172
Int. Zoo Yb. (2008) 42: 172–182
DOI:10.1111/j.1748-1090.2007.00029.x
Management and reproduction of the Komodo
dragon Varanus komodoensis Ouwens 1912 at ZSL
London Zoo
G. SUNTER
Herpetology Department, ZSL London Zoo, Regents Park, London NW1 4RY, UK
E-mail: george.sunter@zsl.org
The Komodo dragon Varanus komodoensis is the world’s
largest living lizard. Despite numerous collections ex-
hibiting these animals, only a small number of institu-
tions worldwide have managed to breed them. Here, we
discuss the new, purpose-built breeding facility at ZSL
London Zoo and recent husbandry changes that resulted
in the oviposition of two fertile clutches of eggs and five
hatchling Komodo dragons for the first time in the United
Kingdom. Four of the Komodo dragons were produced
parthenogenetically, which is the first time this has been
documented in this species.
Key-words: husbandry; incubation; Komodo dragon;
neonate management; nutrition; operant conditioning;
reproduction; ultraviolet light.
INTRODUCTION
The Komodo dragon Varanus komodoensis is
the largest living lizard. Found on only a few
islands in the Lesser Sundas, Indonesia, it
has one of the narrowest naturally occurr-
ing ranges of all large terrestrial predators
(Gillespie et al., 2000; Ciofi & Gibson,
unpubl.). Wild populations are considered
stable on Komodo and Rinca; however, se-
vere declines have been recorded on the
island of Flores (Ciofi & De Boer, 2004; Ciofi
& Gibson, unpubl.). Their limited distribu-
tion makes them Vulnerable to stochastic
events, poaching of prey species, genetic
inbreeding and habitat destruction as a result
of slash-and-burn agriculture (Ciofi, 1999;
Ciofi et al., 1999; Sastrawan & Ciofi, 2002;
IUCN, 2006; Ciofi & Gibson, unpubl.; Jessop
et al., unpubl.).
The first officially documented captive
breeding of the species occurred at the
Gembira Loka Zoo in Indonesia in 1965
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THE DEVELOPING ZOO WORLD
(Osman, 1967; Busono, 1974). However, it
was not until 1992 that the first Komodo
dragons to hatch in the Western hemisphere
emerged at the National Zoological Park
(NZP), Washington, DC, USA (Walsh et al.,
1993).
In the past 20 years, considerable informa-
tion has been compiled and published on
captive husbandry and significant improve-
ments have been made in species manage-
ment (see Visser & Belterman, 2002; Walsh
et al., 2004). Komodo dragons have also been
the subject of considerable field research (see
Auffenberg, 1981; Auffenberg & Auffenberg,
2002). However, comparatively little infor-
mation is available on captive-reproduction
events, egg incubation or management of
young dragons. Although this species is cur-
rently held in over 60 institutions worldwide,
only a few have instructions to breed for
genetic-management purposes, and many in-
stitutions house siblings together for exhibit
purposes only (and not for breeding).
Despite recent husbandry advances, the
European captive population is in crisis,
suffering from a comparatively high , death
rate. Since its initiation in 2000, the European
Endangered Species Programme (EEP) popu-
lation has lost eight adult ,,; all deaths,
except one, are allegedly reproduction re-
lated. There was only one ‘programmed’
hatching (of 17) in 2004, none in 2005 and
only one 2006, which is the one documented
in this article. The EEP studbook population
(December 2005) comprised 16.14 Komodo
dragons, in 12 institutions, and of these
c 2007 The Zoological Society of London
ZSL LONDON ZOO: KOMODO DRAGON MANAGEMENT AND REPRODUCTION
14 ,, only six were adults. Four of these
six adults died in 2006, leaving only two
adult ,, and one sub-adult remaining as
breeding stock.
FACILITIES AT ZSL LONDON ZOO
ZSL London Zoo’s Komodo dragon house
was constructed in 2004 and provides out-
standing accommodation for dragons and an
excellent visitor experience. More than 25 m
of continuous viewing windows give unin-
terrupted views of the dragon’s habitat and
information panels and interactive devices
provide educational input (Plate 1).
The footprint of the building is a half-
ellipse, which faces south to capture as much
natural sunlight as possible. Internally, the
building is divided into two display enclo-
sures separated by an artificial-rock divide, a
reclaimed timber-clad visitor area, off-show
holding dens, a service area and a plant room.
The total area available to the Komodo dra-
gons comprises 220 m2 indoor exhibit space,
60 m2 outdoor exhibit space and 45 m2 of
indoor dens.
The air temperature is controlled via a
dedicated heat-and-humidity-exchange sys-
tem that forces warmed and humidified air
into the exhibit. The air-conditioning system
is controlled by independent controllers for,
heating, cooling, humidity, fresh air and
energy recycling (Table 1). The system is
monitored by a TREND computer, which
automatically adjusts the heating and cooling
in the building as required. The hot air enters
the enclosure at ground level in the centre of
Plate 1. The Komodo dragon Varanus komodoensis
facility at ZSL London Zoo. George Sunter, ZSL
London Zoo.
Int. Zoo Yb. (2008) 42: 172–182.
c 2007 The Author. Journal compilation
173
TEMPERATURE RELATIVE
SEASON ( 1C) HUMIDITY (%)
Winter 22–28 45–55
Mid-season 30–33 45–60
Summer 30–37 45–65
Table 1. Daily and seasonal temperature and relative
humidity changes in the Komodo dragon Varanus
komodoensis enclosure at ZSL London Zoo.
the exhibit, which produces a thermal gradi-
ent of 40–25 1C from the centre to the peri-
meter. The heating profile is designed to
replicate the temperatures found within the
