Beruflich Dokumente
Kultur Dokumente
2006
EKOLOGI SPATIAL TETASAN DAN ANAKAN BIAWAK KOMODO
(Varanus komodoensis) DI TAMAN NEGARA KOMODO, INDONESIA
2006
iii
DECLARATION
I hereby declare that the work in this thesis is my own except for quotations and
summaries, which have been duly acknowledged
ACKNOWLEDGEMENTS
I am grateful upon The Almighty and Merciful Allah S.W.T and his Prophet
Muhammad S.A.W that by His will has bestowed me a good opportunity to explore
one of the tinniest parts of nature’s secret. This thesis has benefited from the help of
many people during all the stages of its maturation. Coming at the end of this work, I
wish to list here all the contributions that made this work possible, by order of
apparition;
Special thanks to Prof. Dr. Zubaid Akbar Mukhtar Ahmad for thesis
supervision and every scientific discussion. I particularly thank Dr. Tim Jessop, my
co-supervisor his assistance in providing research funds and equipments, more over
in directing my field research and scientific writing progress during it. Without their
help, this work had probably not successfully merged.
Avery special thanks also should be given to my parents: Abdul Azid Muchtar
and Aas Sadiah; and my younger brothers and sister: Oris Junisa, Atus Satriawan,
Widia Putri Julika. Last, but not least, a special grateful and dedication should be
awarded to the most amazing woman in my life, my beloved wife, Eva Fairus
Balahmar. Their patience, passionate, and encouragements to kept lifting up my
performance and through the hard time during the research and scientific writing.
This work will not be done very well without their support.
ABSTRACT
ABSTRAK
Tujuan daripada kajian ini ialah untuk menyelidiki pola-pola dalam ekologi spasial
pada tetasan dan anak Biawak Komodo (Varanus komodoensis) di Taman Negara
Komodo, Indonesia. Di dalam penyelidikan ini, kaedah Capture-Mark-Release and
Recapture telah dijalankan di 10 lembah utama ke atas 4 pulau di TNK daripada 2003
sehingga 2005, manakala ‘radiotracking’ dijalankan di Loh Liang, pulau Komodo,
antara bulan Mac sehingga Julai 2004 dan 2005. Didapati bahawa jarak perpindahan
tetasan biawak Komodo lebih rendah (32.62 ± 12.67 m/hari) dari pada anakan
(129.14 ± 41.71 m/hari; t-test t1,9=-3.014, p=0.015). Tetasan memiliki luasan
Minimum Convex Polygon yang secara nyata lebih kecil (3.02 ± 0.73 ha) dari pada
anakan (24.31 ± 8.38 ha; t-test; t1,9 = -3.658, p = 0.006). Terdapat 5 dari pada 6 kes
overlap wilayah aktiviti (4.84 – 91.01 %). Tetasan bersifat arboreal (97.70%) dari
pada anakan yang cenderung terestrial (71.0%) (Chi square test; x2 = 239.22, p ≤
0.001). Tetasan lebih banyak teramati pada ketinggian yang rendah (> 25 m) dari
pada anakan yang lebih banyak teramatai pada ketinggian (25 – 50 m; Chi square
test; x2 = 25.127, p ≤ 0.001). Tetasan menunjukkan kecenderungan untuk
menggunakan pohon asam Tamarindus indica tree (45.37 %) sebagai tempat
berteduh, sedangkan anakan lebih memilih celah-celah diantara batu karang (51.77
%). Terdapat pula korelas yang nyata antara jarak perpindahan skala besar dengan
luasan lembah (Pearson correlation test; r = -0.125, p ≤ 0.001) dan dengan tingkat
kemiringan tebing (Spearman correlation test; r = -0.399, p = 0.007), akan tetapi
terdapat korelasi yang tidak nyata dengan jenis vegetasi (Spearman correlation test; r
= -0.399, p = 0.007). Daripada kajian ini menunjukkan bahawa sedikitnya
perpindahan antara lembah mahupun antara pulau pada tetasan dan anakan biawak
Komodo dapat berdampak kepada rendahnya pertukaran variasi gen pada populasi
pulau. Pengurus mesti merancang strategi pengurusan tetasan dan anakan Biawak
Komodo untuk menyokong pertukaran sumber variasi gen bagi populasi terisolasi.
vii
CONTENTS
Page
DECLARATION iii
ACKNOWLEDGEMENTS iv
ABSTRACT v
ABSTRAK vi
CONTENTS vii
LIST OF FIGURES x
LIST OF TABLES xi
4.1 Introduction 28
4.3 Results 35
4.4 Discussion 44
5.1 Introduction 49
5.3 Results 56
5.4 Discussion 57
ix
6.2 Recommendations 63
REFERENCES 68
x
LIST OF FIGURES
3.1 Study sites across the four islands of Komodo National Park.
Numbers are dedicated to show the locations; 1. Loh Sebita;
2. Loh Liang; 3. Loh Lawi; 4. Loh Wau; 5. Loh Buaya; 6.
Loh Baru; 7. Loh Tongker; 8. Loh Dasami; 9. Nusa Kode;
10. Gili Motang. 19
LIST OF TABLES
Table Page
3.1 Size of each island within KNP and it’s vegetation cover (in
km2). 15
GENERAL INTRODUCTION
The Komodo dragon (Varanus komodoensis) is the largest extant lizard in the world
(King & Green 1999; Mattison 1992; Pough et al. 2001). This species lives in a
relatively small range of distribution in the wild. Komodo dragons can be found on the
islands of Komodo, Rinca, Nusa Kode, Gili Motang, and the western coast of Flores
(Ciofi et al. 2002; Ciofi & de Boer 2004), and were previously known on Padar Island.
Currently, four of these populations are located within the boundary of Komodo
National Park (PHKA 2000). Flores is the largest island within the Komodo dragon’s
distribution and contains two nature reserves: Wae Wuul on the west coast and Wolo
Tado on the north coast (Sastrawan & Ciofi 2002). It is evident that the dragon
populations in these parks are more secure than those outside of the parks, given that
key threatening processes, including habitat loss, altered fire regimes and competition
for key prey with humans, are largely absent within the park (Jessop et al. 2004). The
range of the Komodo dragon has decreased significantly over the last 3 decades due to
several threatening processes including the suspected decline of large prey, such as
Timor deer (Cervus timorensis) and anthropic habitat fragmentation and disturbances
(Ciofi et al. 2002; Ciofi & de Boer 2004). Degradation of the environment is
considered to be a major threatening process that could influence the viability of the
extant dragon populations (Jessop et al. 2004).
Due to its vulnerability and its restricted distribution in the wild, Varanus
komodoensis is listed as “vulnerable” in the IUCN Red Data Book, and is on
Appendix I of CITES (Ciofi et al. 2002). The Komodo dragon is also protected by
2
Indonesian law, including the need for a presidential decree to transfer any
individuals of this charismatic species out of Indonesia.
There are a number of key research areas that require further investigation to
facilitate Komodo dragon management and conservation. An important component of
Komodo dragon management is to understand the spatial requirements and habits,
including movement, activity area, behavior, and habitat use (Phillips 1995; Brill et
al. 1995). These spatial aspects of an animal’s ecology are the key to understanding
important processes such as dispersal, habitat partitioning and reproduction and how
they influence population structure. Understanding these spatial aspects of dragon
ecology is also invaluable in the conservation of this threatened species.
A number of spatial studies have been conducted and have described activity,
area size, movement, and behaviour of Varanid lizards (Ibrahim 2002; Thompson et
al. 1999; Thompson 1992) is relation to patterns of physiology, reproduction and
demography (Christian et al. 1995; Christian & Weavers 1994; James 1996; Phillips
1995). These spatial studies focused on adults, and thus little is known concerning
juveniles and hatchlings. Spatial studies on Komodo dragons by Auffenberg (1981)
and Sastrawan and Ciofi (2002) focused on the home ranges of adult dragons only.
3
In the present study, spatial ecology was defined as the interaction between
young Komodo dragons with environmental conditions. It is hypothesized that
environmental conditions will influence the spatial ecology of young Komodo
dragons. The objectives of this study are to: 1) determine the patterns of daily
movement and the activity areas; 2) identify patterns of habitat use; and 3) assess
long distance movement ability in hatchling and juvenile Komodo dragons. Results of
this research are anticipated to contribute valuable information on the biology of
hatchling and juvenile Komodo Monitors, that can be of benefit to the management
practices of the park, particularly for this species and its habitat.
