IJERD – International Journal of Environmental and Rural Development (2013) 4-2

Research article
erdComparative Reproductive Behavior of α-Male, β-Maleand Subordinate Male Timor Deer (Cervus timorensis
Blainville) Raised under Captivity
DAUD SAMSUDEWA*
University of Diponegoro, Semarang, Central Java, Indonesia
Email: daudreproduksi@gmail.com

SEVERINO S. CAPITAN
University of The Philippines Los Baños, College, Laguna, Philippines

CESAR C. SEVILLA
University of The Philippines Los Baños, College, Laguna, Philippines

RENATO S. A. VEGA
University of The Philippines Los Baños, College, Laguna, Philippines

PABLO P. OCAMPO
University of The Philippines Los Baños, College, Laguna, Philippines
Received 16 December 2012

Accepted 10 June 2013(*Corresponding Author)

Abstract Timor deer were used in an experiment to study the comparative behavior of αmale, β-males and subordinate male raised in captivity. Twelve males (4.25 years old) and24 females (3 to 4 years old) were randomly assigned into three separate cages following
the ratio of 4 males: 8 females. Kruskal-Wallis H test of non-parametric analyses weredone for aggressive, libido and mating behaviors. The α-male was the dominant male interms of aggressive, libido and mating behaviors. Subordinate male showed the highest
frequency of wallowing to reduce stress, and running around for expressing libido.
Successful mating was mostly exhibited by α-male, Subordinate2 (S2)-male did not show
actual mating. The establishment of dominance hierarchy resulted to the manifestation ofdifferences reproductive behaviors.
Keywords reproductive behavior, male Timor deer, captivity

INTRODUCTION
The Timor deer (Cervus timorensis Blainville) is a medium-sized cervid with 60 kg mature bodyweight and 1.00-1.10 m shoulder height (Jacoeb and Wiryosuhanto, 1994). It is currently being
reared in farms in many countries including New Zealand, Australia, Indonesia, Mauritius, New
Caledonia, China, Korea and Russia (Semiadi and Nugraha, 2004).
The productivity of deer farming in Indonesia is still low. The study of Daningsih (2005)
showed that annual population growth of deer was only 0.169 at Kelompok Pemangkuan Hutan(KPH) Bunder, 0.25 at Kulon Progo and 0.17 at KPH Jonggol. The low of the annual population
growth suggests that the productivity of deer farming in captivity can be improved if some key
aspects of management such as feeding, mating, aggressive male behavior, maternal care andnervousness tendency are addressed. Asher et al., (1996) claimed that intensive deer farming withpoor management practices, such as low of the feed quality and high social stress, may lead to poor
cervid reproduction health. This is corroborated by Moberg (1991) who stated that behavioral stresshas adverse effects on the reproduction system of both males and females. Reproductive success is
one of the parameters to monitor stress. Animals that suffer from stress often fail to reproduce
successfully.

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IJERD – International Journal of Environmental and Rural Development (2013) 4-2