Komodo National Park on a daily and seaso-
nal basis.
The exhibit’s internal theme replicates the
dry riverbeds typically found in lowland
valleys in the Komodo National Park. Artifi-
cial rainfall is controlled via an external pump
supplying pre-heated water at 17 1C and an
electronic timer controls the rainfall that is
adjusted to mimic seasonal rainfall patterns.
The exhibit areas are filled with 30 cm of a
porous sand substrate with a perforated con-
crete base below to ensure adequate drainage.
Ponds in each half of the main exhibit are
filled with water maintained at 22 1C, provid-
ing both drinking and bathing opportunities
for the dragons and planted areas provide
shaded patches and offer refuge from the heat
of the day.
All three dens have independently con-
trolled temperatures via stand-alone air-con-
ditioning units. Two 250 W metal–halide
flood lamps (5600 K) on an automated
12:12 hour (light:dark) photoperiod provide
additional illumination. Each den also con-
tains three Osram Ultravitaluxt bulbs posi-
tioned 50 cm above designated basking areas.
The substrate in the dens is a deep layer of
bark chippings covered with leaf litter.
Because of the northerly latitude of Lon-
don, the Komodo dragons are only permitted
outdoor access during the summer months.
Two 3 kW infrared radiant heaters provide
localized hot spots of 45 1C. Between April
and October, the dragons are given access to
their outdoor enclosure when the air tempera-
ture exceeds 22 1C.
c 2007 The Zoological Society of London
174
Komodo dragons require exposure to ultra-
violet light (UVB) between the ranges of 285
and 315 nm to allow for vitamin D3 photo-
biogenesis, calcium absorption and homeos-
tasis (Gyimesi & Burns, 2002). At the time of
writing, it is still unclear how much UVB a
Komodo dragon requires. However, work
conducted by Gillespie et al. (2000) demon-
strated that Komodo dragons that were
housed indoors with no UVB source exhib-
ited clinical lameness, poor bone density and
fractures along several long bones. Further-
more, circulating vitamin D3 levels as low as
14 nmol litre 1 were recorded and these in-
creased to 252 nmol litre 1 with exposure to
1 month of temperate sunlight.
The majority of the Komodo dragon ex-
hibit roof at ZSL London Zoo is constructed
from transparent ETFE (ethylene tetrafluor-
oethylene) plastic that is penetrable to UVB,
with as much as 85% transmission of UVB in
the range of 285–315 nm. Because the United
Kingdom is at a higher latitude from the
equator than the Komodo National Park, the
amount of UVB available at ground level is
lower throughout the year and especially
during the winter months (October–March).
To compensate for this, additional artificial
UVB-emitting lamps are located throughout
the exhibit and in all dens.
Each enclosure has two clusters of spot
lamps. Each cluster contains three Osram
Ultravitaluxt 300 W UVB-producing bulbs,
which generate localized hot spots up to
45 1C and emit a minimum of 250 mW cm 2
of UVB radiation. Ambient and spot lamp
UVB levels are monitored inside and outside
the enclosure throughout the year using a
standard protocol (time, location and orienta-
tion) with a Solar Systems Solar Metert 6.2.
EEP and Association of Zoos and Aqua-
riums (AZA) recommendations suggest that
each Komodo dragon should have access to
an area where available levels of UVB radia-
tion are in excess of 250 mW cm 2, as a
minimum requirement to avoid health pro-
blems and to promote natural growth.
The remainder of the roof, which covers
the viewing area and needs to be darker than
the exhibit space, is a living ‘brown roof’.
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THE DEVELOPING ZOO WORLD
KOMODO DRAGONS AT ZSL
LONDON ZOO
In June 2004, ZSL London Zoo received an
F1 < Komodo dragon ‘Raja’ hatched in 1998,
from Miami Metro Zoo, FL, USA. Shortly
afterwards, an F1 , ‘Nina’, hatched in 1994,
was received from Reptilandia in the
Canary Islands. Nina died shortly after com-
pleting her quarantine period from severe
haemorrhage into the coelomic cavity as
a result of ruptured mineralized vasculature.
This medical condition, diagnosed postmor-
tem for the first time at ZSL London
Zoo, appears to be a recurring issue in
Komodo dragons in captivity (R. Pizzi, un-
publ. data).
Almost a year later, a second F1 , ‘Sungai’
arrived on loan from Thoiry Zoo in France.
Sungai hatched in 1995 and had laid small
clutches of infertile eggs during the 2 years
before her arrival at ZSL London Zoo.
DAILY MANAGEMENT
Each adult animal occupies one-half of the
exhibit separated by the artificial-rock wall.
One of the intentions of the design was to
allow both adults to experience the same
environmental conditions and share olfactory
contact but prevent visual and physical con-
tact until the breeding season.
The Komodo dragons are housed in their
respective dens during the night and released
into the exhibit in the morning. Each day, the
dragons are released from their dens and
allowed unrestricted access to one of the
paddocks for the rest of the day with the
den remaining closed. ZSL London Zoo
operates a non-protected contact regime and
regular keeper interaction with the dragons is
encouraged.
Variations in the timing of key events
such as feeding, keeper interaction, den ac-
cess and rainfall activations are purposely
performed at random times to prevent stag-
nation.
Each dragon returns to a den in the eve-
ning, which has a lower environmental tem-
perature to mimic night-time conditions.
c 2007 The Zoological Society of London
ZSL LONDON ZOO: KOMODO DRAGON MANAGEMENT AND REPRODUCTION 175

