Kelelawar Pemakan Buah (Megachiroptera, Pteropodidae) Dari Gunung Bawakaraeng Sulawesi Selatan

THE FRUIT BATS (MEGACHIROPTERA, PTEROPODIDAE)
FROM BAWAKARAENG MOUNTAIN, SOUTH SULAWESI

ELLENA YUSTI

GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015

STATEMENT ABOUT THESIS, INFORMATION SOURCES,
AND ACT OF SPILLING OVER COPYRIGHTS*
By this writing I clarify that the graduate thesis The Fruit Bats
(Megachiroptera, Pteropodidae) From Bawakaraeng Mountain, South Sulawesi is my
own work under the supervisions of the advising committee and has not been
proposed for any institution. Copied information source of published and unpublished
writing of other author has been mentioned in the text and incorporated in the
references at the last part of this thesis.
By this writing I hand over the copyright of my thesis to Bogor Agricultural
University.
Bogor, February 2015

Ellena Yusti
NIM G352120151

RINGKASAN
ELLENA YUSTI. Kelelawar Pemakan Buah (Megachiroptera, Pteropodidae) Dari
Gunung Bawakaraeng Sulawesi Selatan. Dibimbing oleh BAMBANG
SURYOBORTO dan IBNU MARYANTO.
Kelelawar pemakan buah (Megachiroptera, Pteropodidae) berperan
penting dalam ekosistem sebagai penyebar biji dan polinator, umumnya ditemui
di wilayah perkebunan, hutan primer dan hutan sekunder. Keanekaragaman
kelelawar pemakan buah dipengaruhi oleh sumber pakan, tipe habitat, ketinggian
dan faktor-faktor lingkungan, yaitu curah hujan, angin, fase bulan yang
mempengaruhi aktivitas kelelawar dan jumlah individu yang tertangkap.
Penelitian kelelawar pemakan buah (Megachiroptera, Pteropodidae) di kawasan
Gunung Bawakaraeng, Sulawesi Selatan bertujuan untuk mengetahui komposisi
spesies, termasuk kategori umur dan status reproduksi kelelawar betina,
keanekaragaman jenis kelelawar pemakan buah dan preferensi habitat di berbagai
tipe habitat, serta untuk mengetahui pengaruh dari fase bulan terhadap jumlah
individu yang tertangkap.

Sebanyak 265 individu kelelawar pemakan buah didapatkan dengan
menggunakan perangkap jaring kabut (mist net) pada lima tipe habitat yang
berbeda, yaitu hutan sekunder (1200 m dpl), perkebunan campuran (1453 m dpl),
hutan pinus (1545 m dpl), hutan primer (2000 m dpl) dan gua (2200 m dpl) di
kawasan Gunung Bawakaraeng, Sulawesi Selatan. Kelelawar pemakan buah yang
didapatkan terbagi atas tujuh genus yang meliputi 10 spesies dengan kurva
estimasi menunjukkan 12 spesies yang berarti dua spesies belum tertangkap.
Komposisi kelelawar yang tertangkap meliputi Boneia bidens (159 individu;
60%), Thoopterus suhaeniahae (50 individu, 18.86 %), Rousettus celebensis (21
individu; 7.95%), Thoopterus nigrescens (11 individu; 4.16%), Eonycteris spelaea
(9 individu; 3.40%), Rousettus amplexicaudatus (6 individu; 2.27%), Dobsonia
viridis (4 individu; 2.27%), Styloctenium wallacei (2 individu; 0.75%), Dobsonia
exoleta (2 individu; 0.75%) and Cynopterus luzonensis (1 individu; 0.37%).
Analisis keanekaragaman Shannon-Wiener menunjukkan nilai tertinggi di
perkebunan campuran (1453 m asl) (H=1.80), sedangkan nilai terendah di hutan
pinus (1545 m dpl) (H=0.32). Nilai indeks dominansi tertinggi di perkebunan
campuran (1453 m dpl) (D=0.77) dan nilai dominansi terendah di hutan pinus
(1545 m dpl) (D=0.18). Komponen analisis utama menunjukkan hubungan antara
spesies dan tipe habitat. T. suhaniahae berkorelasi dengan tipe habitat terganggu,
yaitu perkebunan campuran (1453 m dpl) dan hutan pinus (1545 m dpl),

sementara itu T. nigrescens berkorelasi dengan hutan pinus (1545 m dpl). B.
bidens berkorelasi dengan tipe habitat hutan primer dengan aliran sungai (2200 m
dpl) dan R. celebensis berkorelasi dengan hutan sekunder (1200 m dpl).
Berdasarkan fase bulan, keseluruhan spesies ditemukan pada fase bulan baru,
bulan sabit, bulan setengah, bulan cembung dan bulan purnama. B. bidens
merupakan spesies yang ditemukan pada fase bulan purnama, namun dengan
jumlah individu yang sedikit.
Keywords: Chiroptera, kelelawar pemakan buah, distribusi, preferensi habitat,
fase bulan