Timor deer's develop a linear dominance hierarchy consisting of α-males, β-males andsubordinate males through aggressive behavior. Social stress including those caused by conflicts
related to dominance hierarchy has relatively consistent effects on reproductive behavior. This ismanifested by changes in male sexual behavior (Blanchard et al., 2002).
OBJECTIVES
Therefore, in order to develop better management practices to maximize reproductive capacity ofdeer farms, it is necessary to study reproductive behavior of α-male, β-male and subordinate male
Timor deer raised under captivity.
METHODOLOGY
The study was conducted at H. Yusuf Wartono Timor deer captive breeding, Gondosari, Gebog,
Kudus, Central Java. The reproductive behavior observation was carried out for 43 days starting
August 1, 2011 until September 12, 2011. Twelve (12) males (51 ± 6 months old; 68.29 ± 8.41kg
body weight and in same antler stages), and twenty-four (24) females (3-4 years old 40-60 kg bodyweight with normal estrus) were used. Other materials include 3 communal cages (23.5 x 21.5 m)
and ethogram table.
Observation for reproductive behavior was done with behavior sampling (Martin and Bateson,1993). The observation was done by manual recording using a record book and an ethogram table.
Time, frequency, interval, duration and sequence of every behavior were recorded. The data was
supported by audiovisual recorder. Data gathering of reproductive behavior was focused on anumber of aspects including libido behavior, aggression and mating behavior.
Kruskal-Wallis H test of non-parametric analysis was done for comparing α, β andsubordinate male Timor deer in terms of libido, aggressive and mating behavior.
RESULTS AND DISCUSSIONS
Aggressive behavior: Aggressive behavior of male Timor deer was expressed by wallowing,
walking with head up, shouting, rubbing antler, expression of threat, pushing, actual fighting,fleeing and climbing. Aggressive behaviors are commonly expressed when more than one malestart to approach an estrus female, α-males are the ones commonly displaying these behaviors,
especially when β-males or subordinate males are approaching. From the time of actual fighting
and incidence of fleeing, dominance hierarchy was established.
Average values of each aggressive behavior of α, β and subordinate male Timor deer's areshown in Table 1. The result of Kruskal-Wallis H test showed significant differences for all
aggressive behaviors among the different hierarchy of male Timor deer. The α-male as thedominant male had mostly shown dominancy in all of the behavior parameters except in frequency
and interval of wallowing and fleeing. The highest frequency and interval of wallowing were
exhibited by S2-male, which could be an attempt of the animal to reduce stress. S2 male had thehighest frequency and longer interval of wallowing, but shorter duration because he was disturbed
by more dominant males when started. On the other hand, β-male exhibited the highest frequencyof fleeing because of he was the opponent of α-male. The β-male also exhibited the longest intervalof fleeing because in addition to fighting with α-male, in between also fight with S1 or S2-male andexhibited winning.
Indeed, shouting bouts (shouting in between two periods of shouting) were only done by αmales as a sign of dominance. Wallowing is commonly followed by rubbing antler or expression of
threat. In these behavior α-male still shown the most dominant male except for interval of
expression threat. In terms of pushing and actual fighting, α-male also shown dominancy infrequency, duration and interval. Actual fighting was longer time between α and β-male comparewith other male. Climbing as one sign of dominancy, α-male was shown the most frequent maleexhibited this behavior with longest duration and interval. The α-male exhibited climbing everyday© ISERD
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IJERD – International Journal of Environmental and Rural Development (2013) 4-2

and the males usually disturbing other males when they exhibited this behavior.
Table 1 Average values of each aggressive behavior (43 days observations) of male Timor
deer
Behavior
Wallowing- Duration, seconds- Frequency, counts- Interval, minutes
Walking with Head Up- Duration, seconds- Frequency, counts- Interval, minutes
Shouting Bouts- Duration, seconds- Frequency, counts- Interval, minutes
Rubbing Antler- Duration, seconds- Frequency, counts- Interval, minutes
Expression of Threat- Duration, seconds- Frequency, counts- Interval, minutes
Pushing- Duration, seconds- Frequency, counts- Interval, minutes
Actual Fighting- Duration, seconds- Frequency, counts- Interval, minutes
Fleeing- Frequency, times- Interval, minutes
Climbing- Duration, seconds- Frequency, counts- Interval, minutes