CHAPTER II
LITERATURE REVIEW
Habitat loss and fragmentation are currently the most serious threats to wildlife
worldwide, so it is important to understand how patterns and processes of landscape
change will cause individual populations and species to respond to these broad-scale
modifications (Blumstein & Fernandez-Juricic 2004; Collinge 2001). Spatial
heterogeneity processes directly affect ecological systems (Gardner et al. 1989). The
spatial arrangement of individuals within a population will reflect aspects of its
behavior and ecology, and is important in determining population persistence and
gene flow within and between sub-populations (Brown & Downhower 1988; Johnson
2000). Thus, the dynamics of an animal population depends not only on birth and
death rates, but also on an animal’s ability to move into or out of a population
(Dasmann 1964). Determining the number of individuals persisting in an area is a
basic question in ecology, but it is more important to understand how an animal
responds to changing landscape conditions, regardless of whether it is at the
individual, population or community level (Lawson et al. 2006).
Krebs (1999) pointed out that spatial ecology as a science aims to understand
the ecological processes that determine the location of individuals, which are rarely
spread evenly over the landscape. Collinge (2001) concluded that spatial ecology is
an ecological study, which centers upon understanding how landscape configurations
influence the community and population dynamics of organisms. Spatial ecology is at
the very core of the science of ecology (Boyce & McDonald 1999). Whitaker and
Shine (2003) stated that studying spatial ecology can contribute at least three potential
5
The Komodo dragon, Varanus komodoensis, was described for the first time by
Major Peter A. Ouwen in 1912 (Auffenberg & Auffenberg 2002; Dunn 1927). This
6
giant lizard species was placed in the genus Varanus, family Varanidae, order
Squamata, Class Reptiles (Mattison 1992). Varanus salvadorii from Southern New
Guinea and V. varius from Southeastern and Eastern Australia are believed to be the
sister groups of V. komodoensis (King et al. 2002; Molnar 2004). The closest
congeneric species occupying the same region is the Monitor lizard, V. salvator
salvator (Auffenberg 1981).
Auffenberg (1981) reported that this species was called the “Ora” by the local
people of Komodo, Rinca, and West Manggarai. There are several local names
described by Auffenberg (1981) from across its distribution in the Lesser Sunda
region (see Table 2.1). The name “Komodo” was taken from the name of the island
where the first specimens were taken, and which means “rats” (Dunn 1928).
Even though the Komodo dragon is the largest lizard in the world, this species
has the smallest range of any large carnivore (King & Green 1999; Mattison 1992;
Pough et al. 2001). In the early studies of the Komodo dragon, this species was found
in the heart of the Lesser Sunda region on the islands of Komodo, Rinca, Padar, Gili
Motang, Gili Dasami (also known as Nusa Kode), and the Western coast of Flores
Island (Dunn 1928; Fig. 2.1). Five of the islands are within the boundary of Komodo
National Park (PHKA 2000; Fig 2.1). In studies conducted after 1991, this species
could not be found on Padar Island (Ciofi & de Boer 2004; Jessop et al. 2004;
Sastrawan & Ciofi 2002).
7
Figure 2.1 Distribution of Varanus komodoensis. Grey areas represent the current distribution; black areas
represent areas that were identified as part of the distribution by Auffenberg (1981); hatched areas
indicate where V. komodoensis have been reported by villagers.
Source: Modified from Ciofi & de Boer 2004; Jessop et al. 2006
unpublished data.
Even though the Komodo dragon is not threatened by the leather trade, like
the congeneric Water Monitor (V. salvator), and is considered ‘dangerous’ to humans
(King et al. 2002; Shine et al. 1996; Ellis 1998), hunting and trade in this species has
9
been occurring for a long time. Since the 1930’s Komodo dragons and their eggs have
been hunted illegally for zoo collections and for traditional medicine (Primack et al.
1988). Hien (2003) reported that the local people of Riung, Northwest Flores, claimed
that they once illegally caught 50 live specimens of the Komodo dragon for a
foreigner. However, the widespread hunting and trading of other reptiles including
varanids, for their skins and for food (e.g. Shine et al. 1996) should be considered a
potential threat to the Komodo dragon.
on the islands of Java, Bali and throughout Lesser Sunda region, V. salvator may
reach up to 218 cm in length and weigh up to 25 kg (Gaulke & Horn 2004; Horne &
Gaulke 2004).
Komodo dragons can be found from sea level up to about 800 meters in
altitude, mainly in tropical dry and moist deciduous monsoon forests (Ciofi 2004).
Indeed, this species is generally distributed over entire islands within KNP and
western coastal on Flores but is rarely found above 500 meters (Auffenberg 1981).
All varanids are insectivores or carnivores. Unlike other varanids that rely on
smaller prey species, Komodo dragons are able to feed on larger vertebrate species,
such as the Timor Deer (Cervus timorensis), water buffalo (Bubalus bubalis), or
small wild boars (Sus scrofa). Adult Komodo dragons mostly rely on a sit-and-wait
hunting strategy to catch their prey (Auffenberg 1981; Green et al. 1990; King &
Green 1999; Pough et al. 2001). Hatchlings and juveniles, however, feed on a diverse
diet of insects, small lizards, snakes and birds, and use more active hunting strategies
than adults (Auffenberg 1981; Ciofi 1999; Mattison 1992).
Female Komodo dragons are known to breed when they reach a body weight
of around 20 kgs (King & Green 1999). Females begin nesting in August, as
determined by the presence of recent digging activity on the nest, or by repeated
observation of individuals in association with the nest during the nesting period. The
nesting period is from August through November, with egg deposition occurring in
September (Ciofi 1999; Jessop et al. 2004). Up to 38 hatchlings will emerge from the
nest at the beginning of the dry season (Auffenberg 1981; Jessop et al. 2006
11
Generally, mortality in lizards is highest during the first phase of their life because
they are more vulnerable to predation (Mattison 1992). Poor maternal body condition
and stress can decrease the dispersal tendency of juveniles (Meylan et al. 2002).
Typically, natural populations show substantial variation, in locomotor performance
and body size, which is related to offspring survivorship (Clobert et al. 2000). Thus,
the growth and survivorship of offspring in reptiles is greatly affected by the animal’s
environment (i.e. Brockelman 1975; Gans & Pough 1982).
Most juvenile reptiles leave the natal area in which they were born and move
into new habitats that are not already occupied or to avoid cannibalism by adults
(Pough et al. 2001). Greenwood (1984) noted that natal dispersal among juveniles is
also considered as a mechanism to avoid future inbreeding. Heatwole (1976) and
Sarno et al. (2003) described natal dispersal as being driven by the competition for
food resources and territories with adult.
The first scientific study on the Komodo dragon was conducted by Major Peter A.
Ouwen, director of the Zoological Museum in Buitenzorg (now Bogor), Java, in 1912
(Auffenberg & Auffenberg 2002). Ouwen described the species for the first time.
Later, Dunn (1927 & 1928) conducted the first significant observations and gave the
initial information on the habitat and distribution of Komodo dragons. Auffenberg
(1981) contributed to the first comprehensive study on the ecology and behavior of
this species. On the basis of 13 months of field observations, Auffenberg (1981)
provided the base line information on aspects of the Komodo dragon’s ecology and
behavior. The most recent field study on the Komodo dragon was conducted by
Jessop et al. (2006 unpublished data). Since 2002, they undertook a broad scale
investigation in the ecologyl and demography of the Komodo dragon, which has
provided important information and instigated a program to monitor population
trends. Although numerous field studies on the ecology of Komodo dragons have
been conducted, yet there are no detailed studies on the ecology on juvenile Komodo
dragons (Auffenberg & Auffenberg 2002). Walsh et al. (2002) studied growth in
13
GENERAL METHODOLOGY
The study was conducted within the Komodo National Park, East Nusa Tenggara,
Indonesia, approximately 500 km east of Bali. The Park lies in the Lesser Sunda
region between Flores Island and Sumbawa Island (8°35’40” S and 119°25’51”E),
encompassing two main islands; Komodo and Rinca; three smaller islands, Padar, Gili
Motang and Nusa Kode, and numerous smaller adjacent islands. Komodo National
Park established by the government of Indonesia in 1980, enhancing its status from
Wildlife Reserve in 1970. Before declared as national park, this park was declared as a
Man and Biosphere Reserve by UNESCO in 1977. Further, in 1991 UNESCO
designated this park as a World Heritage Site. The park was initially established to
conserve the unique Komodo dragon (Varanus komodoensis) and was identified by
both WWF and Conservation International as a global conservation priority area. This
area is a transitional zone between the Australian and Asian fauna and flora,
containing elements of both (PHKA 2000).
