10 Herpetological Review 37(1), 2006
ZOO VIEW
Herpetological Review, 2006, 37(1), 10?13.
? 2006 by Society for the Study of Amphibians and Reptiles
Wild and Ferocious Reptiles in the Tower of
London
JAMES B. MURPHY
National Zoological Park, Smithsonian Institution
Washington, DC 20008, USA
e-mail: jbmurphy2@juno.com
The Tower of London is best known as a prison where human
inmates were subjected to harsh and cruel punishments so it might
be surprising that this forbidding venue also served as a reposi-
tory for wild animals. The Royal Collection of Wild and Fero-
cious Beasts kept at the Tower of London Menagerie spanned ca.
600 years, beginning in 1245 (Loisel 1912; Keeling 1992; Hahn
2004; Fig. 1). During that time, leopards, lions, tigers, ostriches,
wolves, kangaroos, zebras, elephants, polar bears, monkeys, ba-
boons, eagles and other birds of prey, macaws and other psitta-
cines, and a stunning variety of other creatures could be found in
the Tower as tourist attractions. Although seemingly not an ideal
facility for reptiles with its stone walls and barred dens, at various
times over the span of six centuries, the rattlesnake, the Indian
boa, the anaconda, and the alligator lived in a large outdoor enclo-
sure surrounded by a moat. All of these reptiles are illustrated in
Edward Turner Bennett?s book, ?The Tower Menagerie: Compris-
ing the Natural History of the Animals Contained in that Estab-
lishment, with Anecdotes of Their Characters and History. Illus-
trated by Portraits of Each, Taken from Life, by William Harvey,
and Engraved on Wood by Branston and Wright? in 1829. Bennett
described the difficulties in drawing animals, ?The whole of the
drawings are from the pencil of MR WILLIAM HARVEY, who,
in seizing faithful and characteristics portraits of animals in rest-
less and almost incessant motion, has succeeded in overcoming
difficulties which can only be appreciated by those who have at-
tempted similar delineations.?
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centrations of steroid hormones in layers and biopsies of chelonian
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CLARK, P. J., M. A. EWERT, AND C. E. NELSON. 2001. Physical apertures as
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Maternally derived yolk hormones vary in follicles of the painted turtle,
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Carbon dioxide influences environmental sex determination in two
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FIG. 1. Tower of London. From Bennett in 1829. Credit: Courtesy of
Smithsonian Institution Libraries, Washington, DC.
Herpetological Review 37(1), 2006 11
Alfred Cops worked at the Exeter ?Change in the Strand but left
this job in 1822 to take charge of the Royal Menagerie at the Tower.
Daniel Hahn (2004) described his skills as an animal caretaker,
?Cops was a professional zoologist and an expert on animal be-
havior, a man with extensive training and experience looking af-
ter captive animals . . . It was expected that such a man would be
able to expand and improve the Menagerie, making it exciting
and competitive in a way that it had not been for some years, and
at the same time improve the animals? living conditions sufficiently
to satisfy the newly fashionable anticruelty lobbies.?
Given that rattlesnakes may be delicate to keep in captivity, it is
amazing that over 100 rattlesnakes (identified Crotalus horridus
Linn. by Bennett) survived sea voyages from the New World to be
cared for by Cops in the Tower; perhaps their survival is a testa-
ment to Cops? ability (Fig. 2).
Here is Bennett on rattlesnake behavior, ?It was long believed,
and the notion is still popularly current, that they possessed the
power of fascinating their victims, which were thought to be so
completely under the influence of their glance as to precipitate
themselves of their own accord into the open throat of their en-
emy; but the truth appears to be that they actually inspire so great
a degree of terror that the animals selected for their attacks are
commonly rendered incapable of offering such resistance as might
otherwise be in their power, or even of attempting to escape from
their pursuit.?