OPERANT CONDITIONING Rattus norvegicus, Mice Mus musculus, small

Rabbits Oryctolagus cuniculus, Chicken
It has long been recognized that Komodo
pieces Gallus gallus and chunks of Cow
dragons are behaviourally complex animals
(beef) Bos taurus or Horse meat Equus
capable of learning. See Burghardt et al.
caballus (Gibson, 2006). Considerable evi-
(2002) and Murphy & Walsh (2006) for a
dence from field observations on prey-species
review of behavioural repertoire in captivity.
composition, feeding frequency and seasonal-
Komodo dragons therefore represent ideal
ity (see Auffenberg, 1981) appears to conflict
candidates for training and respond quickly to
with the nutritional guidelines for captive
standardized techniques.
Komodo dragons. In 2005, ZSL London
At ZSL London Zoo, we trained our
Zoo, in conjunction with the EEP, adopted an
Komodo dragons to accomplish five impor-
approach to nutrition designed to mimic field
tant goals.
data.
 To move from one area to another, either
At the time of writing, ZSL London Zoo’s
from den to exhibit or vice versa.
adult Komodo dragons are offered much
 To be box-trained for restraint and phy-
larger individual meals infrequently. This
sical examination by the veterinary de-
is calculated from a budget of 150–
partment.
160 kg year 1, based on observations of the
 To establish a trusting/working relation-
annual intake of a 48 kg wild adult (Auffen-
ship with the keepers.
berg, 1981). Large portions of food, up to
 To allow the animal to differentiate
20% of the animal’s body mass, are offered in
feeding cues from general keeper inter-
one or two feeding events per month.
action and avoid ‘door rushing’ pro-
Correspondingly, the composition of food
blems experienced by other institutions.
items was also adjusted to incorporate larger-
 To enrich the lives of behaviourally
bodied animals. We now offer large portions
complex and intelligent reptiles in cap-
(43 kg) of Muntjac deer Muntiacus muntjak,
tivity.
Goat Capra sp and Wild rabbit O. cuniculus,
To achieve the above objectives, a number
which are all organically sourced. Feeding
of common training techniques were used.
observations and faecal analysis of wild
These include target training and the use of a
Komodo dragons revealed that a greater
‘clicker’ to act as a bridge stimulus. Small
proportion of bone, skin and hair were in-
food items, such as mice (20 g), are used as
gested by adult dragons than most other apex
nutritional rewards and Komodo dragons
predators (Auffenberg, 1981). In accordance,
learn very quickly to recognize the target and
we give our adults portions of carcass rich in
associated ‘click’ for good behaviour. ZSL
bone, fur and cartilage. Commercially pro-
London Zoo’s Komodo dragons are target
duced whole rats, mice and chicks are only
trained on a daily basis and the original goals
given in small amounts during target training
have been extended to include press-related
sessions and constitute a small portion
events, keeper talks, radiography, venipunc-
(o20%) of the annual nutrition budget.
ture and transdermal ultrasonography.
The use of vitamin/mineral supplementa-
tion in dragon diets is not uniform and ZSL
NUTRITION
London Zoo currently uses no supplementa-
Most recommendations for the nutrition of tion. Routine blood samples taken from our
Komodo dragons in captivity are based on the Komodo dragons reveal blood-chemistry
NZP approach of ‘little and often’ (Walsh profiles within normal parameters when com-
et al., 2002). pared with the levels published by Gillespie
Generally, adult dragons of 47 years are et al. (1997, 2000), Gyimesi & Burns (2002)
offered 1.5–3 kg of food per week in two to and Spelman (2002).
three separate feeding events (Walsh et al., Wild Komodo dragons show typical ‘gorge
2002, 2004). The food items consisted of Rats and fast’ feeding behaviour (Auffenberg,