SUMMARY
ELLENA YUSTI. The Fruit Bats (Megachiroptera, Pteropodidae) From
Bawakaraeng Mountain, South Sulawesi. Advisored by BAMBANG
SURYOBORTO and IBNU MARYANTO.
Fruit bats have an important role in ecosystem as seed dispersers and
pollinators. They commonly found in agricultural areas with cultivated plant,
secondary and primary forest that associated with food resources. The distribution
and diversity of fruit bats are influenced by food resources, habitat types,
elevation and environmental factors such as rainfall, wind, moon phases can all
affect number captured and bat activity. Fruit bats study in Bawakaraeng

Mountain aims to determine the fruit bats composition, including age catagorized
and reproduction status of female fruit bats, fruit bats diversity in each habitat
types, habitat preferences and relation between captured individual fruit bats with
the moon phases in Bawakaraeng mountain.
A total 265 individuals of fruit bats were captured using mist net in five
habitat types, which are secondary forest (1200 m asl), mixed garden (1453 m
asl), pine forest (1545 m asl), primary forest (2000 m asl) and caves (2200 m asl).
The sample include ten species and seven genera, with the estimation curve
showed 12 species, that means 2 species were not caught yet. The fruit bats
composition were caught are Boneia bidens ( (159 individuals; 60%), Thoopterus
suhaeniahae (50 individuals; 18.86 %), Rousettus celebensis (21 individuals;
7.95%), Thoopterus nigrescens (11 individuals; 4.16%), Eonycteris spelaea (9
individuals; 3.40%), Rousettus amplexicaudatus (6 individuals; 2.27%), Dobsonia
viridis (4 individuals; 2.27%), Styloctenium wallacei (2 individuals; 0.75%),
Dobsonia exoleta (2 individuals; 0.75%) and Cynopterus luzonensis (1
individuals; 0.37%).
Shannon-Wiener diversity showed a highest value (H=1.80) in mixed
garden (1453 m asl) while the lowest value (H=0.32) in pine forest (1545 m asl).
The highest evenness indices value (D=0.77) in mixed garden and the lowest
value (D=0.18) was in pine forest. The decreasing in Shannon-Wiener indices

mainly depended on species richness in each habitat types.While the evenness
shows the species dominance in each habitat types. All species were found in
mixed garden (1453 m asl). Principal component analysis showed the correlation
among species and habitat type. T. suhaniahae were correlated with degraded
habitat, that are mixed garden (1453 m asl) and pine forest (1545 m asl), while T.
nigrescens were correlated with pine forest. Both of species were correlated with
mixed garden and pine forest. B. bidens were correlated with primary forest with
river stream at 2000 m asl, while R. celebensis were correlated with secondary
forest at 1200 m asl. Based on moon phases, all species mostly found in new
moon, first quarter, third quarter and waxing gibbous, except for B. bidens that the
only species was found in full moon phases with less individuals number. Bats
that were found in full moon phases were more adapted in light moon intensity
than other bats.
Keywords: Chiroptera, Fruit bats, distribution, habitat preferences, moon phases

© Copy Right owned by IPB, 2015
All rights reserved
No parts of this document may be reproduced or transmitted in any form or by
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permission from IPB.

THE FRUIT BATS (MEGACHIROPTERA, PTEROPODIDAE)
FROM BAWAKARAENG MOUNTAIN, SOUTH SULAWESI

ELLENA YUSTI

A Graduate Thesis
In partial fullfilment of Master Science degree in Animal Bioscience
Faculty of Mathematics and Natural Science

GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015

External Member Supervisor : Dr Ir Agus Priyono Kartono, M Si

FOREWORDS
All praise to Allah SWT for all the hope and give that I have got this far. My
study and thesis research would not have been accomplished without the help of

many people. Special thanks to Dr Bambang Suryobroto and Prof (Ris) Ibnu
Maryanto as supervisory committee, for all guidance and encouragement as well
as invaluable academic advices for the whole period of my study and research.
Thanks to Direktoral Jendral Pendidikan Tinggi (DIKTI) for Beasiswa Unggulan,
Goverment of Gowa, South Sulawesi, staff of Biology Laboratory Makassar
University, Tata and Nurdin Family and All staff of Zoology LIPI. I am highly
indebted to my beloved family Juswardi Yacob Zein (father), Tuti Irda (mother),
Lazuardi Akbar (brother) and Thursina (sister) for their love and support who
always inspire and encourage me for higher education. My friends Husni
Mubarok, Agmal Qodri and Rizaldi Triaz thanks for help in the field. All my
friends in BSH 2012 thanks for support and friendship, especially for Silvia
Puspitasari, Wahyudin Karim and Saudia Fitria.
Bogor, February 2015

Ellena Yusti

TABLE OF CONTENTS
LIST OF TABLES

vi

LIST OF FIGURE

vi

LIST OF APENDIXES

vi

1 INTRODUCTION
Background
Aims

1
1
2

2 METHODS
Study Sites
Fruit Bats Sampling

Sample Identification
Sex, Age Catagorized and Reproduction Status
Data Analysis

2
2
3
4
6
6

3 RESULTS AND DISCUSSIONS
RESULTS
Fruit Bats Composition and Identification
Fruit Bats Estimation Number
Age Catagorized and Reproduction Status of Female Bats
Diversity of Fruit Bats
Habitat Preferences
Fruit Bats Individuals Number Captured In Relation To Moon Phases