The park encompasses the extant distribution of the endangered species of the
Komodo dragon and is approximately 1817 km2 in size encompassing both marine
(1214 km2) and terrestrial (603 km2) habitat (PHKA 2000). Komodo and Rinca are the
two largest island, followed by Padar, Gili Motang, and Nusa Kode. Most of the
islands are mountainuous (the highest peak is 800 m asl on Komodo Island),
15
combining a mostly grass-covere rugged topography, with variuous size of flat valleys
on the lower elevation (up to 50 m asl) (Dunn 1928; Monk et al. 1997).
Komodo Island is the largest island with the highest peak reaching 800 m asl
and is mostly covered by grassy savanna habitat (59.38%) (Table 3.1). Rinca the
second largest island is also predominantly covered by savanna habitat (55.02%) and
the highest peak is 712.5 m. Padar, the third largest island, has the highest percentage
cover of savanna habitat (93.48%) amongst other large islands. The other two main
islands, Gili Motang and Nusa Kode are predominantly covered by open deciduous
forest (79.96% and 84.33%, respectively). The other numerous small islands are
predominantly covered by savanna grass.
There are four existing settlement within the park (PHKA 2000), namely
Kampung Komodo (Komodo Island), Rinca, Kerora (Rinca Island), and Papagarang
(Papagarang Island). The settlements were established prior to 1980, before the area
declared as national park. Total population of these settlements was estimated
approximately 2300 people in 1999.
Table 3.1 Size of each island within KNP and it’s vegetation cover (in km2).
The climate is dry and is influenced by monsoons and trade winds. Rainfall
varies with the elevation, averaging less than 500 mm a year, with peak values
between December and February (Monk et al. 1997). Jessop et al. (2006 unpublished
16
data) classified the vegetation cover of each island, with modifications based on
classification by Auffenberg (1980) as shown in Table 3.1:
- Quasi Cloud Forest (QCF) is only found above 500 m asl. This forest
has a moisture and cooler climate and is characterized by moss-covered
rock, bamboo groves, and rattan.
The overall study was conducted across 10 sites on four islands, but the radio
telemetry study was only done in Loh Liang. The study sites are; Loh Sebita, Loh
Liang, Loh Lawi, and Loh Wau on the Komodo Island, Loh Buaya, Loh Baru, Loh
17
Tongker, and Loh Dasami on Rinca Island, Nusa Kode Island, and Gili Motang
Island. The description of each study site is as follows (Figure 3.1);
1) Loh Sebita (SB) (8º32’ N, 119 º32 E). This site is located on the east coast
of Komodo Island. The valley is characterized by a dense mangrove forest
stretching along the east coast of the valley. Towards the west, the
topography increases gently from sea level to about 735 m asl, the second
highest peak of the island. To the south uplifted rugged mountain stands
delimit the valley with smoother hills to the north. Vegetation in Loh Sebita
is dominated by the open deciduous forest type.
2) Loh Liang (LG) (8º34’ N, 119º29 E). This site is located at the north-east
part of the island, approximately 6 km south of Loh Sebita. Loh Liang is a
wide valley and is characterized by ODF. CDF exists along the northern to
western parts. This valley is delimited by smoother hills to the west and
north, which rise from sea level to the highest altitude of the island (800 m
asl to the west and 740 m asl to the northwest). To the East, this valley is
delimited by the rugged mountain stands. Vegetation in Loh Liang is
dominated by the open deciduous forest type, except in the northern deeper
valley, close dense forest exists.
3) Loh Lawi (LW) (8º36 N, 119º26 E). This site is located at the center of
Komodo Island, approximately 7 km southern of Loh Liang. This valley is
characterized by a long (approximate 6 km from the coast line) and narrow
area, delimited by rugged mountatins to the south, west, and north. To the
east, Loh Lawi is covered by a thick mangrove forest along the south coast.
Across the valley, open deciduous forest is predominantly.
4) Loh Wau (WA) (8º41’ N, 119º26’ E). This site is located on the south of
Komod Island, approximately 17 km from Loh Liang. The valley is
delimited by rugged mountainous topography to the west and smoother hills
to both the south and north. The close dense forest vegetation type is
predominantly in this very small valley.
18
5) Loh Buaya (BY) (8º39’ N, 119º43’ E). This site is located on the northern
part of Rinca Island. This valley has a smooth topography. Savanna is
predominant in the valley with a small open deciduous forest to the middle
of the valley.
6) Loh Baru (BR) (8º43’ N, 119º41’ E). This site is located to the east of the
island, approximately 9 km south of Loh Buaya. The local people occupied
this valley before the park was established. Later, the settlements moved to
the nearest villages; Kerora and Rinca village. In Loh Baru agricultural
crops predominantly in the valley, while to the deeper part of the valley
open deciduous forest exists. The Loh Baru site also includes Loh Baru
valley and Sok Niu valley, which is located 2 km north of Loh Baru.
Komodo dragons are known to move reoutinely between these two valleys.
To the west and south, Loh Baru is delimited by rugged mountains which
lead to the highest peak altitude of the island (712.5 m asl).
7) Loh Tongker (TK) (8º45’ N, 119º43’ E). This site is located to the
Southeastern of Rinca Island, approximately 12 km southern Loh Buaya.
This small and narrow valley is predominantly covered by open deciduous
forest. Loh Tongker is delimited by rugged and steep hills.
8) Loh Dasami (DS) (8º46’ N, 119º39’ E). This site is located on the southern
part of Rinca Island, approximately 15 km southern Loh Buaya, and is
characterized by a very closed dense forest. This small valley is delimited
by a rugged mountainous terrain to the west, north and east, and the highest
peak has an altitude of 712.5 m asl.
9) Nusa Kode (NK) (8º47’ N, 119º39’ E). This site is located as a small island
is southern Rinca Island, approximately 1.5 km across Loh Dasami valley.
Nusa Kode is a rugged mountainous island and has few small and narrow
valleys. The island is predominantly covered by open deciduous forest from
sea level to the peak (400 m asl). Savanna exists on the eastern side of the
island.
19
BORNEO
KOMODO
1
4
PADAR
6
RINCA
8
N 7
0 5 10 Kilometers
9 10
Figure 3.1 Study sites across the four islands of Komodo National Park.
Numbers are dedicated to show the locations; 1. Loh Sebita; 2. Loh
Liang; 3. Loh Lawi; 4. Loh Wau; 5. Loh Buaya; 6. Loh Baru; 7. Loh
Tongker; 8. Loh Dasami; 9. Nusa Kode; 10. Gili Motang.
20
10) Gili Motang (GM) (8º48’ N, 119º47’E). This mountainous island is located
8 km southeast off Rinca Island, and 3 km southwest off Flores. This
isolated island is predominantly covered by open deciduous forest
dominated by Schoutenia ovata. Along the western to the southern part of
the island, savanna exists up to the peak (425 m asl) with small closed dense
forest on the peak. This island has received less attention as compared to the
other places within the park in regards to conservation management (Jessop
et al. 2006 in press).
Hatchlings among species in Varanids vary greatly in size (King & Green 1999). They
range from 18 grams in weight and approximately 17 cm in length in V. rosenbergi,
80 to 127 grams in weight and are about 30 cm in length in V. komodoensis.
Auffenberg (1981) found that average total length of Komodo dragon hatchling was
30.4 cm in SVL and weight 80.3 grams, and were considerably longer and larger than
the other large Varanid species.
The juvenile in Komodo dragon were estimated below maturity sexual age,
which is estimated 5 years (King & Green 1999). Lemm et al. (2002) defined those
juvenile Komodo dragons which less than three years had 38 – 40 cm in SVL and 0.75
– 1.30 kg in weight. Large Komodo dragons (>2.3 m in total length and >38 kg in
weight) are likely to be males, but younger and smaller animals are difficult to sex
(Jessop et al. 2006 unpublished data; Auffenberg 1981). Within this study, hatchling
was defeined as recently hatched animal (below one year), while juvenile was above
yearling animal. Animal thus met all of the measurements ranges as shown in table 3.2
below were classified either as hatchling or as juvenile.
21
- Hatchling
Hatchling has a brown pattern with large, distinct orange spots on the
dorsal region. Yellow stripes present on it’s neck to the dorsal region.
Light yellowish spot pattern found on its four legs from the temporal
region to the front. Front legs are brown with white flecks disposed in
round, horizontal circles. The ventral part of the body is light yellow
with large dark spots. Tail has some light yellow strip circling.