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c 2007 The Zoological Society of London
176
1981; Murphy et al., 2002). The impact of a
regular diet in captivity has long been ques-
tioned (see Walsh et al., 2002). Studies by
Walsh et al. (1999) and Wikramanayake et al.
(1999) on the thermal biology of wild and
captive dragons demonstrated that captive
Komodo dragons have a significantly lower
body temperature than their wild counter-
parts. The authors theorized that small, fre-
quent feeds may be one factor that reduces
the internal body temperature in Komodo
dragons in captivity.
Data collected at ZSL London Zoo using
Thermocront data loggers ingested by our
dragons support the Walsh et al. (1999)
hypothesis. When frequent small prey items
(200 g) were regularly ingested in 2005, the
core body temperature (CBT) peaked at a
maximum 33 1C. However, after the inges-
tion of larger, infrequent meals (  2 kg), after
2005, the CBT of our < dragon increased to
peaks of 37–38 1C, independent of ambient
air temperature. Gastric transit times also
increased, with large meals taking signifi-
cantly longer to digest.
Walsh et al. (2002) state that Komodo
dragons are capable of physiologically up-
grading their metabolic rate during digestion
in a manner similar to White-throated moni-
tors Varanus albigularis (Secor & Philips,
1997). Similarly, Anderson et al. (2005)
demonstrated that the Burmese python
Python molorus is capable of accelerating
postprandial metabolic rate by 40% relative
to fasting rate. This may have further implica-
tions for Komodo dragons, as the increased
basking time needed to generate a higher
CBT may affect photo-biogenesis of circulat-
ing 25(OH)D3. Gyimesi & Burns (2002) and
Nijboer et al. (2003) specifically state that
UVB has a direct effect on the circulating
levels of 25(OH)D3. Further studies are re-
quired to clarify the relationship among
meal size, CBT, metabolism and circulating
vitamin D3.
MATING AND REPRODUCTION
Varanid reproduction has been studied in
considerable detail; see Horn & Visser
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THE DEVELOPING ZOO WORLD
(1997) for a good overview. However,
Komodo dragon reproduction in captivity
has been studied in less detail. Only a handful
of institutions, Cincinnati Zoo, Miami
Metro Zoo, Honolulu Zoo, NZP, Toronto
Zoo, Reptilandia and ZSL London Zoo, have
achieved reproduction outside of Indonesia.
In October 2005, we began to integrate our
two Komodo dragons for breeding. Both had
been acclimatized to captive conditions at
ZSL London Zoo for over 6 months and both
had experienced exactly the same environ-
mental cues and nutritional regime before
introduction. Before any physical introduc-
tion, efforts were made to familiarize each
animal with its potential mate to reduce
aggression. Opaque barriers were removed
to give the dragons visual access to each
other. Shed scales and faeces were swapped
between enclosures and tactile contact was
permitted through vertical bars. At each inter-
action, the , behaved defensively, while the
< showed threat displays. During this safe-
contact period, the posturing and defensive
behaviour reduced in intensity and frequency
with increasing exposure.
On 8 November 2005, the animals were
allowed full physical contact. Initially, the
< was given access to the ,’s side of the
exhibit. The , was the first to show aggres-
sion. The < remained passive throughout and
folded his limbs to his body and laid flat
against the ground. On a couple of occasions,
the < managed to suppress the , but remained
dominant for only a couple of minutes. After
30 minutes, the < suddenly showed aggres-
sion and asserted his dominance by biting the
, on the neck. At this point, the animals were
separated.
After a 7 day ‘cooling-off’ period, the
, was led into the <’s side of the exhibit for
the first time. A series of ritualized bipedal