7
7
7
8
9
9
10
11

DISCUSSIONS
Fruit Bats Captured and Distribution
Fruit Bats Estimation Number
Age Catagorized and Reproduction Status of Female Bats
Diversity of Fruit Bats
Habitat Preferences
Fruit Bats Individuals Number Captured in Relation To Moon Phases

13
13
14

14
14
15
16

4 CONCLUSIONS

17

REFERENCES
APPENDIXES
BIOGRAPHY

17
21
35

LIST OF TABLES
1 Habitat types description in Bawakaraeng mountain
2
2 Individual number of bats effort mist net/nights captured in different habitat
types
8
3 Individuals number of species in each habitat types and variations in both
Shannon-Wiener and Simpson indexes for fruit bats diversity
10
4 Effort of individuals captured mist net/nights based on moon phases
12

LIST OF FIGURES
1
2
3
4

Collecting specimens sites
Ilustration of mist net replacement
External morphology measurement
Skull measurement

3
4
5
5

5 Estimation curve number of bats were calculated based on the individuals
number each night with Jack 1 Mean value
8
6 Female reproduction condition each month, R=reproductive, NR=non
reproductive, P=pregnant, N=nursing
9
7 Principal Component Analysis (PC1, PC2, PC3)
8 Principal Component Analysis of Bat species and Moon Phases

11
12

LIST OF APPENDIXES
1 Habitat types of Fruit Bats in Bawakaraeng mountain
2 Morphology and skull of captured fruit bats in Bawakaraeng Mountain
3 Measurement (mm) for external morphology and skull

22
24
27

1

1 INTRODUCTION
Background
Bats belong to the order Chiroptera and can be distinguished from all other
mammals by their ability to fly (Mickleburgh et al. 1992; Feldhamer et al. 1999;
Suyanto 2001). Bats are a remarkably successful group, the second largest
mammalian order in biodiversity after the Rodentia (Corbet and Hill 1992;
Feldhamer et al. 1999). There are about 950 species of bats are widespreadly
distibuted in the world (Mickleburgh et al. 1992). Bats divided into two suborder,
they are Megachiroptera (fruit-eating bats) and Microchiroptera (insect-eating
bats) (Corbet and Hill 1992; Feldhamer et al. 1999; Suyanto et al. 2002).
Fruit bats consists of a single family, Pteropodidae, which includes 44
genera and 166 species widespreadly distributed in sub tropical and tropical areas,
from Africa, Madagascar, East Mediterania, Southern Asia, Southeast Asia to
Australia, New Caledonia and islands in the Pasific Oceans (Feldhamer et al.
1999; Mickleburgh et al. 1992). Fruit bats are commonly found in agricultural
areas with cultivated plant, secondary and primary forest that associated with food
resources (Corbet and Hill 1992; Medellin et al. 2000; Fukada et al. 2002;
Maryanto et al. 2011). Fruit bats have an important role in ecosystem as seed
dispersers and pollinators (Mickleburgh et al. 1992; Maryati et al. 2008). There
were 14 families, at least 141 plant species that polinnated with fruit bats (Fujita
and Tuttle 1991; Mickleburgh et al. 1992). The distribution and diversity of bats
are influenced by food resources, habitat types, elevation and environmental
factors such as rainfall, wind, moon phases can all affect number captured and bat
activity while bats generally found in the dark moon phases than full moon phases
(Barlow 1999; Bork 2006).
In Indonesia, there are 25 genera and 77 species of fruit bats and some of
them are endemic in certain areas (Maryanto and Higashi 2011). Sulawesi island
(Indonesia) is remarkable for complex geological history and high biodiversity
fauna, especially for fruit bats (Campbell et al. 2007; Maryanto and Higashi
2011). The diversity of fruit bats (Pteropodidae) is higher on Sulawesi than other
islands, which 29 species of fruit bats are widespread in various areas with 10.7 %
endemism level after Papua (Maryanto and Higashi 2011). Some species were
endemic in Sulawesi are Boneia bidens, Cynopterus luzoniensis, Dobsonia
exoleta, Harpyionycteris celebensis, Neopteryx frosti, Pteropus pumilus,
Rousettus celebensis, Rousettus linduensis, Styloctenium wallacei and Thoopterus
suhaniahae (Suyanto et al. 2002).
Some studies about fruit bats diversity in Sulawesi has been recorded in
several areas and found a new species. Maryanto et al. (2011) found 16 species of
fruit bats in Lore Lindu National Park, Central Sulawesi in 17 different habitat
type with the altitude ranging from 300 to 2100 meter above sea level (m asl).
One of the new species Rousettus linduensis was found in 600-1800 m asl. The
study revealed that the diversity of fruit bats decreases as the altitude increases.
While Maryanto et al. (2012) were found a new species Thoopterus suhaniahae
from Sulawesi and adjacent island which different with the sympatric species
Thoopterus nigrescens. Species diversity and distribution of fruit bats

2
(Megachiroptera) in other parts of Sulawesi remain poorly unknown.
Bawakaraeng mountain, Gowa, South Sulawesi is the one highest mountain in
South Sulawesi, with peak at 2830 m asl were located 90 km from Makassar
(Sumaryono and Dasa 2011). The forest types of mountain Bawakaraeng are
lowland forest, secondary forest and primary forest that suspected to have high
diversity of fruit bats (Hasnawir and Kubota 2006).
Aims
This study aims to determine the fruit bats composition, including age
catagorized and reproduction status of female fruit bats, fruit bats diversity in each
habitat types, habitat preferences and relation between captured individual fruit
bats with the moon phases in the mountain region of Bawakaraeng.