Cops nearly bred the Tower python, called Indian Boa (Python
Tigris Daud., now Python molurus) by Bennett (Fig. 3). The py-
thon laid a clutch of eggs which did not hatch. Bennett described
the event, ? The individual figured at the head of the present ar-
ticle is a female; a fact which was proved by the remarkable cir-
cumstance of her producing in May last, after having been more
than two years in the Menagerie, a cluster of eggs, fourteen or
fifteen in number, none of which, however, were hatched, although
the mother evinced the greatest anxiety for their preservation, coil-
ing herself around them in the form of a cone, of which her head
formed the summit, and guarding them from external injury with
truly maternal solicitude.? This account is certainly one of the first
to describe brooding behavior in pythons. Three years later, natu-
ralist Lamarre-Picquot lectured before the Academie de France
about the brooding of a Python molurus bivittatus in the Jardin
des Plantes in Paris (Fig. 4) and claimed that the female not only
coiled around the clutch but produced ?noticeable warmth,? a find-
ing expressed again in a note in 1842. This observation was widely
criticized as speculative, hazardous, and questionable by mem-
bers of the Academie, especially Auguste Dum?ril (1842). The
thermogenesis controversy was finally settled over 130 years later
when sophisticated temperature and gas exchange recording de-
vices were available. Victor H. Hutchison, Herndon G. Dowling,
and Allen Vinegar published two important papers on metabolism,
energetics and thermoregulation in brooding female pythons at
the New York Zoological Society (Hutchison et al. 1966; Vinegar
FIG. 3. Tower python, called Indian Boa (Python Tigris Daud., now
Python molurus) by Bennett (1829), laid a clutch of 14?15 eggs which
did not hatch. Credit: Courtesy of Smithsonian Institution Libraries, Wash-
ington, DC.
FIG. 2. Over 100 rattlesnakes (Crotalus horridus Linn.) lived in the
Tower of London. From Bennett in 1829. Credit: Courtesy of Smithsonian
Institution Libraries, Washington, DC.
FIG. 4. Indian python (Python molurus) incubating eggs at Menagerie
Jardin des Plantes. Illustration reproduced from S. G. Goodrich?s
?Johnson?s Natural History, Comprehensive, Scientific, and Popular, Il-
lustrating And Describing The Animal Kingdom With Its Wonders And
Curiosities, From Man, Through All The Divisions, Classes, And Orders,
To The Animalculae In A Drop of Water; Showing The Habits, Structure,
And Classification Of Animals, With Their Relations To Agriculture,
Manufactures, Commerce, And The Arts. Volume 2 in 1870.? Credit: Cour-
tesy of Smithsonian Institution Libraries, Washington, DC.
12 Herpetological Review 37(1), 2006
et al. 1970). The abstract in the first paper reads: ?At varying en-
vironmental temperatures, measurements of body temperatures and
gas exchange of a female Indian python (Python molurus bivittatus)
show that during the brooding period this animal can regulate its
body temperature by physiological means analogous to those in
endotherms. Ambient temperatures below 33?C result in spasmodic
contractions of the body musculature with a consequent increase
in metabolism and body temperature.?
The Anaconda, called Python Tigris Var. by Bennett, were likely
Ceylonese pythons, Python molurus pimbura (Fig. 5). Cops dis-
covered that captive snakes may have impressive feeding responses
when one of his pythons grabbed his hand and threw two coils
around his neck while being fed. Bennett told the story, ?His own
exertions, however, aided by those of the under keepers, at length
disengaged him from his perilous situation; but so determined was
the attack of the snake that it could not be compelled to relinquish
its hold until two of its teeth had been broken off and left in the
thumb.?
Coote (2001) speculated that the first American Crocodile,
Crocodylus acutus, received by the London Zoological Society in
FIG. 5. The Anaconda, called Python Tigris Var. by Bennett, were likely
Ceylonese pythons, Python molurus pimbura. Credit: Courtesy of
Smithsonian Institution Libraries, Washington, DC.
1831, may have been the Tower ?Alligator? (Fig. 6). However,
the drawing by William Harvey looks like an American alligator
and is called Crocodilus lucius Cuv., an early name for the alliga-
tor. Robert Huish?s drawing in 1830 also appears to be an alligator
(see Fig. 2 in Herpetological Review 36(3):230). In both illustra-
tions, the alligator?s head is abnormally small.
The Tower collection of animals was presented by the King to
the Zoological Society of London in 1831 (Peel 1903:180). Eigh-
teen years later, the London Zoo in Regent?s Park opened the
world?s first zoo reptile building (Fig. 7).
