Download Malay Civet Population Project

IL242 Ecology of the top mammalian predator in the forests of Sulawesi, Indonesia
Dr Adrian Seymour
The Malay civet Viverra tangalunga is one of 35 species of small to medium sized carnivores
in the family Viverridae (Nowak 1999). This family, along with the mongooses which some
authorities now include within the Viverridae, consists of mostly omnivorous tropical forestdwelling animals that are often the most numerous members of mammalian rainforest
predator communities in Asia and Africa (Charles-Dominique 1978, Rabinowitz 1991, Heydon
& Bulloh 1996, Ray 1997, Colón 2002). However, despite their potential importance in Old
World rainforest ecosystems as predators, frugivores and pollinators, very little is known
about their ecology, largely due to their secretive nature and the inaccessibility of their
jungle habitats.
This project explores two crucial elements of Malay civet ecology: 1) population dynamics
(the study of how and why populations change over time) and 2) spatial organisation and
social behaviour (a long-standing hot topic in carnivore ecology because of the great range
of social behaviours displayed within the order Carnivora e.g. Macdonald 1983, Bekoff et al.
1984). Each of these two topics including the findings of the project up to date will be
discussed separately in sections below. But first the study needs to be put into geographical
context – the project study site on Buton Island, southeast Sulawesi, is out the Malay civet’s
original range and is ecologically quite different.
The Malay civet on Buton
The Malay civet is found on the Malay Peninsula, Sumatra, Borneo, Sulawesi, the Molluccas
and the Philippines. On Buton Island, Sulawesi and nearby islands both the plants and
animals are quite different to those in other parts of the Malay civet’s range. Notably, on
Buton the Malay civet shares its range with a much more limited community of mammalian
predators. This is potentially important, because in other habitat types, including many
temperate habitats and tropical savannahs, intraguild predation and competition has been
shown to be very important in affecting the ecology of mammalian carnivores (e.g.
Palomares et al. 1997, Palomares & Caro, 1999, Fedriani et al. 2000). On Buton Island the
Malay civet is one of probably only two species of wild mammalian carnivores: although the
presence of a second species, the endemic Sulawesi palm civet Macrogalidia
musschenbroekii has not yet been confirmed, many of the people living around the forest
have a name for a second species of civet matching the description of the Sulawesi endemic,
and so it is likely that the elusive Sulawesi palm civet is indeed present in the forests of
Buton Island. Feral domestic carnivores including both dogs Canis familiaris and cats Felis
catus are also present in parts of the forest near villages and roads. In contrast, west of the
Wallace line on Borneo, Sumatra and the Malay Peninsula the Malay civet shares its habitat
with a much bigger suite of mammalian predators. On Borneo, for example, the Malay civet
shares its rainforest habitat with 22 other species of mammalian carnivore including eight
species of civet (Nowak 1999).
Not only do conditions on Sulawesi and Buton differ ecologically from elsewhere in the
Malay civets range, because of their isolation the civets themselves may be genetically
distinct from elsewhere in their range. Unlike the big islands of Borneo, Java and Sumatra
which were attached to mainland Southeast Asia via the Sunda shelf during the last ice ages,
Sulawesi and surrounding islands including Buton remained separated from both the Asian
and Australian continents by deep ocean. The Malay civet was introduced to the Wallacea
region including Sulawesi and the Molluccas from Borneo by human agency. It’s not known
when they first arrived on Buton; it could have been as far back as prehistoric times, brought
along with the expansion of humans into Sulawesi, as early as 7000 BC (Whitten et al. 1987)
or as recently as the seventeenth century when European demand for civet musk for the
medical and perfume industries may have triggered the transport of Malay civets to new
islands including Sulawesi (Dannenfeldt 1985). Either way, there are perceptible differences
in the sizes of Malay civets on Buton from those on Borneo and the Malay Peninsula
(Jennings et al. 2006, Jennings et al. 2009), but whether or not these differences have a
genetic component is unknown.