- Juvenile
Juvenile pattern is gradually lost with age. They still show a lighter skin
color, but have a darker brown pattern. Small light yellow spot pattern
still exists on its four legs. Tail has lighter color (as compared to
hatchlings) strip circling.
Juveniles
min 6.50 35.00 0.600
max 13.50 80.10 10.000
3.2.2 Radiotelemetry
process, and the proximate causes of mortality. White and Garrot (1990) emphasized
the use of radiotelemetry to investigate habitat requirements, and demography. Similar
techniques have been demonstrated in studying spatial ecology of reptiles (i.e.
Fitzgerald et al. 2002; Sastrawan & Ciofi 2002; Webbs & Shine 1997).
Komodo National Park, which is being conducted by the Center for Conservation and
Research of Endangered Species, Zoological Society of San Diego since 2002 (Jessop
et al. 2006 unpublished data). Hatchling and juvenile Komodo dragons were captured
using several techniques; including nest enclosure traps (using metal sheeting fence),
baited (box and enclosed-pipe) traps, noose or by hand. Following capture, for safety
reasons, captured Komodo dragons were restrained either by electric tape or by rope
on its four legs and mouth taped. Capture and measuring processes were no more than
30 minutes. No anesthesia was used in handling the animals. To avoid injury to
animals, physical restraint in handling the animals within this study was followed
Friend et al. (1996).
Each trap (300 X 60 X 60 cm) was constructed from alloy and aluminum metal
sheets with two open-ends with sliding doors. Only one door (front side) was kept
open to allow animals to enter the trap. The front door was connected by a metal wire
with bait at the other end of the wire. Each trap was baited with goat-meat, where two
or three pieces were placed on the floor to attract the juvenile to enter and approach
the hanging bait. The front door would close whenever the hanging bait was pulled.
Eight traps were placed along a line transect with an interval of 250 - 450
meters between traps depends on the terrain and covered the valleys as much as
possible. Traps were checked twice a day (morning and afternoon session) and moved
for every three days to allow for sufficient production of the volatile oils that were
released as the bait decomposed. Trap locations were marked by GPS (Garmin Etrex
Summit). Recaptured and resighted animal within the same trapping session in each
location were recorded and considered as Within Trip Recapture or Resight.
To capture hatchling, this study was also employing enclosed PVC pipes 1m in length
and 0.1 m in diameter where the bottom side is closed. To attract the hatchling, this
trap was baited using meat. Similar techniques were implemented with other reptiles
species as reported by Mathies et al. (2001) and Schemnitz (1996). Each pipe-trap was
placed on a tree in 100 m line transects at a 5 m distance and approximately 1–2 meter
above ground to attract hatchling and avoid disturbance by bigger Komodo dragon.
Pipe-traps were placed near the active nests to anticipate escaping hatchlings and near
25
ranger station of station where occasionally hatchlings were found every hatchling
seasons.
The noosing technique has been demonstrated by many authors, such as Olsson and
Madson (2001) and Wapstra and Swain (2001). The noosing technique required two
persons and relies on a systematic search. Searching was conducted opportunistically
by foot and covered almost entire the valley. Noosing technique employed one 2
meters of woody “Y” stick with lasso.
Female Komodo dragons are known to begin nesting in August, (Jessop et al. 2004),
and lay eggs in September (Ciofi 1999). Up to 40 hatchlings will emerge from the nest
at the beginning of dry season (February to March) and weigh between 60–130 grams
and are about 20-60 cm in length (Auffenberg 1981; Ciofi 1999; King & Green 1999).
The spatial ecology of juvenile Komodo dragons was assessed based on following
spatial parameters; spatial movement, activity areas, habitat use, daily activity
patterns, and annual movement. These parameters were used by Fitzgerald, et al.
(2002); Ibrahim (2002), Piepgrass and Lang (2000), and Olupot et al. (2001), in
assessing spatial ecology in reptiles. Detailed descriptions of each analysis will be
included in the relevant chapters.
Spatial movement of each size class was determined using data obtained from the
radiotelemetry study of hatchlings and juveniles. Spatial movement is defined as a
straight line moved between two consecutive points (Samuel & Fuler 1996). This
measurement will provide a minimum distance that an animal could have traveled to
reach the new location. This technique was also employed by Ibrahim (2002) and
Fitzgerald et al. (2002). Spatial movement data will be calculated by means of a
computer program ESRI ArcView 3.2 (ESRI 1999) with X Tools and Animal
Movement program (Hooge et al. 1999).
Long distance movement in Komodo dragons was analyzed using data obtained from
the Mark-Recapture study (Husak & Fox 2003; Lancia et al. 1996; & Nietfield et al.
1996). This long distance movement is used to measure the ability of hatchling and
juvenile Komodo dragons to move to adjacent valleys or adjacent island. Long
distance movement ability was measured as a straight distance moved between two or
more consecutive points (Samuel & Fuler 1996) between capture locations from
subsequent days and years and were calculated by means of computer program of
ESRI ArcView 3.2 (ESRI 1999) with X Tools and Animal Movement program
(Hooge et al. 1999).
CHAPTER IV
4.1 INTRODUCTION
The study was conducted in the Loh Liang valley on Komodo Island, (8°33’40” S and
119°29’51”E) in Komodo National Park (KNP), East Nusa Tenggara, Indonesia.
General descriptions on vegetation cover of KNP are described elsewhere (e.g.
Auffenberg 1981; Jessop et al. 2006 unpublished data; Monk et al. 1999). Komodo
Island is a rugged, mountainous island covered predominantly by savannah.
Deciduous monsoon forest dominated by Tamarind tree (Tamarindus indica).are
found in coastal valleys, while closed evergreen forest persists on hills above 500 m.
30
Five hatchling Komodo dragons with a mean snout to vent length (SVL) of 20.16 ±
0.85 cm (SEM) (range 18.25 – 22.6 cm) and body mass of 0.11 ± 0.01 kg (range
0.095 – 0.135 kg) were captured following emergence from their nest. Hatchlings
were captured by hand from fenced nests or using PVC pipe traps (10 cm diameter
and 100 cm in length). Seven juvenile Komodo dragons (mean ± SEM of SVL 55.81
± 2.97 cm and body mass of 3.03 ± 0.56, range 1.4 – 5.7 kg) were captured by hand
or in goat meat baited box traps. Trapping methods are described in Chapter III in this
thesis. Measurements of individual animals are summarized in Table 4.1.
This study was conducted between March 2004 and June 2005. In 2004, two
hatchlings and four juveniles were radio-tracked, and in 2005 three hatchlings and
four juveniles were followed. Telemetry equipment consisted of activity-sensitive
AVM G31V transmitters (AVM Instruments Co. Ltd.), a AVM LA12Q receiver, and
a three-element yagi antenna. Each transmitter was attached to the dragon’s base of
tail using duct tapes. In 2004, 8 transmitters were attached to four hatchlings and four
juveniles and in 2005, 6 transmitters were attached to three offspring and three
juveniles.
Table 4.1 Summary of SVL, weight and duration of radio-tracking for hatchling
and juvenile Komodo dragons.
Observations
PIT tag ID Year SVL (cm) Weight (kg) Positional
#
Hatchling
64E4218 2004 22.6 0.14 50 12
64CDC09 2004 21.3 0.11 26 6*
63DFB2A 2005 18.25 0.1 66 11
64D4C0E 2005 18.3 0.09 51 18
63C1383 2005 20.35 0.11 67 14
Mean ± SEM 20.16 ± 0.85 0.11 ± 0.01 52.00 ± 16.60 12.2 ± 1.96
Juveniles
64CF2AO 2004 61.85 3.5 18 17
639E332 2004 68.35 5.7 39 17
643761C 2004 58.05 3.5 18 15
64E2E95 2004 55.45 3.4 14 14
63DE6F1 2005 51.15 1.65 57 16
63D9C4B 2005 44.45 1.4 44 15
63DB3B7 2005 51.35 2.05 7 6*
Mean ± SEM 55.81 ± 2.97 3.03 ± 0.56 28.14 ± 7.78 14.29 ± 1.44
* Excluded from movement and activity area calculation due to lack of fixes.
To quantify and compare the habitat use of juvenile and hatchling lizards, habitat and
shelter site parameters concurrently with positional data of lizards were collected
32
during radio-tracking (Olupot & Waser 2001). Four parameters were measured and
defined as follows;
1) Habitat strata
Use of either terrestrial or arboreal strata
2) Habitat type
The presence of lizards in one of three key habitat types: open deciduous forest,
closed dense forest, and savannah grassland. These habitat types were readily
distinguished by canopy closure and floristic composition.