2 METHODS
Study sites
This study was conducted at Bawakaraeng mountain, Gowa and Sinjai
Barat, South Sulawesi from September until December 2013. Sampling was done
in different habitat types with a various levels of elevation (1200 to 2200 m asl)
(Apendixes 1). Five sites were set up at different elevations and habitats : mixed
garden (1453 m asl), pine forest (1545 m asl), secondary forest with a river stream
(1200 m asl), primary forest with a river stream (2000 m asl) and cave (2200 m
asl) (Table 1, Figure 1 and Appendix 1).
Table 1 Habitat types description in Bawakaraeng mountain
No

Habitat Types

1

Mixed Garden

Elevation
( m asl)
1453

Description

2

Pine Forest

1545

The dominant vegetation which are pine, shrubs,
herbs, and ferns. This area were directly adjacent
to forest plantation.

3

Secondary Forest

1200

The forest area has a cliffs and waterfalls.
Vegetation in this habitat in the form of fruit
trees, ferns, shrubs, and herbs.

4

Primary Forest

2000

The forest area has a cliffs, small caves and
river. Vegetation in this habitat which are trees,
ferns, shrubs, and herbs.

5

Cave

2200

This area were located in primary forest with
cliffs. The area of forest were dominated by
trees, ferns, shrubs and herbs.

The dominant vegetation which are trees,
shrubs, herbs, and flowering plants. In this area
were found a variety of fruit plants, such as
guava and banana.

3

Figure 1 Collecting specimens sites (1) Mixed garden (1453 m asl); (2) Pine forest
(1545 m asl); (3) Secondary forest (1200 m asl); (4) Primary forest (2000
m asl) and (5) Cave (2200 m asl). The habitat types description refer to
Table 1.

Fruit Bats Sampling
Mist nets were used to trap bats placed in 2-3 meter above ground (Figure
2). Each sampling site was sampled by using six and four mist nets. Mist nets size
were 9 m x 2.5 m (four mist net) and 12 m x 3 m (two mist net). Mixed garden use
six mist net with , pine forest, secondary forest and primary forest use four mist
net. The mist nets were set up started from 18.00 pm to 06.00 am. Bats were
trapped placed in a sealed bag which contains cotton wool soaked in the
anaesthetic (Chloroform). The bats were left in bag for at least 10 to 15 minutes
and then labelled. Sex and age catagorized were recorded. Specimens were
preserved in 10 % formalin for further identification.

4

Figure 2 Illustration of mist net replacement (Kunz 1988)

Sample Identifications
Samples identification were based on Corbet and Hill (1992), Suyanto
(2001) and observed against voucher bats specimen in Indonesian Institute of
Science (LIPI). Identification were done until species level, including sex, age
category, reproduction status and measurement of external morphology and skull.
External morphology measurement
Measurement of external morphology including, second digit metacarpal
length (D2MCL), third digit metacarpal length (D3MCL), third digit first
phalange length (D3P1L), second phalange length (D3P2L), fourth digit
metacarpal length (D4MCL), fourth digit first phalange length (D4P1L), second
phalange length (D4P2L), fifth digit metacarpal length (D5MCL), fifth digit first
phalange length (D5P1L), second phalange length (D5P2L) (Maryanto and Yani
2003; Rahman and Abdullah, 2010) (Figure 3).

5

Figure 3 External morphology measurement (Rahman and Abdullah 2010)
Skull Measurement
Measurement of skull including Greatest skull length (GLS), Postorbital
width (POW), Zygomatic breadth (ZB), Braincase width (BW), Mesopterygoid
fossa width (MSF), Bulla length (BL), Dentary length (DL), outside upper canine
width (C1-C1), upper canine to third molar (C1-M3), upper third molar widht (M3M3), Condilo Canine Length (CCL), Condilo Braincase Length (CBL), Lower
first canine (C1-C1), Lower canine to third molar length (C1-M3), Lower molar
width (M3-M3) (Figure 4) (Kitchener and Maharadatunkamsi 1999; Helgen
2005).