α-maleβ-maleS1-maleS2-male

χ2

69.56
1.18
43.15

67.50
1,06
26.60

63.97
1,28
30.58

56.26
1,93
134.72183.94*72.02*
114.10*13.70
3.50
173.3611.65
1.42
127.09

5.61
0.63
36.81

0.00
0.00
0.00

374.69*404.36*241.47*

33.49
4.08
151.69

0.00
0.00
0.00

0.00
0.00
0.00

0.00
0.00
0.00

39.06
5.26
148.56

16.27
0.60
28.55

4.68
0.15
1.64

0.17
0.01
0.00

374.44*400.83*407.85*5.76
4.22
159.314.66
1.57
170.47

2.33
0.64
35.65

0.29
0.07
0.00

353.30*415.32*250.80*

6.14
3.92
162.154.95
1.61
156.53

2.57
0.68
40.78

0.47
0.11
0.37

348.38*402.98*247.04*229.72
3.55
163.64247.01
1.96
206.17204.091.39
88.69

107.360.68
38.41

261.66*342.14*156.82*

0.00
0.00

1.82
204.08

1.32
71.29

0.69
41.14

341.99*189.89*13.61
3.00
143.27

11.31
1.33
75.06

6.02
0.64
26.41

1.50
0.13
0.00

317.80*365.55*231.32*

Legend: α-male (The most dominant male); β-male (The second dominant male); S1-male (Subordinate 1 male); S2-male(Subordinate 2 male)

Libido behavior: Libido behavior of male Timor deer was expressed by wallowing, roaring,seeking female, crowning, flehmen, spraying of urine, rutting, climbing and running around.
Libido behavior was shown by male as expression of mating desire. In this present study, duration,
frequency and interval of each libido behavior was measured. The most behavior exhibited by male
Timor deer was seeking female (6.55 counts/day) followed by flehmen, spraying urine rutting andclimbing (6.40, 6.32, 4.88 and 4.31 counts/day, respectively). Average values of each libido
behavior of α, β and subordinate male Timor deer's are shown in Table 2.
The result of Kruskal-Wallis H test showed significant differences (P<0.05) for all libido
behavior among the different hierarchy of male Timor deer. The α-male as the dominant male had
mostly shown dominancy in all the libido behavior parameters except running around. The highest
frequency and longest duration and also interval of running around was exhibited by S2-male,which might be an attempt to express libido.
The α-male has shown dominancy for wallowing and spraying urine (most frequent and also
longest duration and interval). Spraying urine and wallowing could be related to release of odors
and attract the estrus female, that activity will be repeated when the estrus female shown nonreceptive behavior prior to mating. Libido behavior of α-males was found to be related with scentmarking (urinary spray and rutting), which was more common compared with other males. Roaring
was commonly done by α-males to display their dominancy. Blanchard et al., (2002) reported that
dominant male sugar gliders (Petaurus breviceps) display frequent scent marking behavior as partof their libido behavior.The α-male also showed dominancy on crowning behavior, which could be© ISERD
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IJERD – International Journal of Environmental and Rural Development (2013) 4-2

included as a sign of dominance and an act of attracting female. Flehmen is one of the interesting
libido behaviors of Timor deer's. It is an important sign signaling the start of a mating behavior.
Flehmen is done to facilitate exposure of the vomeronasal gland for a scent or pheromone(Messang-Nallley, 2006). Climbing was also mostly done by α-male. When other males exhibited
this behavior, α-male would disturb them. This is the reason for shorter duration and interval of
climbing in other males.
Table 2 Average values of each libido behavior (43 days observations) of male Timor Deer
Behavior
Wallowing- Duration, seconds- Frequency, counts- Interval, minutes
Roaring Bouts- Duration, seconds- Frequency, counts- Interval, minutes
Seeking Female- Duration, seconds- Frequency, counts- Interval, minutes
Crowning- Duration, seconds- Frequency, counts- Interval, minutes
Flehmen- Duration, seconds- Frequency, counts- Interval, minutes
Spraying Urine- Duration, seconds- Frequency, counts- Interval, minutes
Rutting- Duration, seconds- Frequency, counts- Interval, minutes
Climbing- Duration, seconds- Frequency, counts- Interval, minutes
Running Around- Duration, seconds- Frequency, counts- Interval, minutes