Population dynamics of mammalian carnivores
Probably the most famous and long-running of any published time-series of a mammalian
carnivore is Elton and Nicholson’s (1942) data on the number of pelts of Canadian lynx Lynx
canadensis obtained by the Hudson’s Bay Company each year between 1821 and 1912.
These data revealed a dramatic ten-year cycle in the fluctuation of lynx numbers, which
were seen to track the abundance of snowshoe hares Lepus americanus. Population cycling
is easily generated in model predator-prey systems and arises due to “delayed density
dependence”, the time lag between the effect of the predator abundance on prey
populations and vice versa (e.g. Begon et al. 1990). The lynx population was tracking the
abundance of their main prey, the snowshoe hares, though it seems the hares themselves
were being affected by the quality and availability of their plant food supply (Sinclair et al.
1988) rather than predation by lynx. Populations of mammalian carnivores are likely to
track prey abundance when they specialise on particular prey species and if those prey
themselves are prone to fluctuations. This situation is common for specialist predators such
as stoats and weasels in boreal regions (e.g. Hanski et al. 1991, Hellstedt et al. 2006). The
situation for generalist mammalian Carnivores is less clear, since generalist species can easily
switch to other prey types one prey type becomes scarce. This is especially true of
omnivorous members of the order Carnivora such as foxes, raccoons, badgers, and many
species of civet including the Malay civet, as they may even switch from animal to plant
material such as fruit when animal food becomes scarce. For these carnivores, other factors
are likely to play an important role in shaping population dynamics.
Disease has shown to be an important factor influencing the abundance of mammalian
carnivores. Red fox Vulpes vulpes populations in Europe have been shown to crash
dramatically in response to outbreaks of sarcoptic mange Sarcoptes scabei infection
(Lindström, 1994; Soulsbury et al. 2007). The populations of a range of mammalian
carnivores in the Serengeti-Masai Mara ecosystem including lions Panthera leo and spotted
hyenas Crocuta crocuta have been shown to be profoundly affected by an outbreak of the
canine distemper virus in 1994 (Roelke-Parker et al. 1996). Civets may be affected by a
number of diseases including canine distemper (Ikuyo et al. 2009) and the SARS coronovirus
(Bell et al. 2004), though it is not known what roles these pathogens play on civet population
dynamics.
Population dynamics of Malay civets on Buton
The numbers of adult Malay civets caught each year has fluctuated from 9 in 2004 to 17 in
2008 (Figs. 1 and 2).
Fig. 1. Minimum number alive count of civets caught in the Buton study area by year
25
Juveniles
Subadult Male
Subadult Female
Adult Male
Adult Female
20
Total adults
Total adults + subadults
N
15
10
5
0
2003
2004
2005
2006
2007
2008
2009
2010
Fig. 2. Abundance estimates ± 95% CL using closed capture-mark-recapture models
60
Civet abundance
50
40
30
20
10
0
2003
2004
2005
2006
2007
2008
2009
2010
The low abundance of civets in 2004 coincided with behavioural signs of disease
(unsteadiness on feet, and slow and erratic departure when released from traps), and low
adult body mass (Fig. 3). Although data from the civets caught in 2011 has not yet been
processed, initial indications suggest that abundance is again low and adults also show low
body masses and signs of disease. Further investigation clearly needs to examine blood
tissues for signs of disease including canine distemper. It is interesting that the first signs of
disease since 2004 are happening in 2011 after a sustained run of high civet abundance from
2007-2010. Could it be possible that a disease and civets are showing host-pathogen cycles?
Fig. 3. Mean civet body mass (kg) ± 95% CI
One of the clearest observations from the capture-mark-recapture is that females are much
more likely to be recaptured between years than males (Fig. 4). It is not clear whether this is
due to higher female survival or increased female philopatry, but either way, females remain
in the study site for longer. Between year recapture rates are much higher in females than
males for young age classes (Juvenile, Subadult, Young adult), but this pattern is reversed in
the
age classhistory
(Fig. 5).for male and female civets from 2003 to 2010.
Fig.adult
4. Capture
Males
Females
2003
M1
M2
M3
M4
M5
M6
M7
M8
M9
M10
M11
M12
M13
M15
M16
M17
M18
M19
M20
M21
M22
M41
M23
M24
M25
M26
M27
M28
M29
M30
M31
M32
M33
M34
M36
M37
M38
M39
M40
M?