Figure 4 Skull measurement (Rahman and Abdullah 2010)

6
Sex, Age Catagorized and Reproduction Status
Male and female bats were differentiated by observing their external
reproductive organs, penile and testes in males, vagina and mammary gland in
females (Racey 1988). Age categories divide into infant, juvenile, sub adult and
adult (Suyanto 2001; Racey 1988). Juveniles and sub adult were distinguished
from adults by joints of finger bone, cartilaginous ends of the bones in juveniles
appear paler and more translucent than the joints of adults (Cristian and Helversen
2005). Female reproductive condition were divided into four categories, (1)
reproductive (R) is a female in adult age, (2) non-reproductive (NR) is a female
individual that does not reproduce, (3) pregnant (P) are individual pregnant
females, (4) lactating (L) are female individuals that are nursing their baby (Racey
1988). Pregnancy in females could be observed with full and enlarged size in
abdomen. Lactating females could be observed by enlarged nipples, which when
gently massaged will express milk (Racey 1988).
Fruit Bat Individuals Number in Relation to Moon Phases
Bats activity based on moon phases was determined by calculating effort
(number of individuals / night / nets). The data from different nights are pooled
according to moon phases (Morrison 1978; Bork 2006). The moon phases were
used in this study are based on Bork (2006), they are new moon, wanning
crescent, third quarter, waxing gibbous and full moon. The moon phases data
among nights were download in www.moonphasesconnection.com.
Data Analysis
Estimation on species was analyzed with Estimates S 8. Species captured
were calculated by sampling effort to standardised individuals number of bats
(Sampaio et al. 2003). Shannon-Wiener’s indexes and Evenness (Maguran 2004)
were used to compare the diversity of bats in each habitat types. Bats reproductive
condition and age categories were count manually. Principle Component Analysis
(PCA) was employed to group bats according to habitat preferences and
correlation between number of fruit bats individuals and moon phases with
Paleontological Statistic (Hammer et al. 2001).

7

3 RESULTS AND DISCUSSIONS
Results
Fruit Bats Composition and Identification
Composition bats species were captured during 40 nights included Boneia
bidens (Jentink, 1989) (159 individuals; 60%), Thoopterus suhaeniahae
Maryanto, et al. 2012 (50 individuals, 18.86%), Rousettus celebensis K.
Andersen, 1907 (21 individuals; 7.95%), Thoopterus nigrescens (Gray,1870) (11
individuals; 4.16%), Eonycteris spelaea Jentink, 1888 (9 individuals; 3.40%),
Rousettus amplexicaudatus (Geoffroy, 1810) (6 individuals; 2.27%), Dobsonia
viridis (Heude, 1896) (4 individuals; 2.27%), Styloctenium wallacei Gray, 1866 (2
individuals, 0.75%), Dobsonia exoleta (2 individuals, 0.75%) and Cynopterus
luzoniensis (Peters, 1861) (1 individuals; 0.37%) (Appendix 2). Six of ten fruit
bats species were found endemic to Sulawesi which are Boneia bidens,
Cynopterus luzoniensis, Dobsonia exoleta, Rousettus celebensis, Styloctenium
wallacei and Thoopterus suhaeniahae. Two species were widespreadly distribute
in entire Indonesia are Rousettus amplexicaudatus and Eonycteris spelaea.
Boneia bidens has a dark brown hair in dorsal and ventral body part. B.
bidens were resemble to Rousettus genera, primarly Rousettus celebensis. B.
bidens were differentiated with R. celebensis in body size, where the HBL=
128.25, FA= 100 and GSL = 45.78, while R. celebensis were medium size, with
HBL= 96.89, FA= 74.68 and GSL= 37.39. R. celebensis has a dark brown hair, its
different with R. amplexicaudatus which has a pale brown in dorsal and ventral
body part with slightly hair with HBL= 100.17, FA= 80.29 and GSL= 35.22.
Eonycteris spelaea has a brown hair in dorsal and ventral. This species were
resemble to R. amplexicaudatus, but E. spelaea has not a claw in second digiti,
where the HBL= 108.15, FA= 77.82 and GSL= 35.18 and has a gland like kidney
in anus part. Thoopterus suhaniahae and Thoopterus nigrescens were
differentiated in forearm length and pelage colouration. T. suhaniahae has a
brown grey in under part from chest to abdomen and dark borwn in under part and
forearm, while T. nigrescens has a brown pale under part hair. Forearm length in
T. suhaniahae FA= 75.37, HBL= 98.03 and GSL= 35.36 while T. nigrescens HBL
= 96.45, FA= 72.90 and GSL= 34.32. Cynopterus luzoniensis were differentiated
by pelage colouration, where in neck part has a orange-brown hair. In addition,
this species were differentiated with other species by small body size with HBL=
79.04, FA= 64,38 and GSL= 32.22. Styloctenium wallacei were diffrentiated by
white stripe mark in face part and a brown molar colour. This species has HBL=
145.22, FA= 94.55 and GSL= 48.91. Both Dobsonia exoleta and Dobsonia viridis
has not a hair in dorsal body part. D. viridis has a brown hair in dorsal, with a
brown hair in head part, while D. exoleta has a dark brown and dark orange hair in
neck and D. exoleta has a dark yellow claw, but D. viridis has a dark green claw.
The all external morphology measurement and skull measurement in Appendix 3.