bouts ensued, which terminated with the
< asserting his dominance, having won at
least 90% of the wrestling bouts. This process
was repeated every day for the next 6 days.
Each time the animals were introduced, the
aggression levels reduced until eventually
the , became passive in the <’s presence
(Plate 2).
c 2007 The Zoological Society of London
ZSL LONDON ZOO: KOMODO DRAGON MANAGEMENT AND REPRODUCTION
Characteristically, the < would lay along
side the ,, one foreleg would be placed over
her back and the corresponding hind leg was
scraped repeatedly across her pelvis and tail.
The <’s tongue flicked continually over the
neck and head of the , and he jerked his head
horizontally to stimulate her. Voluntary tail
lifting by the , was witnessed on several
occasions. Hemipenal eversion and intromis-
sion were observed twice and seminal fluid
was clearly emitted. Intromission was brief,
c. 2–5 minutes, and both animals remained
still. After 3 weeks of courtship, the , began
to refuse food and showed signs of abdominal
swelling.
Oviposition
Before oviposition, the , lost all appetite for
food and became unresponsive and listless.
Regular blood samples were taken ventrally
from the tail to monitor calcium serum levels
and transdermal ultrasounds were performed.
After several days in this condition, it was
possible to detect ova that were c. 6 cm in
diameter, although calcium serum levels
appeared to be stable and within normal
parameters (Gillespie et al., 1997, 2000;
Spelman, 2002). After 10 days of immobility,
the ,’s activity levels increased and she
resumed periodic feeding and began excavat-
Plate 2. Raja (<) and Sungai (,) Komodo dragons
Varanus komodoensis during courtship. Claire
Cunningham, ZSL London Zoo.
Int. Zoo Yb. (2008) 42: 172–182.
c 2007 The Author. Journal compilation
177
ing potential nest sites. Initially, she dug
exploratory holes and then abandoned them.
Auffenberg (1981) also documented this be-
haviour in wild Komodo dragons and it is
believed to be anti-predatory in nature.
A nesting site was provided in one of the
off-show dens. This comprised a large mound
of earth, sand and leaf litter 250 cm 
200 cm  80 cm deep. A false ceiling was
incorporated to prevent tunnel collapse
following Walsh et al. (2002). The nesting
den was heated to an ambient 28 1C with
localized basking spots provided for the ,.
The temperatures inside the oviposition site
were 27–29 1C.
Two separate clutches of eggs were laid by
the ,: clutch 1 (22 eggs) was laid on 5 August
2005 before introduction to our < and clutch
2 (six eggs) was laid on 6 January 2006
following successful mating.
Incubation
All eggs were artificially incubated and were
set at a ratio of 1:3 (water:vermiculite) by
mass in containers with airtight lids. Initially,
all eggs were incubated together and infertile
eggs were removed as the incubation pro-
gressed. Eventually, eggs were incubated in
pairs within a larger box. The lids were
initially removed weekly for aeration of the
incubation environment and daily in the latter
stages of development. Following NZP re-
commendations, the eggs were incubated at
29 1C for the first 7 days. The temperature
was then increased to 29.5 1C for the next
180 days before being reduced to 28.5 1C.
Miami Metro Zoo also adopted this strategy
in 1998 when 27 of 29 eggs hatched
(S. Conners, pers. comm.).
After 110 days of incubation, the eggs
began to collapse and decline in mass despite
the relative humidity being 485%. The
incubation boxes were then placed inside a
larger box containing shallow water to in-
crease the humidity. Water was dripped onto
the eggs daily for the next 8 weeks and the
eggs began to increase in mass (Table 2).
The progress of the eggs was followed
using traditional candling techniques. We
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178 THE DEVELOPING ZOO WORLD