M42
M43
M44
M45
M46
M47
M48
M49
M50
M51
M52
M53
M54
M57
M58
M59
M60
M61
M62
M63
M64
M65
Ken
LR
Beng2 !L
Neil
R
Rob
R
Ady
LR
Frank
L
Sardin R
Bule
R
Jim Lad R
Matt
L
Steve
R
Bintang L
Martin
Laduni
Floyd
Koti
Stuart
Marcus
Paul
Zuni
Djembe
Extra Joss
Mark
Wahyu
Dave
Pete
Arthur
Mr nanda
CJ
Elvis
The Gimp
Alex
Ed
Angur
Alan
Juliadin
Bruce
Scott
Itchy
no name
Sudi
Dan
Sam
Harry
Daddy
Colin
Storm
Bob Marley
Baz
James
Lalra
Ashby
Ned
Frank2
Clive
Jeff
Badger
Orio
Bernie
James Bond
Big Ed
Marmite
2004
LR
!LR
2005
2006
LR©
L©
2007
2008
2009
2010
2003
F1
F2
F3
F4
F5
F6
F7
F8
F9
F10
F11
F12
F13
F14
F15
F16
F17
F18
F31
F19
F20
F21
F23
F24
F25
F26
F27
F28
F29
F30
F32
F33
F35
F40
F41
F42
F44
F46
F47
F48
F49
F51
F52
!R
L
R
R
LR
R
R
R
LR
R
R
R
!LR
LR
R
LR
LR
LR
LR©
!LR
LR©
LR
LR
R
LR
!R
LR
LR
LR
L
LR
LR†
LR
!R©
LR
LR
LR
LR
LR
LR©
L
R
Rm
LR
LR©
LR
LR©
LR
LR
LR©
LR
LR
LR
LR©
LR
LR
LR
LR©
Penelope!LR
Vicky
L
Anna
R
Mandy LR
Nina
L
Big MamaR©
Jenny
!R
Sarah
R
Laura
L
Cindy
!R
Jen
Magdalena
Elen
Rachel
Esti
SJ
Sahudin
Christina
Kit
Wawa
Uni
Dewi
Jamie
Noeleen
Kirsty
Aisla
Mary
Tia
Claudia
Maria
Tine
Liv
Foxy
Alex2
Jamie2
Bintang2
Peggy
Ladune2
Neo
Shaka Khan
Ruby
Fiona
Tamsin
2004
2005
2006
2007
2008
2009
2010
L
L
LR
!R
L
!LR
LR
!L
LR
!R©
!R©
!R©
L
L©
LR
!LR
LR
LR
L
LR
L
R
LR
L©
LR
LR
LR
LR
LR
LR
LR
!R
R
!R
!R
R
LR
!R
LR©
LR
LR
L©
LR
R©
LR
LR
!LR©
!L
L
LR©
LR
LR
LRR
!LR
LR
LR©
LR
R
LRm
m
!L©
!R
!LR
L
L
L
L
!L
!m
LRm
!L
LRm
Rm
Rm
Rm
Rm
Rm
LRm
LR
R
L
Rm
Rm
Rm
Rm
!Rm
Rm
Rm
R
R
Fig. 5. Percentage between year recapture for male and female civets.