8
Fruit Bats Estimation Number
Total number of species exist in Bawakaraeng Mountain was estimated
based on the number of individuals captured each nights. Total night sampling
was 40 nights, where successful trapping were 27 nights. Estimation of bats
species number during 27 nights were 12 species (Figure 5). Sampling effort on
this study were almost maximal, because only two species estimated was not
caught yet, the bats were captured included 10 species from 7 genera (Table 2).
The total sampling effort (5112 m2) were resulted 265 individuals of fruit bats
representing 10 species (Table 2).
Table 2 Individual number of bats effort mist net/nights captured in different
habitat types
Habitat

Altitude

Bb*

Ts*

Tn

Rc*

Ra

Es

Sw*

De*

Dv

Cl*

(m asl)
SF

1200

0.00

3.25

0.00

15.9

0.00

0.00

0.00

0.00

0.00

0.00

MG

1453

2.35

10.29

2.35

55.6

1.76

2.65

0.59

0.59

1.76

0.29

PiF

1545

0.00

11.10

2.06

0.00

0.00

0.00

0.00

0.00

0.00

0.00

PF

2000

55.60

4.40

2.2

4.4

0.00

0.00

0.00

0.00

0.00

0.00

Abbreviations of habitat and species names MG=Mixed Garden, PiF=Pine Forest,
SF=Secondary Forest, PF=Primary Forest; Bb=Boneia bidens, Ts=Thoopterus
suhaniahae, Tn=Thoopterus nigrescens, Rc=Rousettus celebensis, Ra=Rousettus
amplexicaudatus, Es=Eonycteris spelaea, Sw=Styloctenium wallacei, De= Dobsonia
exoleta, Dv=Dobsonia viridis, Cl=Cynopterus luzoniensis (*= endemic species in
Sulawesi and adjacent island).

Figure 5 Estimation curve number of bats was calculated based on the
individuals number each night with Jack 1 Mean value.

9
Age Catagorized and Reproduction Status of Female Bats
Based on age catagorized, 13 infants (4.90%), 16 juveniles (6.03%), 17
sub adults (6.41%) and 219 adults (82.64%). The composition of the fruit bats
captured by sex was 121 females (45.66%) and 144 males (54.33%). Based on
female bats reproduction categories, the composition of female individuals in
reproductive period or adult were 73 individuals (60.33%), not reproducing or
non-adult were 22 individuals (18.18%). Of the female reproductive category, 6
individuals (4.95%) are pregnant and 20 individuals (16.52%) are nursing. Some
species that were found in pregnant and have an infant and nursing are B. bidens,
D. viridis, E. spelaea, T. suhaniahae, R. amplexicaudatus, and R. celebensis. The
reproductive female were found higher in November and December (Figure 6).

Figure 6 Female reproduction condition each month, R=reproductive, NR=non
reproductive, P=pregnant, N=nursing

Diversity of Fruit Bats
Shannon-Wiener diversity showed a highest value in mixed garden (1453
m asl) H’=1.80, while the lowest value in pine forest (1545 m asl) H’=0.32. The
highest dominance indices value D = 0.77 in mixed garden and the lowest value D
= 0.18 in pine forest (Table 3). The decreasing in Shannon-Wiener indices mainly
depended on species richness in each habitat types.

10
Table 3 Individuals number of species in each habitat types and variation in both
Shannon-Wiener and Simpson indexes for fruit bats diversity
Number of Individuals
Species
Boneia bidens
Thoopterus suhaniahae
Thoopterus nigrescens
Rousettus celebensis
Rousettus amplexicaudatus
Eonycteris spelaea
Dobsonia viridis
Dobsonia exoleta
Styloctenium wallacei
Cynopterus luzoniensis
Number of Individuals
Number of Species
Shannon-Wiener Index (H')
Simpson Index (D)

MG
1453 m asl
8
35
8
7
6
9
4
2
2
1
82

PiF
1545 m asl
0
9
1
0
0
0
0
0
0
0
10

SF
1200 m asl
0
2
0
10
0
0
0
0
0
0
12

PF
2000 m asl
52
2
2
4
0
0
0
0
0
0
60

10

2

2

4

1.80
0.77

0.33
0.18

0.45
0.28

0.53
0.24

Abbreviations of habitat MG=Mixed Garden, PiF=Pine Forest, SF= Secondary
Forest, PF=Primary Forest

Habitat Preferences
Result of principal component analysis revealed that PC1, PC2, and PC3
for 97.92% in total variance, accounted 52.19% (PC1), 24.10% (PC2) and 21.61%
(PC3) (Figure 6). Principal component analysis showed the correlation among
species and habitat types. T. suhaniahae and T. nigrescens were correlated with
degraded habitat, that are mixed garden (1453 m asl) and pine forest (1545 m asl).
B. bidens and T. nigrescens were correlated with primary forest with river stream
at 2000 m asl, while R. celebensis were correlated with secondary forest at 1200
m asl. Another six species, that are C. luzoniensis, D. viridis, D. exoleta, E.
spleaea and S. wallacei were correlated with mixed garden (Figure 7).

11

Figure 6 Principal Component Analysis (PC1, PC2, PC3)

Figure 7 Principal Component Analysis (PC1, PC2, PC3) explained the
correlation between fruit bats and habitat

Fruit Bat Individuals Number Captured In Relation To Moon Phases
Species that were captured each night was calculated by sampling effort to
standardised individuals number of bats (Table 4). Principal component analysis
was revealed that PC1, PC2, and PC3 for 96.18% in total variants, accounted for
53.02% (PC1), 37.16% (PC2) and 5.88% (PC3). Principal component analysis
showed the correlation between species and moon phases. The three moon phases
which are new moon, waxing gibbous and full moon correlated with B. bidens and
R. celebensis. Two species that were related to wanning crescent and third quarter
phases, they are T. suhaniahae and D. viridis, whereas species that were related
with waxing gibbous phases is R. celebensis (Figure 8).