Acknowledgments.?Smithsonian National Zoological Park Librarian
Alvin Hutchinson, and Leslie Overstreet and Daria Wingreen in the Jo-
seph F. Cullman 3rd Library of Natural History (from the SI Special Col-
lections Department) helped locate historical literature. Judith Block and
Jon Coote reviewed early drafts of this manuscript. I thank Kraig Adler
for providing the illustration of the reptile building at London Zoo.
This contribution is dedicated to Jon Coote, who has expanded our
understanding and appreciation of early herpetocultural practices through
his writings.
LITERATURE CITED
BENNETT, E. T. 1829. The Tower Menagerie: Comprising the Natural His-
tory of the Animals Contained in that Establishment, with Anecdotes
of Their Characters and History. Illustrated by Portraits of Each, Taken
from Life, by William Harvey, and Engraved on Wood by Branston
and Wright. Printed for R. Jennings, London.
COOTE, J. G. 2001. A history of western herpetoculture before the 20th
century.  In W. E. Becker (ed.), 25th International Herpetological Sym-
posium on Captive Propagation and Husbandry, pp. 19?47. Interna-
FIG. 6. Tower ?Alligator? called Crocodilus lucius Cuv. by Bennett in
1829. Note the abnormally small head. Credit: Courtesy of Smithsonian
Institution Libraries, Washington, DC.
FIG. 7. Description of London Zoo reptile building from The Illustrated
London News on 2 June 1849. The beginning of the article reads as fol-
lows: ?The new reptile house in the Gardens of the Zoological Society in
the Regent?s Park will ultimately form one of the most instructive, as it is
the most novel and original feature in this delightful institution. The col-
lection already contained in it is so unexpectedly brilliant, considering
the small number of reptiles previously exhibited in the Menagerie, that
we cannot but anticipate the most important results in the study of this
singularly interesting division of the animal kingdom.? Credit: illustra-
tion provided by Kraig Adler.
Herpetological Review 37(1), 2006 13
tional Herpetological Symposium, Detroit, Michigan.
DUM?RIL, A. 1842. Sur le d?veloppement de la chaleur dans les oeufs des
serpents, et sur l?influence attribu?e ? l?incubation de la mere; par M.
Dum?ril (Concerning the production of heat in the eggs of snakes, and
the influence attributed to the incubation of the mother; by M. Dum?ril).
Comptes Rendus Acad. Sci. Paris 14:193?210.
HAHN, D. 2004. The Tower Menagerie. Jeremy P. Tarcher/Penguin, New
York.
HUISH, R. 1830. The Wonders of the Animal Kingdom Exhibiting Delin-
eations of the Most Distinguished Wild Animals in the Various Menag-
eries of the Country. Thomas Kelly, London. [description of the alliga-
tor housed in the Royal Menagerie of London (pp. 51?52).].
HUTCHISON, V. H., H. G. DOWLING, AND A. VINEGAR. 1966. Thermoregula-
tion in a brooding female Indian python, Python molurus bivittatus.
Science 151:694?696.
KEELING, C. H. 1992. A Short History of British Reptile Keeping. Clam
Publications, Guilford, UK.
LAMARRE-PICQUOT, P. 1835. L?lnstitut 3: 70.
______. 1842. Trois?me m?moire sur l?incubation et autres ph?nom?nes
observ?s chez les ophidiens (Third report on the incubation and other
phenomena observed in the snake house). C. R. Acad. Sci. Paris 14:164.
LOISEL, G. 1912. Histoire des m?nageries de l?antiquit? ? nos jours (His-
tory of Menageries from Antiquity to Present Times). O. Doin et fils,
Paris.
PEEL, C. V. A. 1903. The Zoological Gardens of Europe. Their History
and Chief Features. F. E. Robinson & Co., London.
VINEGAR, A., V. H. HUTCHISON, AND H. G. DOWLING. 1970. Metabolism,
energetics, and thermoregulation during brooding of snakes of the ge-
nus Python (Reptilia, Boidae). Zoologica (New York) 55:19?48.
ARTICLES
Herpetological Review, 2006, 37(1), 113?120.