90
9
80
% recaptured
70
8
60
17
50
9
Males
Females
14
40
30
20
9
12
10
7
0
Juvenile
1 2
Subadult
Young
Adult
Adult
Age at first capture
Social organisation of carnivores
Old Adult
Many members of the order Carnivora form complex social groups showing cooperative
behaviour, with large species such as wolves Canis lupus and lions Panthera leo clear
benefiting from living in groups by cooperative hunting and in the case of lions by
cooperative defence (female defence of cubs from infanticidal males), and other species
such as meerkats Suricata suricata and banded mongooses Mungos mungo benefiting from
group vigilance and alloparental care (Macdonald 1983, Bekoff et al. 1984). But there are
many other species from other Carnivoran families, including Canids (e.g. red fox Vulpes
vulpes), Mustelids (e.g. European badgers Meles meles), Procyonids (e.g. Kinkajou Potos
flavus) and Viverrids (e.g. African palm civet Nandinia binotata), where the adaptive
significance of group living is much less clear, since these species usually hunt alone, but will
often form groups at other times (Charles-Dominique 1978, Baker et al. 1998, Kays &
Gittleman 2001). Explaining why these groups are formed has been much harder, and many
theories have been put forward. These include the resource dispersion hypothesis (the
passive tolerance of additional group members due to the abundance and distribution of
food resources: Macdonald 1983, Johnson et al. 2002), the territory inheritance hypothesis
(inclusive fitness benefits from allowing offspring to stay in the natal territory when dispersal
costs are high: Lindström 1986) and the “anti-kleptogamy” hypothesis (males space
themselves to monopolise as many females as possible, defending these territories to
ensure exclusive access to several females: Roper at al. 1986). Understanding the ecological
conditions that allow multiple stable territories to overlap thus opening the possibility of
repeated interaction between individuals is a crucial step in understanding how group living
and sociality may have evolved.
Most rainforest Viverrids are considered to be solitary animals, who only come together to
mate or when mothers are raising young offspring (Nowak 1999). However, our radio
tracking research has shown that the Malay civets on our study site have frequently
overlapping ranges (both within and between sexes: Fig.6), and that meetings between
collared civets occur quite frequently – any given civet is likely to meet a neighbour at least
once per night. Sometimes civets spend long periods in close proximity to each other.
Females being the more sedentary, may form long-lasting relationships, however, there
seems to be no difference in the frequency or duration of male-male, male-female or
female-female interactions, though more analysis needs to be done.
M28 (Mr Nanda) Transient adult
#
14.5 ha
Detected 50% of days (n=10)
#
2005 Movement ranges
#
#
#
#
M01 (Ken) Resident adult
57.1 ha
Detected 92% of days (n=24)
#
#
#
#
1 km
#
#
#
Lapago camp
M26 (Pete) Transient adult
43.9 ha
Detected 69% days (n=13)
#
F07 (Jenny) Resident adult
45.5 ha
#
Detected 95% of days (n=22)
#
#
#
#
F09 (Laura) Resident adult
49.1 ha
Detected 94% of days (n=34)
#
#
M01 (Ken) Resident adult
53.1 ha
Detected 58% of days (n=33)
#
#
#
1 km
2006 Movement ranges
#
#
Lapago camp
F17 (Sahudin) Resident adult
7.7 ha
Detected 67% of days (n=21)
F14 (Rachel) Transient
adult
29.3 ha
2007 Movement ranges
S
#
S
#
#
M31 (Gimp) Resident
adult
37.0 ha
M34 (Angur) Resident adult
46.6 ha
#
#
#
F29 (Claudia) Resident
adult
26.9 ha
#
#
#
M40 (Itchy)
Resident
adult #
90.6 ha
#
#
#
#
M25 (Dave) Resident
adult
48.7 ha
F05 (Nina) Resident adult
42.5 ha
#
#
#
#
F16 (SJ) Resident
adult
43.7 ha
F07 (Jenny) Resident
adult
55.1 ha
F26 (Aisla)
Resident
1
adult
km
37.3 ha
#
#
#
#
#
#
#
#
#
2008 Movement ranges
#
F26 (Aisla) #
Resident adult
65.0 ha
Detected 100%
of days (n=23)
#
#
#
#
F35 (Foxy)
Resident subadult
15.1 ha
Detected 100% of
days (n=11)
M40 (Itchy)
Resident
adult
66.9 ha
Detected
60% of days
(n=25)
#
#
#
M50 (Baz)
Transient adult
38.2 ha
Detected 100%
of days (n=20)
#
#
#
#
Lapago
camp
M46 (Daddy) Resident adult
31.8 ha
Detected 46% of days (n=24)
F17 (Sahudin)
Resident adult
13.0 ha
Detected 52% of days
(n=27)
1
km
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