12
Table 4 Effort of individuals captured mist net/nights based on moon phases
Moon
Phases

Individuals number of bats captured
Bb*

Ts*

Tn

Rc*

Ra

Es

Sw*

De*

Dv

Cl*

A

104.74

25.70

14.00

15.99

12.35

6.17

0.00

6.17

0.00

0.00

B

8.37

48.48

3.97

5.95

3.97

8.37

2.42

1.98

3.97

0.00

C

2.06

20.7

9.00

5.85

2.06

1.74

1.74

0.00

12.80

2.06

D

43.65

27.78

0.00

39.68

3.97

0.00

0.00

0.00

0.00

0.00

E

22.22

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

Abbreviations of moon phases A=new moon, B=wanning crescent, C=first
quarter, D=waxing gibbous, E=full moon. Abbreviation of species name refer to
table 2)

Figure 8 Principal Component Analysis of Bat species and Moon Phases

13

Discussions
Fruit Bats Captured and Distribution
The fruit bats that were found in Bawakaraeng Mountain were also widely
distributed in Sulawesi and adjacent island (endemic species). These species
included B. bidens, C. luzonensis, D. exoleta, S. wallacei, T. suhaniahae and T.
nigrescens (Suyanto et al., 2002), Two species which are D. viridis and R.
celebensis were endemic to Sulawesi and Molluccas. Meanwhile, R.
amplexicaudatus and E. spelaea are found widely distributed throughout
Indonesia (Corbet and Hill 1992; Suyanto et al.2002). R. celebensis was found in
Sulawesi, but R. amplexicaudatus were found in entire Indonesia (Corbet and Hill
1992). Rousettus genera entirely distributed in Indonesian islands, but some of the
species are endemic in several areas (Corbet dan Hill 1992; Suyanto et al. 2002).
Endemic Sulawesi Rousettus is Rousettus linduensis, a new fruit bat that was
found in Lore Lindu National Park, Central Sulawesi (Maryanto and Yani 2003).
Corbet and Hill (1992) were recorded the distribution of four Rousettus species in
Indonesia, they are R. amplexicaudatus (Sumatra, Borneo, Sulawesi, Molluccas,
Java, Bali and Lesser Sunda), R. celebensis (Sulawesi), R. leschenaultii (Java), R.
spinalatus (Sumatra and Borneo).
Boneia bidens were morphologically resemble with Rousettus genera, but
the different are Boneia has larger size than Rousettus (Bergmans and Rozendaal
1988). Boneia bidens were endemic to Sulawesi, this species were recorded in
Sulawesi and adjacent island (Suyanto 2001; Suyanto et al. 2002). Cynopterus
genera were widespreadly distribute in entire Indonesia. This genera has seven
species, they are C. brachyotis (Sumatra, Java, Borneo, Bali, and Molluccas), C.
Horsfeldi (Sumatra, Java and Borneo), C. luzoniensis (Sulawesi), C. minutus
(Sumatra, Borneo, Java and Sulawesi), C. nusatenggara (Nusa tenggara), C.
sphinx (Sumatra, Java, Borneo, Sulawesi) and C. tithaecheilus (Sumatra, Java,
Bali, Lombok and Timor) (Kitchener and Maharadatunkamsi 1991; Corbet and
Hill 1992; Suyanto et al. 2002).
Dobsonia genera were widespreadly distribute in east Indonesia region.
That genera has nine species, they are D. beauforti (West Papua), D. crenulata
(Sangihe island, Tongian, Banggai island and Halmahera), D. emersa (West
Papua), D. exoleta (Sulawesi), D. magna (West Papua), D. minor (Sulawesi and
West Papua), D. moluccensis (Molluccas), D. peroni (Bali and Nusa Tenggara)
and D. viridis (Sulawesi and Molluccas) (Bergmans and Rozendaal 1988; Corbet
and Hill 1992). Eonycteris genera have two species that were found in all
Indonesian, they are E. spelaea (all Indonesia islands) and E. major (Borneo)
(Corbet and Hill 1992; Suyanto 2001; Suyanto et al. 2002).
Styloctenium genera were only have one species in Indonesia Styloctenium
wallacei (Stripe fruit bats), this species were endemic to Sulawesi (Corbet an Hill
1992; Suyanto et al. 2002). Another species from that genera were Styloctenium
mindorensis were distributed in Philiphine islands (Esselstyn 2007). Thoopterus
genera were widely distributed in Sulawesi and Molluccas (Bergmans and
Rozendaal 1988). This genera has two species, they are Thoopterus nigrescens
and Thoopterus suhaniahae that a sympatric species (Maryanto et al. 2012).
Thoopterus suhaniahae were a new species from Sulawesi, this new species were