3) Shelter site
Included a description of resting or sleeping location based on the use of
vegetation (identified to species), or substrate type. Further, with relation to
vegetation, particularly trees, the tree diameter at breast-height (Dbh) and tree
height were also recorded. Tree Dbh was obtained by measuring the tree
circumference at breast-height and calculated using the formula circumference =
2πr (where: π = 3.14, 2r = diameter). Tree height and Komodo dragons’ position
above ground was calculated using a Suunto Clinometer PM5 (Suunto, Finland).
4) Elevation
Ground elevation at which a lizard was located was recorded alongside every fix
locations.
Directionality and tortuousity were analyzed to describe the pattern of path movement
(linearity) of Komodo dragons (Nams & Bourgeois 2004). Directionality was
analyzed by measuring the turning angle of movement on each point in a compass
direction (Claussen et al. 1997). The turning angles indicate the movement angles
between two consecutive points (Higham et al. 2001). Tortuosity was measured by
33
analyzing the fractal dimension (D) (Milne 1991), whose values range between 1-2:
D = 1 the animals’ path of movement is straight; D = 2 (i.e. maximum) the animals
path of movement is so tortuous as to completely cover a plane (Nams & Bourgeois
2004). To describe site fidelity, Site Fidelity Test was ran with 1000 replications to
obtain the r2 value, the lower r2 value, the highest fidelity of the animal (Hooge &
Eichenlaub 1997). Directionality and Site Fidelity were computed using the Animal
Movement Program extension (Hooge et al. 1999) and X Tools of ArcVIew 3.2
(ESRI) while Fractal dimensions were computed using the computer program Fractal
(Nams 2004a).
B) Activity area
In this study, the activity area was defined as the area occupied by an animal during
the study period. Core area was defined as the area in which studied animal mostly
visited or occupied during the study period. Activity areas were calculated using three
methods: the 100% Minimum Convex Polygon (MCP), 95% and 50% Adaptive
Kernell Analysis (AK). MCP is the most common and simplest method used to
estimate the activity area of animals by calculating the smallest possible convex
polygon around the outermost animal locations (Kenward 1993; Samuel & Fuller
1996). Sastrawan & Ciofi (2002) applied MCP to measure adult Komodo dragon’s
activity area size. However, MCP only uses a proportion of location data; it is
sensitive to outliers and incorporates large areas that may have never been used by the
animal (Melville & Swain 1999; White & Garrot 1990). The 95% AK is also suitable
34
To increase the quality of data, radio-tracking animals with less than 10 fixes
(data for hatchling 64CDC09 and juvenile 6eDB3B7) were excluded. Daily
movements and activity areas were analyzed based on data subset of four hatchlings
and six juveniles with minimum 11 fixes. Activity area estimations were calculated
using the Animal Movement Program (Hooge et al. 1999) and X Tools extension of
ArcVIew 3.2 (ESRI). A similar technique for studying spatial ecology was applied by
King and Gurnell (2004) and Piepgras and Lang (2000).
C) Habitat Use
Patterns in habitat use were analyzed by comparing the frequency of particular habitat
types (arboreal and terrestrial), types of vegetation (closed forest, open forest, and
savanna), elevation of location, shelter sites, tree species, tree DBH and tree height,
and animal position above ground. To determine if Komodo dragons were using their
habitat in a non-random fashion, their observed use of habitat was compared with a
random selection of locations calculated using the program Ecological Methodology
V. 6.1 (Krebs 1999; 2002) to generate 100 random latitude and longitudes positions
within Loh Liang. These random locations were then assigned their habitat type and
elevation drawn from digital topographic maps.
35
D) Statistical Analysis
Prior to analysis all data was log transformed (Sutherland et al. 2000) to meet the
assumptions of parametric tests (Zar 1999). Log transformed data that did not meet
the assumptions was statistically analyzed using a nonparametric statistical
procedures. All data is presented in tables and graphs of the means and standard error
of the mean (SEM) of back-log transformed data. T-test, Chi square test and Mann-
Whitney U tests were used to determine the in habitat types used (King & Gurnell
2005; Rheaney & Whiting 2003). Linear regression test and ANOVA was used to test
differences in time spent on a tree between hatchlings and juveniles. For all statistical
tests, significance was inferred at P < 0.05.
4.3 RESULTS
During the study, hatchlings dispersed in a predominantly linear fashion with little
movement over previously occupied areas (Figure 4.1). In contrast, juvenile dragons
exhibited more tortuous movement paths reflecting increased movement over the
same area (i.e. use of an activity area) (Figure 4.2). Fractal analysis indicated that the
linearity of the movement paths undertaken by hatchlings D = 1.11 ± 0.01 was
significantly more directed (i.e. strait) than that of juvenile dragons (D = 1.39 ± 0.05;
Mann-Whitney U test; Z = -2.558, p = 0.01). The degree of site fidelity in hatchlings
(r2 = 729502.69) was significantly lower than that in juveniles (r2 = 2729220.96; T-
test t1,9=-39.123.01, p < 0.001).
(Chi square test; x2 = 51.68, p < 0.001) based on an activity index calculated using the
relative proportion of movement fixes compared to the proportion of stationary fixes.
OVERLAPPED
VK ID MCP (ha) OVERLAP (ha) OVERLAP (% )
VK ID
64CF2A0 12.53 2.55 20.34 639E332
639E332 32.37 2.55 7.88 64CF2A0
643761C 4.02 3.66 91.01 64E2E95
643761C /
64E2E95 17.42 6.65 38.20
63DE6F1
63DE6F1 61.77 2.99 4.84 63DE6F1
37
#a
8ー33'55"
8ー33'05"
8ー34'00"
#c
Latitude (South)
#b 8ー34'05"
#d
8ー33'10"
8ー34'10"
8ー34'15"
N N
0 50 100 Meters 0 100 200 Meters
8ー33'15"
8ー34'20"
119ー29'45" 119ー29'50" 119ー29'55" 119ー30'00" 119ー29'50" 119ー29'55" 119ー30'00" 119ー30'05"
Longitude (East)
Figure 4.1 Movement patterns of hatchling Komodo dragons. Arrows indicate starting point and general direction of movements.
Letter refers to hatchlings ID as follow; a) 64D4C0E, b) 63DFB2A, c) 64E4218, d) 63C1383.
38
a b
8ー33'10" 8ー33'50"
8ー33'15" 8ー33'55"
8ー33'20" 8ー34'00"
Latitude (South)
8ー33'25" 8ー34'05"
8ー34'10"
8ー33'30"
8ー34'15"
8ー33'35"
N N
0 100 200 Meters 0 100 200 Meters
8ー34'20"
8ー33'40"
119ー30'20" 119ー30'25" 119ー30'30" 119ー30'35" 119ー30'40" 119ー30'45" 119ー29'40" 119ー29'45" 119ー29'50" 119ー29'55" 119ー30'00"
Longitude (East)
Figure 4.2 Movement patterns of two juvenile Komodo dragons (a: 639E332, b: 63D9C4B). Circles indicate shelter sites.
39
a b
8ー32'45" 8ー32'45" #4
#3 #5 #3
#
8ー33'00" #1 8ー33'00"
#5 #4 #1
8ー33'15" 8ー33'15"
#2
#2
8ー33'30" 8ー33'30"
Latitude (South)
#5
8ー33'45" 8ー33'45"
#6
8ー34'00" 8ー34'00"
#6
8ー34'15" 8ー34'15"
N N
0 500 1000 Meters 0 500 1000 Meters
8ー34'30" 8ー34'30"
119ー29'30" 119ー29'45" 119ー30'00" 119ー30'15" 119ー30'30" a
119ー30'45" b
119ー29'30" 119ー29'45" 119ー30'00" 119ー30'15" 119ー30'30" 119ー30'45"
Longitude (East)
Figure 4.3 Activity area overlaps of six juvenile Komodo dragons as calculated by MCP (a) and 50% Adaptive Kernell (b). Numbers
refer to juvenile ID as follow: 1) 64CF2A0, 2) 639E332, 3) 64E761C, 4) 64E2E95, 5) 63DE6F1, 6) 63D9C4.