EGG MASS (g)

2005 2006
EGG NO. 6 AUG 10 NOV 21 NOV 13 DEC 24 DEC 18 JAN 9 FEB 1 MAR 15 MAR 22 MAR
1 2
2 154 145 152 159 164 171 166 152 146 140
5 179 1661 166 172 181 1862 180 174 172 164
9 142 1291 139 147 154 1562 153 144 139 134
10 170 1561 158 166 170 1732 168 161 157 151
1
First date when water was dripped onto eggs.
2
When additional water was stopped.
Table 2. Mass change of viable Komodo dragon Varanus komodoensis eggs from clutch 1 (laid 5 August 2005)
during artificial incubation at ZSL London Zoo.

also used a ‘Buddy’ digital egg monitors,

which uses a single or triple light-emitting
diode light source emitting infrared light and
a light sensor that detects the variation in
intensity of absorbed versus reflected infrared
light caused by arterial pulsatile vasculature
within the developing fertile egg membranes
(Strike et al., 2006, 2007). The eggs were
monitored three times weekly, with the high-
est heart rate of 62 beats per minute (b.p.m.).
At around day 217 of incubation, the heart
rates reduced to below 30 b.p.m. Forty-eight
to 72 hours before hatching, heart rate would
characteristically increase to 60 b.p.m. until
hatching (M. Taema & T. Strike, pers.
comm.) This is the first time this technology
has been applied to monitor the eggs of this
species and is part of an ongoing study.
The first Komodo dragon began to hatch Plate 3. A Komodo dragon Varanus komodoensis
on 24 March 2006, followed by three hatching at ZSL London Zoo. George Sunter, ZSL
others, each c. 2 days apart (Plate 3). The London Zoo.
incubation period was 251–256 days. Incuba-
tion periods recorded in other institutions were housed together in a 1 m3 plastic vivar-
are in the range of 205–256 days, with an ium with a wiremesh ceiling. The single
average of 220 days (Walsh et al., 2002). The animal that hatched from clutch 2 was housed
first four Komodo dragons to hatch had a separately. One Osram Ultravitaluxt bulb
body mass of 96–121 g. Walsh et al. (2002) was suspended over the rearing vivarium to
found a mean body mass at hatch of 97 g, provide a basking spot of 35 1C. The ambient
whereas Auffenberg (1981) gave a lower temperature was held at 25–28 1C. Branches
figure of 80.3 g based on Busono (1974) and cork-bark tubes were used as cage dec-
(Tables 3 and 4). oration and provided ample refuge and
climbing opportunities. Regular spraying
Neonate management maintained humidity above 65%.
The Komodo dragons took their first feed 5–
All hatchling Komodo dragons were retained 7 days after being removed from the incubator
inside the incubator for up to 7 days post- and were given pre-killed rat Rattus sp pups.
emergence to reduce stress and allow yolk After their initial meals, the dragons were
absorption. All four dragons from clutch 1 offered c. 25 g each per week in three to five

Int. Zoo Yb. (2008) 42: 172–182.

