to test a null hypothesis that perches of lizards of differ-
ent sizes do not vary.

At various times of day and under varying weather condi-
tions (which were often rainy during our stay), we searched
microhabitats where anoles were most likely to be found, con-
centrating on areas near where they were first observed in 2004.
For each anole sighted, we recorded time, size class, height above
ground, perch diameter (if applicable), orientation, and various
behaviors. Anoles were categorized into three classes: “1” for
adult males, “2” for subadult males and adult females (which are
sometimes difficult to distinguish from a distance), and “3” for
juveniles of indeterminate sex. Orientation on a perch was
numerically interpreted as 1 = vertical, facing up; 2 = horizon-
tal; 3 = vertical, facing down. We used StatView 5.0 (SAS
Institute, Inc., Cary, North Carolina) for statistical analyses.
Means are presented ± 1 standard error.

Results
We were unable to find Anolis carolinensis. Shrubs and intermin-
gled vines in the area where they were most abundant in 2004
had been cleared, although considerable, apparently suitable
habitat remained, including large Ficus trees where individuals
had been observed two years previously.

Anolis sagrei remained abundant in the area where first seen
in 2004, and these lizards had expanded their range to nearby
areas and to a peninsula characterized by only marginal habitat.
Considerably more apparently suitable habitat exists throughout
much of the island, including sites adjacent to those where we
found lizards. We found no Brown Anoles in the area that had
been occupied by A. carolinensis in 2004.

For obvious reasons, our investigation of microhabitat use
was restricted to A. sagrei. We made 82 observations (12 adult
males, 50 subadult males/adult females, 18 juveniles, and 2 for
which size class was not recorded). Most (n = 65) were under
cloudy skies or during rain. Seven observations were of individ-
uals on the ground, 18 on rocks, 17 on artificial perches (e.g.,
pipe, box, debris, planter), 8 on stumps or logs, and 32 on veg-
etation. Of the latter, 20 were on leaves. We observed one indi-
vidual eating (small arthropod), one mating pair, one other inci-
dence of courting behavior, and two territorial displays. Fifteen
instances of elicited escape behavior included 10 individuals
jumping to the ground or crawling under an object on the
ground and five lizards jumping onto vegetation, four of them
from rocks or the ground.

Mean perch height for all size classes was 35.1 ± 2.8 cm (n
= 79), mean perch diameter was 8.7 ± 1.9 cm, and mean orien-
tation was 1.9 ± 0.1, or predominantly horizontal.

Perch heights of adult males (53.8 ± 6.6 cm, n = 12) were
significantly higher than those of subadult males and adult
females (34.1 ± 3.5 cm, n = 48) and those of juveniles (29.1 ±
5.1 cm, n = 17; ANOVA, P = 0.02). Mean perch height differ-
ences between classes 1 and 2 (19.7 cm) and between classes 1
and 3 (24.6 cm) were significant (Fisher’s PLSD, both P = 0.01).
However, mean perch height differences between classes 2 and
3 (5.0 cm) were not significant (P = 0.45).

Perch diameters used by anoles in different size classes did
not differ significantly (ANOVA, P = 0.82), nor were any dif-
ferences between any two classes significant (Fisher’s PLSD,
between classes 1 and 2 P = 0.65, between classes 1 and 3 P =
0.54, between classes 2 and 3 P = 0.69). Similarly, orientations
did not differ significantly among classes (ANOVA, P = 0.97),
nor between individual classes (Fisher’s PLSD; not valid between
classes 1 and 2 due to limited sample sizes; between classes 1 and
3, P = 0.85; between classes 2 and 3, P = 0.80).

Discussion
The apparent absence of Anolis carolinensis may represent one of
the few documented failed colonization attempts by any invasive
reptile. Despite the altered vegetation in the area where these
lizards were most commonly encountered in 2004, the presence
of apparently suitable habitat, including some used by Green
Anoles in 2004, suggests that other factors are involved. Because
we found no A. sagrei anywhere near where A. carolinensis had
been found previously, we have ruled out displacement as a con-
sequence of interspecific competition. In a concurrent survey of

Distribution of Anolis sagrei in 2004 (narrow dashed line) and in 2006
(solid narrow line) and of A. carolinensis in 2004 (broad solid line).

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An adult male Cuban Brown Anole (Anolis sagrei) seeks shelter in vegetation near the base of a large Banyan Tree (Ficus benghalensis) on Coconut
Island in 2006. Ecologically versatile, these anoles readily exploit large trees, small bushes and shrubs, as well as walls, buildings, and human debris.

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birds, we did find many more introduced Red-Vented Bulbuls
(Pycnonotus cafer) and a lesser number of Red-whiskered Bulbuls
(P. jocosus) than had been observed during similar surveys in 2001
and 2004. Both species are known predators of diurnal lizards,
both in their native range (Indian Subcontinent through south-
eastern Asia) and in Hawai‘i. The combination of altered vegeta-
tion and presumably increased predation pressures may have
combined to eliminate Green Anoles from Coconut Island.

Although Anolis sagrei remains present and has expanded
its range on the island, the extent of the expanded range in light
of abundantly available and apparently suitable habitat is much
less than we had expected. Because vegetation in the areas occu-
pied by Brown Anoles had not been substantively altered, bul-
buls may be implicated in the slower-than-expected exploitation
of suitable habitats by A. sagrei.

Activity may have been suppressed by inclement weather,
but observed perch choices and escape behaviors were compat-
ible with expectations for a trunk-ground anole. Males selecting
higher perches and the sexual size dimorphism evident in these
lizards may serve to spatially partition resources among different
size classes, thereby reducing intraspecific competition. Larger
animals (i.e., adult males) perching higher than smaller con-
specifics may indicate that the former are able to take advantage
of their size to monopolize microhabitats that facilitate optimal
foraging. These more elevated positions also may result in more
distinct territorial boundaries between conspecific males.

Data regarding perch diameters were equivocal due to too
few observations of adult males and juveniles (n = 1 in each
instance). This does reflect the large number of observations of
animals using perches other than those for which diameter data
were appropriate (e.g., the ground, rocks, and artifical perches
such as boxes, refuse, and even buildings). This conforms to
observations for this species in other areas, and to expectations
for a trunk-ground ecomorph. The same frequency of occurrence
on the ground and common use of rocks and artificial perches
were undoubtedly responsible for the fact that most anoles were
oriented horizontally, which did not vary among size classes.

Acknowledgments
We thank Avila University instructors David Wissmann and
Robert Powell for the opportunity to conduct this project on
Moku O Lo‘e and the Hawai‘i Institute of Marine Biology for
permission to work on the island. Robert W. Henderson and
Sandy Echternacht made helpful suggestions on an earlier draft
of this manuscript.

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