40
Activity areas of hatchlings and juveniles ranged between 1.72 – 4.83 ha and 4.02 –
61.77 ha for MCP; 12.14 - 20.12 ha and 5.49 – 33.23 for 95% AK; and 2.51 – 4.74
and 0.56 – 10.48 ha for 50% AK (Table 4.2, Figure 4.3). Hatchlings used significantly
smaller activity areas, of approximately 8 times less than juveniles for the 100% MCP
(T-test; t1,9 = -3.66, p = 0.001). However, there was no significant differences in 95%
AK (T-test t1,9 = -0.02, p = 0.18) and neither for 50% AK (T-test t1,9 = 0.396, p =
0.70). In five of six cases, the activity areas of juvenile dragons overlapped with one
(n = 4) or two (n = 1) conspecifics. The extent of overlap of activity areas ranged from
4.84 – 91.01 % or 2.55-6.65 ha (Table 4.4; Figure 4.3.a).
120
255
100
a
40
53
20
6
0
Terrestrial Arboreal
Habitat Strata
120
b
Frequency of arboreal activity (%)
100
80
60
40
20
10 20 30 40 50 60 70 80
SVL cm
80 42 73
c
Hatchling
Juvenile
Frequency of Use (%)
60
40
20 16
6 7
5
0
Closed Forest Open Forest Savanna
Habitat Type
Figure 4.4 Differences in terrestrial and arboreal habitat use by hatchling and
juvenile Komodo dragons (a). Figure b depicts the relationship
between body size (SVL) and arboreal activity in the radio tracked
Komodo dragons.
42
Shelter sites were defined as those places used by hatchlings and juveniles for resting
(also as sites between movements - for juveniles only) or as overnight refuges.
Hatchlings used trees exclusively for shelter. They significantly used live (84.71 %)
over dead trees (15.29 %; Chi square test; x2 = 122.859, p < 0.001) (Table 4.4).
Amongst live trees (15 species recorded), there was a significant tendency for
hatchlings to occupy Tamarindus indica tree (45.37 %) (Chi square test; x2 = 122.86,
p < 0.001) (Table 4.5). Juveniles used predominantly crevices under or among rocks
(51.77 %) as shelters. These shelters could be a part of a larger forested area with
significant granitic exfoliation forming rocky fields or a clump of rocks associated
43
with trees. Trees (9 species recorded) were occasionally used by juveniles as locations
for resting or basking (Table 4.5).
With respect to arboreal habitat use, hatchlings used significantly thinner (tree
DBH; T-test t1,10 = -2.634, p = 0.025) and shorter trees (T-test t1,4 = -2.915, p = 0.043)
than juveniles (Table 4.6). However, there was no significant difference in tree
elevation use between hatchlings and juveniles (T-test t1,4 = 2.23, p = 0.089).
Table 4.5 Major tree preferences used by hatchlings and juveniles (%).
Table 4.6 Total number of tree species, and mean values of tree parameters
used, and animal position on tree.
4.4 DISCUSSION
Hatchlings displayed several key differences with respect to their spatial ecology
compared to juvenile Komodo dragons. In particular, their post-natal movements were
essentially linear compared to those of juveniles that appeared to have already
developed a defined area of established activity. This predominantly linear movement
could possibly underpin a deliberate type of dispersal behavior that enables hatchlings
to move as far as possible away from their nesting sites relative to the actual distance
traveled. Based on the limited number of hatchlings radio-tracked, there was no
indication that hatchlings were heading towards a consistent bearing similar to natal
dispersal in other reptiles including hatchling Green sea turtles (Chelonia mydas) that
display well-oriented post-emergence dispersal (Salmon & Wyneken 1987). Rather,
the movement of hatchling dragons was similar to patterns of natal dispersal
demonstrated by hatchlings of Blanding turtles (Emydoidea blandigii) (McNeil et al.
2000) and the Australian freshwater crocodile (Crocodylus johnstoni) (Tucker et al.
1997).
Pough et al. (2001) described that lizard hatchlings moved out of their natal
areas to find a suitable patches of habitat which are not already occupied by larger
conspecifics or to avoid cannibalism by adults. As reported by Sarno et al. (2003),
dispersal in juvenile Guanacos occurred in order to avoid conflict with adults.
Cannibalism among Komodo dragons has been observed (Auffenberg 1981) as well as
in other Varanids such as V. griseus, V. gouldii and V. gigantheus (King & Green
1999). This early life-stage is dominated by high mortality because hatchlings are
particularly vulnerable to predation (Mattison 1992).
From the data, hatchlings displayed a consistent pattern in habitat use. They
were arboreal during their first stage of life. Following emergence, hatchlings
immediately climbed the nearest tree (Auffenberg 1981). Their arboreal nature is
presumably a strategy for avoiding their terrestrial based larger conspecifics and
possibly aerial predators such as birds of prey (Auffenberg 1981; King & Green
1999). Thompson (1993) reported similar behaviour in V. caudolineatus. Hatchlings
of Blanding’s turtles hide in vegetation soon after emergence (McNeil et al. 2000).
Arboreal habits could provide advantages for hatchlings to test differences in escaping
tactics from predators (Higham et al. 2001). Arboreal habitat also provides resources
such as food and shelter (Reaney & Whiting 2003). Hatchlings were observed to feed
on Gecko gecko (PIT tag ID 64E4218 and 64D4C0E) which are commonly on trees in
Komodo island (Auffenberg 1980). Unfortunately, this study could not determine how
long the hatchlings spent their life as being arboreal. Observations from KNP rangers
suggest that hatchlings could lead an arboreal life for up to five years.
Even thought juveniles were terrestrial, three out of six juveniles (SVL < 60
cm) were found occasionally on trees for overnight shelter. Auffenberg (1981)
reported a similar arboreal characteristic on juveniles less than 70 cm in SVL.
Arboreality in juvenile of other terrestrial Varanid species were also reported in V.
doreamus (Bohme et al. 2004) and V. salvator (Gaulke & Horn 2004).
The rate and distance moved by hatchlings within their natal valley was
relatively slow and small compared to juveniles, suggesting that this early dispersal
phase was not particularly important for the overall pattern of dispersal from natal
sites. Movements in hatchlings were shorter than in juveniles. Mean juvenile daily
movement was four times larger than that of hatchlings, but six times smaller than
adults (Sastrawan & Ciofi 2002). As reported by Perry and Garland (2001), terrestrial
lizards have larger movement and activity area size than arboreal individuals. In
contrast, Pough et al (2001) noted that arboreal reptiles have a low movement pattern.
King and Green (1999) noted that arboreal monitors such as V candolineatus and V.
brevicauda showed a similar characteristics to Komodo dragons.
46
Movement in juvenile reptiles increase with age (McMaster & Herman 2000).
The study indicated that the larger the animal, the larger the daily movement and the
longer the movement. Sastrawan and Ciofi (2002) reported similarity in adult male
and adult female Komodo dragons. Similar patterns in other species reptiles were also
reported, such as in arboreal snakes (Fitzgerald et.al 2002; Pearson et al. 2005).
This study also suggested that juveniles were able to explore and expand their
foraging areas better than hatchlings, which had to familiarize with their new
environment after emergence. Activity area size increased as Komodo dragons grew
older. Average adult activity area is 19 times larger than in juveniles (see also
Auffenberg 1981; Sastrawan & Ciofi 2002). Smith et al. (1999) reported that the adult
Angonoka turtle Geochelone yniphora has a larger activity area size compared to
juveniles. According to Pough et al. (2001) many authors reported a positive
correlation between activity area size and body size in many species of reptiles.
Christian and Waldschmidt (1984) demonstrated that the lizard’s body size and sex
were significantly correlated to size of activity area. Perry and Garland (2002) also
demonstrated that activity area is correlated with body size in different species of
lizards.
According to this study, both hatchlings and juveniles had small core activity
areas. Komodo dragons used core areas as shelter sites rather than for foraging. Small
core area sizes were also reported for the Massasauga (Johnson 2000). In hatchling
Komodo dragons, the small size of the core area was probably due to less active
behavior in this age class as far as exploration of their new environment concerned.
Yet, juveniles preferred particular spots suitable to their needs. Although juveniles
moved away from their overnight shelter, they returned to their shelter in the
afternoon. Nevertheless, some juveniles remained at the same overnight shelter for a
47
few days (e.g. 639E332, and 63D9C4B). However, many authors reported that there is
a significant correlation between age or sex class, activity and core area in other
reptile species, such as snakes and turtles (e.g. Melville & Swain 1999; Piepgrass &
Lang 2000; Whitaker & Shine 2003). King and Green (1999) suggested that core
activity areas varied seasonally. In this study, however, it was not possible to
investigate seasonal movements and seasonal activity areas.
size. Auffenberg (1981) described that since vegetation could limit the vision of adult
Komodo dragons, it was important for small Komodo dragons to hide from the adults.