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ZSL LONDON ZOO: KOMODO DRAGON MANAGEMENT AND REPRODUCTION 179

EGG NEONATE
-------------
INCUBATION
EGG NO. LENGTH (mm) WIDTH (mm) MASS (g) FERTILE (Y/N) PERIOD (DAYS) BODY MASS (g)

1 89
6 59
1 169
2 ? 7
2 86 58
6 154
4 Y 251 109
8
3 88
3 61
9 169
2 ? 7
4 83
7 55
7 146
6 N
5 94
3 57
5 179 Y 255 121
1
6 95
3 53
7 184 ? 7
7 83
8 55
3 159 Y 64
8 78
2 57 143 ? 7
9 84
6 55
4 142 Y 253 96
6
10 85
3 59
5 170
8 Y 256 108
4
11 78
8 49
7 92 N
12 89
5 45
8 96 N
13 83
6 51
6 107 N
14 83
7 56
5 110
6 N
15 84 52
5 113
4 N
16 72
1 51
5 103 N
17 84 50
2 84
4 N
18 88
2 45
7 100 N
19 83
2 44
1 102
4 N
20 85
6 48
9 92
8 N
21 78
2 45 94
4 N
22 83
5 54
7 91
4 N

Table 3. Clutch 1 data, including clutch size, egg dimensions, incubation period (to point of death or hatch) and
body mass at hatch, for parthenogenetically produced Komodo dragons Varanus komodoensis at ZSL London
Zoo.

EGG NEONATE
-------------
INCUBATION
EGG NO. LENGTH (mm) WIDTH (mm) MASS (g) FERTILE (Y/N) PERIOD (DAYS) BODY MASS (g)

1 88 57
1 145
2 ? 47
2 95
7 62
3 185
7 N
3 89
3 60
7 171 N
4 83
1 52 125 Y 224 73
5 84 54 109 N
6 103 53 152 N

Table 4. Clutch 2 (laid 6 January 2006) data, including clutch size, egg dimensions and incubation period, and
body mass at hatch for the Komodo dragon Varanus komodoensis produced sexually at ZSL London Zoo.

small feeds, comprising one appropriate-size
furred mouse M. musculus, one part-incubated
quail egg and multiple gut loaded insects
(Gryllus sp, Blaberus sp and Shistocerca sp).
The mice and eggs were always taken with-
out problem but the Komodo dragons
required significant encouragement to recog-
nize insects as food. Initially, the insects
were pre-killed and wrapped in mouse fur or
scented with blood before they were accepted
(Fig. 1).
Health checks were carried out after the
first 3 months of growth and blood samples
were taken. Radiography was carried out to
check bone density, and transdermal ultra-
sound performed to check internal organs.
Samples were also sent off for DNA-sexing,
as described by Halverson & Spelman
(2002). The results confirmed that the four
animals from the first clutch were all << and
had been produced parthenogenetically; see
Watts et al. (2006) for a full description.

Int. Zoo Yb. (2008) 42: 172–182.