α-maleβ- maleS1- maleS2- maleχ2

252.403.24
146.21225.781.40
110.65139.98
0.73
21.58

37.64
0.20
0.00

345.92*359.41*233.78*38.87
4.41
155.73

0.00
0.00
0.00

0.00
0.00
0.00

0.00
0.00
0.00

228.82
4.06
151.57

194.01
1.59
154.56102.93
0.79
86.39

17.71
0.11
3.37

335.81*384.72*170.91*61.51
2.81
170.96

16.76
0.43
23.82

6.08
0.16
7.44

0.00
0.00
0.00

366.77*388.85*365.40*8.06
4.13
156.666.38
1.53
149.81

3.02
0.69
72.96

0.21
0.05
3.37

348.34*399.34*184.10*

11.98
4.04
151.2110.14
1.50
143.78

5.07
0.68
64.14

0.96
0.10
0.00

317.07*388.25*
195.76*41.13
4.19
156.90

13.48
0.57
27.13

2.68
0.12
2.74

0.00
0.00
0.00

393.76*404.09*403.62*

11.86
3.33
145.53

5.70
0.63
25.82

2.66
0.29
12.58

0.51
0.06
4.10

294.60*329.22*307.65*

0.00
0.00
0.00

0.00
0.00
0.00

45.95
0.41
24.18

129.901.58
145.31331.10*373.43*195.57*

Legend: α-male (The most dominant male); β-male (The second dominant male); S1-male (Subordinate 1 male); S2-male(Subordinate 2 male)Mating behavior: In the present study, mating behavior of male Timor deer was expressed byfollowing, sniffing, kissing, flehmen, kicking, nudging, mounting, erection, intercourse, ejaculation,refractory and bisexual. The highest frequency of mating behavior of male Timor deer was
manifested in following (6.91 counts/day) followed by sniffing and kissing (6.18 and 5.84
counts/day, respectively). Average values of each mating behavior of α, β and subordinate male
Timor deer's for duration, frequency and interval are shown in Table 3.
The result of Kruskal-Wallis H test showed significant differences for all mating behaviors
among the different hierarchy of male Timor deer. The α-male as the dominant male had mostly
shown dominancy in all of the mating behavior parameter except bisexual. Bisexual only observedin the subordinate male. S2-males showed the highest frequency (0.71 counts/day), longest
duration (3.16 seconds) and longest interval (37.41 minutes) of bisexual behavior among the maledeer's. The Subordinate male Timor deer exhibited bisexual behavior when they cannot mate withfemale deer's due to intimidation of α-male. Mating behavior was observed to be more frequent in
α-males than other males. Subordinate males had the lowest frequency of mating behavior. Indeed,S2 males only exhibited following, sniffing, kissing and bisexual behavior (0.08, 0.03, 0.02 and0.71 counts/day). Blanchard et al., (2002) also reported that subordinate males of albino mice

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IJERD – International Journal of Environmental and Rural Development (2013) 4-2(Rattus norvegicus), deer mice (Peromyscus maniculatus) and lesser mouse lemurs (Microcebus
murinus) have the least frequent of the mating behavior.
Table 3 Average values of each mating behavior (43 days observations) of male Timor deer
Behavior
Following- Duration, seconds- Frequency, counts- Interval, minutes
Sniffing- Duration, seconds- Frequency, counts- Interval, minutes
Kissing- Duration, seconds- Frequency, counts- Interval, minutes
Flehmen- Duration, seconds- Frequency, counts- Interval, minutes
Kicking- Duration, seconds- Frequency, counts- Interval, minutes
Nudging- Duration, seconds- Frequency, counts- Interval, minutes
Mounting- Duration Inter, seconds- Frequency Inter, counts- Interval Inter, minutes
Erection- Duration, seconds- Frequency, times- Interval, seconds
Intromisian & Ejaculation- Duration, seconds- Frequency, times- Interval, seconds
Refractory- Duration, seconds- Frequency, times- Interval, seconds
Bisexual- Duration, seconds- Frequency, times- Interval, seconds