Besides using the Tamarind tree, hatchlings were often found hiding on the
Gebang tree (Corypa utan). Live Gebang trees contain dead branches at the base of
their trunks that provide cover for hatchlings to hide, while dead Gebang tree provide
holes at relatively high elevations above that ground. Blazquez and Rodriguez-Estrella
(2001) in their study of Spiny-tailed iguanas, Ctenosaura hemilopha, reported that
selected trees could allow hatchlings and juveniles to take shelter, to feed, basking in
the morning, and minimize the risk of predation.
In their early life stage, Komodo dragon showed their tendency to discover
their new environments in order to find suitable and safest space from predators.
Exploration was demonstrated by the increasing in size of movement and activity
areas along with their age. McMaster and Herman (2000) documented that along with
increasing of age, movement in juvenile reptiles become larger. Greenwood and
Swingland (1984) noted that animals are out of their natal sites to exploit available
resources. Immature Komodo dragons demonstrated their requirements of hidden
places to avoid conflict with their conspecific or other predators, including adults, as
noted by Mattison (1992) that mortality in lizards is highest during the early phase of
their life because they are more vulnerable to predation. Further, long-term and
comprehensive studies are important to investigate detail patterns in spatial use during
early life stage of Komodo dragons.
CHAPTER V
5.1 INTRODUCTION
There are three critical mechanisms for dispersal: (1) leaving the natal site
(natal dispersal), (2) dispersing in terms of accession to breeding site (breeding
dispersal), and (3) settling into a new home range (immigration) (Lebreton et al.
2003). Natal dispersal is the movement of individuals from their place of birth to their
first breeding location (Greenwood 1980). According to Pough et al. (2001) hatchling
reptiles move out of their natal areas to find suitable patches of habitat that are not
already occupied by larger conspecifics. This allows access to specific resources and
avoids cannibalism by adults. Greenwood (1980) also defined breeding dispersal as
the movement of individuals between successive breeding sites, either within a
breeding season or between breeding seasons. Bancroft and Smith (2005)
demonstrated how movement into a new home range was correlated with a specific
dispersal mechanism.
50
Dispersal can also play an important role in the behavior and social systems of
animals (Wolff 1999). Caro (1994) and Gompper (1996) suggested that dispersal
could also contribute to increased foraging success and thus decreased competition
among males. Female breeding dispersal is related to mate choice in order to improve
mate quality and locate resources that facilitate offspring fitness (Otter & Ratcliffe
1996).
The absence of gene flow, on the other hand, increases inbreeding and genetic
drift and, in small populations, the probability of extinction (Konuma et al. 2000;
Madsen et al. 1996). Low genetic variability is generally correlated with inbreeding.
Rosenfield and Bielefeld (1992) reported a relationship between low natal dispersal
and high inbreeding incidence in Cooper’s hawk. Inbreeding can cause severe low
fitness for small populations (Lande 1998).
51
To date, there has been no detailed study on the dispersal capacity of Komodo
dragon (Auffenberg & Auffenberg 2002; Sastrawan & Ciofi 2002). The objective of
this study is to investigate the dispersal capacity in immature Komodo Dragons (i.e.
hatchlings and juveniles). Sutherland et al. (2000) suggested that movement from the
natal site in the early life stage of an animal is important to the processes of
demography, population dispersion, colonization, and gene flow. As parental care is
rare on reptiles, it is suggested that post-natal or early life-phase dispersal should be
common (Olson & Shine 2003; Shine 1988).
The study was conducted over 10 sites across four islands in Komodo National Park.
These four islands encompass the extant distribution of this species within Komodo
National Park and include populations from Komodo (393.4 km2), Rinca (278.0 km2),
Gili Motang (10.3 km2) and Nusa Kode (referred to as Gili Dasami) (9.6 km2). Study
site description and area size of selected study sites and mark-recapture techniques
were described in Chapter III.
52
During the study period a total of 149 hatchlings and 304 juveniles were
captured and tagged (Table 5.1). Within three years of study, 1 hatchling and 76
juveniles were recaptured in the subsequent years. To increase the sample size for
hatchlings, data on 1 recaptured hatchling from March 2006 was added. These
recapture data were then analyzed for distance of movements (see below).
h
s= ×α (Equation 5.1)
d
Prior to analysis all data were log transformed (Sutherland et al. 2000) to meet the
assumption of parametric tests (Zar 1999). Log transformed data that did not meet the
assumptions was statistically analyzed using nonparametric statistical procedures. All
data is presented in tables and graphs of the means and standard error of the mean
(SEM) of back-log transformed data. The Chi square test was used to determine
differences among slope degree between valleys (King & Gurnell 2005; Rheaney &
Whiting 2003). Paired T-tests was used to determine differences in distance of
movement between WTR data and annual mark recapture data. Pearson correlation
test was used to determine relationship between distances of movement with the size
of the valley. For all statistical tests, significance was inferred at P < 0.05.
Figure 5.1
Frequency (%) Frequency (%) Frequency (%)
0
5
10
15
20
25
0
5
10
15
20
25
0
10
20
30
40
50
60
c
1 0 1 00 10 10 0
b
10 10 0
a
0- 0- 0-
2 0 2 00 20 200 20 200
0- 0- 0-
3 0 3 00 30 300 30 300
0- 0- 0-
4 0 4 00 40 400 40 400
0- 0- 0-
5 0 5 00 50 500 50 500
0- 0- 0-
6 0 6 00 60 600 60 600
0- 0- 0-
Distance (m)
11 -11
Distance (m)
1 1 - 11 1 1 -1 1
0 0 00 00 00 00 0 0
1 2 - 12 12 -12 1 2 -1 2
0 0 00 00 00 00 0 0
1 3 - 13 13 -13 1 3 -1 3
0 0 00 00 00 00 0 0
1 4 - 14 14 -14 1 4 -1 4
0 0 00 00 00 00 0 0
-1 -1 -1
50 50 50
>1 0 >1 0 >1 0
50 50 50
0 0 0
5.3 RESULTS
The study showed a low number of recapture events for hatchlings. From the total of
149 tagged hatchlings only two hatchlings were recaptured. One hatchling of 2004
was recaptured in 2005 at the same location of its first capture in Loh Baru, Rinca.
Another hatchling was recaptured a year (March 2006) after its emergence in 2005
approximately 1500 m from its initial capture location in Loh Buaya, Rinca (Figure
5.1a). On the contrary, juveniles showed a higher degree of recapture than hatchlings.
From a total of 304 juveniles tagged, 74 of them were annually recaptured during the
study period. Most juveniles displayed a very short distance of movement (< 100 m;
Figure 5.1b, 5.1c). Twenty-two (28.95 %) individual juveniles were recaptured within
the same locations of their first captured.
Slope degree was negatively correlated with the size of the valley and the
nearest peak elevation (Pearson correlation test; r = -0.75, p ≤ 0.01 and 0.66, p ≤ 0.01)
(Table 5.3). On the other hand, slope degree was not significantly different among
valleys (Chi square test; x2 = 6.18, p ≤ 0.52) (Table 5.3). Distance of daily movement
and farthest daily movement from WTR data during trapping session showed a
significant correlation with area (valley) size (Pearson correlation test; r = 0.32, p ≤
0.01 and r = 0.41, p ≤ 0.01) in juveniles. There was also a significant correlation
between distance annual movement and area size (Pearson correlation test; r = -0.12, p
57
≤ 0.01) and between farthest annual movement and area size (Pearson correlation test;
r = 0.34, p = 0.02). Distance in annual movement and farthest annual movement
showed a significant negative correlation with slope degree (Pearson correlation test; r
= -0.399, p ≤ 0.01, and r = -0.399, p ≤ 0.01, respectively). In contrast, both annual
movement and farthest annual movement distance were not significantly correlated
with vegetation types (Pearson correlation test; r = 0.07, p ≤ 0.63, and r = 0.16, p =
0.30).