c 2007 The Author. Journal compilation

c 2007 The Zoological Society of London
180
1200
1000
body mass (g)
800
600
400
200
0
0 7 13
Fig. 1. Body-mass growth of four < Komodo dragon Varanus komodoensis at ZSL London Zoo.
Parthenogenesis is defined as the production
of offspring without fertilization by a <. The
offspring’s genotype is derived entirely from
its mother (Groot et al., 2003). In Varanus sp,
,, have dissimilar chromosomes (‘Z’ and
‘W’) whereas the recombination ZZ produces
<<. Because the parthenogenetic mechanism
can only produce homozygous individuals,
all resulting offspring are <. A later, second
clutch laid by a different , at Chester zoo also
produced parthenogenetic offspring despite
being raised at a different institution under
different conditions.
The second clutch from ZSL London
Zoo’s Komodo dragon produced a single ,
that was the result of sexual reproduction.
Facultative parthenogenetic reproduction was
previously unknown in this species and sug-
gests that sexual reproduction is not a fixed
reproductive trait for this species (Watts et al.,
2006).
As the juveniles grew, they were moved
into a larger enclosure. The four << contin-
ued to be housed together and the single ,
was housed separately but still in visual and
olfactory contact. The four << share a 15 m2
den furnished with a deep substrate of
bark chips and leaf litter, ample climbing
branches, live plants and multiple Osram
Ultravitaluxt basking lamps.
The young Komodo dragons appear to
cohabit well and fighting is rarely observed.
The younger , is separated in a wire mesh
enclosure, which affords her protection from
her much larger half-brothers. As the four <<
grew, it quickly became evident that a dom-
inance hierarchy was being established and
each animal had its place in the pecking order.
Int. Zoo Yb. (2008) 42: 172–182.
c 2007 The Author. Journal compilation
THE DEVELOPING ZOO WORLD
4598
4599
4600
4601
60 90 120 180 300
days
Ritualized threat displays were often seen
during feeding and submissive displays ob-
served from the lower-ranking animals. Ef-
forts were made to ensure that each animal
received equal amounts of food and periodic
shifting of troublesome individuals to a sepa-
rate area helped maintain a balance in the
group.
Studies are currently underway to assess
and monitor the long-term health status of the
four < parthenogenetic dragons and better
understand the mechanism underpinning fa-
cultative parthenogenesis in this species.
CONCLUSION
The facility at ZSL London Zoo combines an
attractive exhibit with a complex environ-
ment that provides optimal environmental
conditions, including adequate levels of
UVB, for Komodo dragons. The suitability
of the conditions and husbandry regime are
confirmed by the first hatching of Komodo
dragons in the United Kingdom. Further-
more, significant improvements to the nutri-
tion and management of the species at ZSL
London Zoo have been adopted in the Eur-
opean Association of Zoos and Aquaria
(EAZA) EEP guidelines for the species.
The discovery that Komodo dragons are
capable of facultative parthenogenesis is an
unexpected breakthrough with implications
far beyond the scope of this paper.
ZSL has made a major commitment to
Komodo dragon conservation through the
development of a large, dedicated breeding
facility and supporting field research and
conservation initiatives on the islands of
c 2007 The Zoological Society of London
ZSL LONDON ZOO: KOMODO DRAGON MANAGEMENT AND REPRODUCTION
Komodo, Gili Motang and Flores. The pro-
gramme on the latter is the only long-term
conservation initiative for the species outside
the Komodo National Park and is funded by
donations from all EAZA EEP participating
institutions.
ACKNOWLEDGEMENTS
I am grateful to the ZSL herpetology team, including
Richard Gibson, Terry March, Matt Fagg, Glynn Hennes-
sey and Ian Stephen, and the ZSL veterinary team,
including Wayne Boardman, Taina Strike, Matt Rendle,
Maged Teama and Romain Pizzi. Thanks are also owing
to Steve Conners and Tom Condie (Miami Metro Zoo,
FL, USA), Gerard Visser (Rotterdam Zoo, the Nether-
lands), Colomba de la Panouse (Thoiry Zoo, France) and
Trooper Walsh (Private consultant). Matt Rendle and
Richard Gibson commented on and edited the manuscript.
PRODUCTS MENTIONED IN THE TEXT
‘Buddy’s: digital egg monitor, manufactured by Avi-
tronics, PO Box 107, Truro, Cornwall TR1 2YR, UK
(http://www.avitronics.co.uk); supplied by Vetronic Ser-
vices, 4 Brunel Buildings, Brunel Road, Newton Abbot,
Devon TQ12 4PB, UK (http://vetronic.co.uk).
ETFE: UV stable copolymer Ethylene Tetra Fluoro
Ethylene, manufactured by Vector Foiltec UK, Studio 4,
The Cooperage, 91 Brick Lane, London E1 6QL, UK.
http://www.vector-foiltec.com
Metal–halide flood lamps: 250 W lamps, manufactured
by Lyco Direct limited, Clarke Road, Bletchley, Milton
Keynes MK1 1ZR, UK. http://www.lyco.co.uk/
Osram Ultravitaluxt: 300 W light bulbs, manufactured
by Osram, German Headquarters, Hellabrunner Strasse 1,
81543 Munchen, Germany.
Solar Systems Solar Metert 6.2: digital UV meter,
manufactured by Solartech Inc., 26101 Harbour Pointe
Drive N, Harrison Township, MI 48045, USA. http://
www.solarmeter.com
Thermocront: temperature logger, manufactured by
OnSolution, PO Box 1007, Baulkham Hills, 2153 Aus-
tralia. http://www.onsolution.com.au/
TREND: computer software, manufactured by Keypad
Control Systems Ltd, 35 Salcombe Drive, Earley, Read-
ing RG6 7HU, UK.
Vent-Axia Patio Heaters: 1.3 kW quartz patio heaters
(IP54), manufactured by Vent-Axia, Fleming Way, Craw-
ley, West Sussex RH10 9YX, UK. http://www.vent-
axia.com/
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c 2007 The Zoological Society of London