? 2006 by Society for the Study of Amphibians and Reptiles
Reproduction and Growth of Seven Species of
Dwarf Geckos, Sphaerodactylus (Gekkonidae), in
Captivity
RUB?N REGALADO
4841 West 3rd Avenue
Hialeah, Florida 33012, USA
e-mail: regalado05@bellsouth.net
A diversity of reproductive, growth, and survival strategies
(Shine and Charnov 1992) have made lizards model organisms in
the investigation of how environmental factors and phylogenetic
constraints shape life-history traits (Clobert et al. 1998; Dunham
1978; Shine and Greer 1991; Sinervo and Adolph 1994; Tinkle et
al. 1970). However, ecological studies on the family Gekkonidae,
the second most speciose family of lizards (Uetz 2000), are few
(see Vences et al. 2004 and references therein). Small and usually
invariant clutch sizes (Kluge 1987, but see Colli et al. 2003; Shine
and Greer 1991), nocturnal activity, and microhabitats that are
usually not conducive for easy field observation and manipula-
tion may have biased researchers against selecting geckos for life-
history study. The lack of life history information is even more
evident in dwarf geckos, genus Sphaerodactylus (Krysko et al.
2003; L?pez-Ortiz and Lewis 2002). However, some dwarf gecko
species are believed to reach the highest known population densi-
ties of terrestrial vertebrates in some areas (Rodda et al. 2001).
Miniaturization in Sphaerodactylus has led to even more secre-
tive habits and, thus, amplified the problems of tractability.
Because most Sphaerodactylus species are typically
parapatrically distributed, are similar in shape, and published de-
scriptions of their microhabitat are usually general, they have been
assumed to be ecologically similar (but see Thomas et al. 1992).
Still there may be important differences in ecological factors that
have been overlooked. For instance, selection acting on a gecko
that inhabits and defends the underside of a rock should be differ-
ent from those acting on a congeneric inhabiting the adjacent leaf
litter; likely, they would face different predators, hunt different
prey, and experience different social environments, but have been
reported simply as living in the leaf litter.
 The major objective of this study is to describe reproduction
and growth in seven Sphaerodactylus species kept in captivity.
Although data from captive populations may be different from
wild populations (e.g., growth rates/clutch sizes may vary in rela-
tion to proximate climate factors and food availability [Stamps
and Tanaka 1981]), they may be useful for uncovering differences
among related species that could generate testable hypotheses,
suggest field research projects, and help interpret field data.
Sphaerodactylus nicholsi, S. townsendi, and S. roosevelti were
collected from backyards and dumpsites on the southwestern coast
of Puerto Rico (Guanica, Ensenada, Punta Verracos, Punta Pestillo,
Paso Seco, and Salinas) in 1995, 1997, and 1999. Individuals of S.
nigropunctatus granti, S. argus, and S. pimienta were obtained
from eggs collected in eastern Cuba (Holgu?n, Belic, and El Yarey,
respectively). S. savagei adults, from the northeastern of Domini-
can Republic, were obtained from a South Florida dealer in 2001.
The natural history for most of these species is summarized by
Schwartz and Henderson (1991). Nothing is known about the natu-
ral history of S. pimienta.
Lizards were housed individually or in male-female pairs in plas-
tic containers (1.9 L), were kept in the shade indoors, and were
exposed to natural light cycles (through open windows) typical of
South Florida. Containers were provided with a substrate made of
soil and peat moss, a few small rocks, dead leaves and pine bark;
they were misted with water three times a week, and exposed to
daily and seasonal temperature and humidity changes. Mean air
temperature ranged from 24?C to 33?C during April through Oc-
tober and from 14?C to 23?C during November through March,
from 1996 to 2001. Vitamin and mineral-dusted insect larvae and
adults (fruit flies, humpback flies, crickets, and ants) were pro-
vided at two or three day intervals.
Containers were inspected weekly to find newly laid eggs, usu-
ally under leaves, bark, or in the substrate 1?2 cm below the sur-
face. The largest and smallest egg diameters were measured to the
nearest 0.1 mm with an ocular micrometer. Eggs were placed in-
dividually in small plastic containers (30 cc) with a sterile soil/
peat moss substrate and incubated in a plastic box (20 x 20 x 30
cm) in the same room, exposed to daily and seasonal temperature
fluctuations. Eggs were checked every week during the first month
of incubation, and later in the study every day. To explore the po-
tential effect of temperature on incubation duration (Andrews
2004), individual records for each species were pooled into two
groups: eggs incubated during May?October (high temperature)