Distance to the nearest island was significantly different among islands (Chi
square test; x2 = 14036.88, p ≤ 0.001) and significantly correlated with size of the
island (Pearson correlation test; r = 0.90, p ≤ 0.01) (Table 5.5). Daily and farthest
daily movement was significantly correlated with size of the island (Pearson
correlation test; r = 0.88, p ≤ 0.01 and r = -1.00, p ≤ 0.01). There was also a significant
correlation between daily and distance between island (Pearson correlation test; r =
0.61, p ≤ 0.01) and between farthest daily and the distance (Pearson correlation test; r
= 0.95, p ≤ 0.01). Annual and farthest annual movement were showed a negative
significant correlations with both size of island and distance between island (Pearson
correlation test; r = -0.99, p ≤ 0.01, r = -0.79, p ≤ 0.01, r = -1.00, p ≤ 0.01, and r = -
0.88, p ≤ 0.01, respectively).
5.4 DISCUSSION
Many authors define dispersal as movement away from one location to another
location, being it to move away from natal sites, to look for suitable breeding
opportunities, to find an unoccupied habitat, or to either avoid a conflict or
competition with other conspecifics (van Dyck & Baguette 2005; Grant 1978;
Greenwood 1980). In this study, dispersal was defined as movement of young
Komodo dragons to the adjacent valleys or adjacent islands in term of accession of
gene flow exchange between local populations. The aim of this study was to measure
movement patterns and site fidelity in the early life stages of Komodo dragons. Based
on three years of mark recapture data, this study has documented the first information
on dispersal ability of Komodo dragons during early life stages. In general, movement
58
Distance to
Area Size Nearest Slope Vegetation
Location the nearest
(km2) Peak (m) degree type
peak (m)
Loh Sebita 5.81 181.24 889.93 4.33 2.82
Loh Liang 8.94 181.24 889.93 4.33 2.82
Loh Lawi 10.03 231.82 954.73 3.46 2.61
Loh Wau 0.83 301.67 909.27 9.11 2.60
Loh Buaya 5.50 126.81 913.46 3.61 2.78
Loh Baru 5.48 253.11 597.82 7.84 2.12
Loh Tongker 2.64 272.10 533.67 6.74 2.14
Loh Dasami 3.54 326.76 830.43 5.22 1.00
TOTAL/MEAN 5.35 234.34 814.90 5.58 2.36
During the study, there was no record of within trip recapture or resight on
hatchlings, thus daily movement of hatchlings could not estimated. No captured
hatchling were recaptured nor even resighted after captured, either from nest or from
traps, during the mark-recapture study session. This study showed a low degree of
recapture opportunities for hatchlings. This might be due to arboreal tendency of
hatchlings and their tendency to stay in the same tree for a long period of time
(approximately 9 days; see chapter IV).
After a year, one hatchling was recorded did not move at all from its first
capture location. This hatchling was recaptured at the same location one year after its
first capture. In contrast, another hatchling displayed long distance movement during
the first year of its life. Radiotracked hatchlings (see chapter IV) displayed a relatively
slow and short distance movement during their first month after emergence. However,
it is difficult to conclude that hatchling Komodo dragons exhibit either site fidelity or
long distance natal dispersal during their early stage of life.
During their early life stage, Komodo dragons exhibit poor long distance
movement ability. Juveniles did not move more than 2000 m, either for daily
movement or for annual movement, and there was no record of any movement to
adjacent valleys, or to adjacent islands. In contrast, most juveniles displayed short
distance movement (<500 m). This result is similar to short distance dispersal pattern
in juveniles of the Tasmanian Snow skink (Niveoscincus microlepidotus) (Olsson &
Shine 2003). Sutherland et al. (2000) documented that dispersal in juvenile birds and
mammals often involve short distance movement, while long distance was
uncommon.
60
During the three years of the study, there was no evidence of juvenile Komodo
dragon dispersing to adjacent islands. Even though Komodo dragons are able to swim,
the ocean may be a main geographical barrier preventing dispersal between adjacent
islands (Ciofi 2002). Manel et al. (2003) suggested that landscape features, i.e.
mountains, stream, and ocean, might influence the pattern of gene flow and population
structure.
In addition, this study suggested that movement and fidelity in hatchling and
juvenile Komodo dragons may have an implication to the gene flow. Ciofi (2002)
described genetic variability among island populations for Komodo dragon. High
degrees of similarity were found between Rinca and nearby islands (Nusa Kode and
west coast of Flores) whereas, populations from Komodo and Gili Motang showed
significant differentiation from other island populations.
GENERAL DISCUSSION
An important finding from this study was that, during the early stage of their
life, Komodo dragons have small activity areas with a high degree of overlap, and do
not move long distances. Hatchlings displayed a linear fashion of movement from
their nest that appeared to underpin a deliberate type of dispersal behavior to move as
far as possible from their nesting sites. Juveniles, in turn, displayed their ability to
explore their environment and occupied larger areas than hatchlings. Movement in
hatchlings and juveniles are important in term of finding suitable habitat to avoid
competition with larger conspecifics, expanding their distribution and maintain
genetic variability of the population (Lebreton et al. 2003; Pough et al. 2001; Sarno et
al. 2003; Sutherland 2000; Trakhtenbrot et al. 2005).
Forest coverage plays an important role for both hatchling and juvenile
Komodo dragons as shown by the extensive use of arboreal habitats. Both hatchling
and juvenile Komodo dragons displayed their preference for a particular tree (i.e.
Tamarindus indica) and particular terrestrial setting (i.e. rock crevices) for shelter.
This tendency showed the importance of trees during their early life-stage particularly
63
Given their limited dispersal abilities, this age class of Komodo dragons are in
a precarious situation, and vulnerable to both natural and anthropic threats. Ciofi
(2002) mentioned that potential natural threats included active volcanoes adjacent to
the habitat. Ciofi and de Boer (2004) noted that decline in the extant population of
Komodo dragons was influenced by habitat fragmentation and poaching activities. It
has been documented that natural and anthropogenic factors are the major causes of
species extinction within last decades (Kull et al. 2006).
6.2 RECOMMENDATIONS
Dispersal in animals plays an important role in their population dynamics and genetic
variability (Trakhtenbrot et al. 2005; Grant 1978). Dispersal was suggested as a
mechanism to maintain small populations and stabilized the overall population
through recolonization and as a source of genetic variability (Reed et al. 1998).
Blouin-Demers and Weatherhead (2002) described how movements had a significant
64
impact to the gene flow mechanism in black rat snakes (Elaphe obsoleta) which was
represented by the mating system between male and females from different
subpopulations.
The absence of dispersal may put small populations at risk of local extinction
due to inbreeding depression and genetic drift (Madsen et al. 1996; Konuma et al.
2000). Limited inter-island dispersal over long periods of time may result in insular
alteration (Sinclair 1998). Losos (2004) documented that the adaptation of the Greater
Antillean anoles was correlated to isolation by insular and within island mechanisms.
Ciofi (2002) has documented the genotypic differences among insular populations,
particularly for the most isolated island, Gili Motang. This sub-population is the most
severely at risk (Jessop et al. 2006 in press). Limitation in genetic variation due to
insular adaptation has the potential to harm the extant populations of Komodo dragons
which are declining due to natural or anthropic causes.
Finally, further long-term studies are needed to address the lack of detailed
knowledge on the spatial ecology of this vulnerable species. Detailed ecological
studies may provide a critical basis for management and conservation planning (Webb
& Shine 1977). In addition, dispersal models can be used to predict which group or
populations is likely to be the most vulnerable to disturbance and allow managers to
test the merits of alternative habitat conservation strategies (Sutherland et al. 2000).
CHAPTER VII
CONCLUSION
This study has documented that, during their early life stage, Komodo dragons have
small activity areas with a high degree of overlap, and do not move long distances.
There was no record of either hatchlings or juveniles performing a long distance
movement to the adjacent valleys, neither to adjacent islands. Movement in Komodo
dragons, particularly young, was constrained by landscape features.
This study also showed that forest coverage plays an important role for both
hatchling and juvenile Komodo dragons. Both hatchling and juvenile Komodo
dragons displayed a preference for open forest with a particular tree (i.e. Tamarindus
indica) and particular terrestrial setting (i.e. rock crevices) as shelter. This showed the
importance of trees during their early life-stage particularly in avoiding cannibalism
by adult Komodo dragons.
Since the dispersal of this species during their early life stage is limited, it may
put the extant population of Komodo dragon at risk of local extinction due to
67
inbreeding depression and genetic drift. Several feasible options to manage the
population should be considered before the population becomes too small for
conservation options. Protecting remain suitable habitats is the most important aspect
for the conservation of this species. Augmentation of this endangered species could be
considered by managers as one of conservation strategies in order to maintain genetic
variability and population viability. Finally, further long-term studies, especially on
the hatchling and juvenile stages, are needed to address the lack of detailed knowledge
on the spatial ecology of this vulnerable species.
68
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