α-maleβ-maleS1-maleS2-maleχ2

229.685.11
111.57

95.72
1.22
72.80

49.63
0.50
36.43

4.88
0.08
3.23

326.96*351.55*217.04*9.96
4.87
113.75

3.64
0.92
67.01

1.68
0.39
26.78

0.13
0.03
3.23

353.15*281.83*243.24*13.55
4.65
118.15

4.80
0.84
62.10

2.14
0.33
20.92

0.10
0.02
0.00

365.10*372.71*
267.72*

6.82
4.51
116.16

2.00
0.69
39.97

0.90
0.27
16.83

0.00
0.00
0.00

383.80*327.54*298.17*

5.61
4.49
117.68

1.59
0.63
33.74

0.70
0.24
13.40

0.00
0.00
0.00

379.90*281.66*313.31*

17.69
4.47
114.64

4.86
0.55
21.44

2.11
0.21
10.41

0.00
0.00
0.00

382.44*386.38*355.73*20.28
4.20
114.12

5.84
0.43
7.58

1.64
0.09
3.11

0.00
0.00
0.00

331.45*403.67*416.05*37.99
4.20
112.14

7.64
0.24
1.27

2.42
0.09
3.11

0.00
0.00
0.00

422.22*422.33*446.74*4.67
4.20
112.28

0.88
0.22
1.27

0.28
0.08
0.26

0.00
0.00
0.00

393.19*425.15*454.14*212.504.19
112.49

39.41
0.22
1.27

13.11
0.08
0.26

0.00
0.00
0.00

407.73*425.16*454.14*

0.00
0.00
0.00

0.05
0.01
0.00

1.56
0.33
18.44

3.16
0.71
37.41

153.41*156.46*41.09*

Legend: α-male (The most dominant male); β-male (The second dominant male); S1-male (Subordinate 1 male); S2-male(Subordinate 2 male)Mating behavior of Timor deer was observed to occur mostly between 5 am and 8 pm. Timor
deers stopped mating activity when the temperature fell down below 21 0C, which commonlyoccurred after 8 pm. These findings were confirmed by Ismail (2008) in Ranca Upas, Indonesia
who reported the same observation. Timor deer still continue to mate under rain condition but will
look for shelter when temperature dramatically drops. But, in Ranca Upas this condition would
happen when temperature dramatically drops until around 16 0C. It was observed that Timor deer
started to rest from 8 pm. Almost all of them would stay and lay in canopy, only a small number
would ruminate outside of the canopy. Mating behavior of Timor deer is different compared withSambar deer (Rusa unicolor) in India. Savanth et al., (2011) reported that Sambar deer arenocturnal, therefore during day time mounting and intercourse could not be observed more thanonce, but other activities related to breeding like chasing females, sniffing and flehmen wereobserved many times. Territorial behavior, holding the head up high, fighting, spraying urine on its
own body and face were also observed a few times.

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CONCLUSIONS
The most aggressive males were α-males. The common sequence of aggressive behaviors was
expression of threat, followed by pushing, actual fighting and fleeing when one of the males lose
the fight. Libido behavior of male Timor deer was expressed by wallowing, roaring, seeking female,crowning, flehmen spraying of urine, rutting, climbing and running around; α-male was the mostactive in libido behavior. Subordinate male expressed libido through running around. Farmmanagement practices should be given emphasis as an important key for reproductive success. This
may include the provision of mud bath in the cage to support expression of normal behavior.
Further study on the grouping of the males in relation to the number of males in each cage must be
conducted. The possible influence of dominance hierarchy on the subsequent sex ratio of fawns
from captive breeding of Timor deer should also be studied.
ACKNOWLEDGMENTS
This research was funded by The Southeast Asian Regional Center for Graduate Study andResearch in Agriculture (SEARCA) and German Academic Exchange Service (Deutthscher
Akademischer Austausch Dienst e.V./DAAD), with SEARCA-DAAD Scholarship for graduate
study. In addition, thanks to Mr. Yusuf Wartono and Mr. Deka Hendratmanto, the owner of“Cervidae” captivity breeding, Kudus, Central Java, Indonesia for all the facilities used in thisresearch.
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