14
found in several areas at Talaud, Wowoni islands and Rorekatimbu (Maryanto et
al. 2012).
Fruit Bats Estimation Number
Estimated curve were discuss differences between species number of
collected and expected species (Figure 3). Sobs (Mao Tau) is the total number of
species that was captured in this study. Jack 1 mean value is the first order
Jacknife richness estimator of species number (Gotelli and Colwell 2011). Some
factors that suspected causing other bats species were not caught including
sampling duration, mist net efficiency, including the placement and regular check
and environmental conditions, such as heavy rain, wind and the moon phases
(Barlow 1999; Lang et al. 2004; Larsen et al. 2007). In this study, it is possible to
get 12 species of fruit bats if adding night sampling was conducted. Mist net
position in this study were held in some area, there were near fruit trees, over
stream area, but some mist net were placed far from fruit trees. Mist net was not
regularly checked each night and environment factor that were observed there was
no bats were caught in heavy rain and wind. In addition, in the full moon phases
there were only a few individuals of bats captured.
Age Catagorized and Reproduction Status of Female Bats
Age catagorized and pregnant female bats shows the correlation between
breeding season and rainfall in Neotropics area (Coates and Estrada 2001). In this
study, the most age catagorized were caught are adult and some are pregnant
female. Coates and Estrada (2001) reported that flowering peaks occur in dry
season, where the fruiting peaks in the rain season. That study shows that the
reproductive peaks (adult age catagorized) were coincident with these periods. In
this study, a reproductive peaks (adult catagorized) were found in December. In
this study, the adult female more dominance than non reproductive, pregnant and
nursing. The adult females number mostly found in November and December.
Generally, bats were produce one offspring in a year with periods 3-4 month, with
nursing period about one year (Suyanto 2001).
Diversity of Fruit Bats
The decreasing in Shannon-Wiener indexes mainly depended on species
richness in each habitat type (Magguran 2004). Shannon-Wiener indexes show the
species richness value, while the evennes indexes shows the species dominance in
sampling area (Maguran 2004). The fruit bats diversity to be maintained by
various factors; including the diversity of food sources (Hall et al. 2004;
Hodgkinson et al. 2004), the availability of roosts (Zubaid 1993) and the
heterogenity of forest structure (Hall et al. 2004; Hodgkison et al. 2004). In this
study, there is abundance of food in habitat types, they are mixed garden, primary
forest and secondary forest with stream. The number individuals captured of fruit
bats were highest in mixed garden than another habitat types. The similar resulted
were reported by Heaney et al. (1989), Medellin et al. (2000) and Hall et al.
(2004), where the highest individuals number of bats captured were found in

15
agricultural areas. Another study were also found the highest abundance of fruit
bats in secondary and primary forest, some habitat types with a river stream with
flower and fruit availability (Heideman and Heaney 1989; Wiantoro and Ahmadi
2011).
Some studies reported that the number of fruit bats species also influence
with the elevation pattern. Maryanto et al. (2011) reported that the highest
diversity of fruit bats were stable in lower mountain association (300 and 1500 m
asl) and decreased in upper mountain association (2100 m asl). Bruce et al (1998)
in study about elevation in relation among birds, bats, and rats diversity in
Amazon mountain, Peru reported that the highest bats diversity were found in
500-1500 m asl and decreased in 2000-3500 m asl. Both studies suggest that
species richness decreases with elevation and habitat type that associated with
food avaibility.

Habitat Preferences
Fruit bats were commonly found in different habitat types which closely
related with fruit and flower availability (Corbet and Hill 1992; Maryanto et al.
2011). Habitat used in fruit bats were vary as roosting habitat and foraging habitat
(Mickleburgh et al. 1992), such as agroforestry, mixed garden, forest area
(primary and secondary forest) and caves (Bergmans and Rozendall 1988; Corbet
and Hill 1992; Wijayanti 2011). In this study, all species were found in mixed
garden (1453 m asl) (Table 1). The species were simpatrically found in mixed
garden, they are C. luzoniensis, D. viridis, D. exoleta, E. spelaea, R.
amplexicaudatus and S. wallacei. Another four species which are B. bidens, R.
celebensis, T. nigrescens and T. suhaniahae were found simpatrically in primary
forest (2000 m asl). Previous study also found Thoopterus suhaniahae has been
collected sympatrically with T. nigrescens, R. celebensis and C. luzoniensis
(Maryanto et al. 2012). Some species were found sympatrically in the same
habitat shows the present of fruit bats were depend on habitat types that closely
related with fruit and flower availability (Whitmore 1989; Corbet and Hill 1992).
Cynopterus was generally found in garden and roosting in leaf of trees and
some caves with hight light intensity (Storz 2000). This genera were adapted in
light condition (Storz 2000; Wijayanti 2011). In this study, Cynopterus were
found in mixed garden with open area (1453 m asl), this species possibly used the
bamboos trees as roosting habitat. Some studies reported that Cynopterus were
prefered to roosting in trees, but some studies were found the species in this
genus, Cynopterus horsfieldii and Cynopterus brachyotis in some caves with hight
light intensity at karst Gombong, Central Java (Wijayanti 2011). Another species
such as B. bidens, R. celebensis, T. suhaniahae, and T. nigrescens were generally
found in forest with a shelter rock and caves as roosting habitat, but were found in
agroforestry as foraging habitat (Bergmans and Rozendall 1988; Maryanto and
Yani 2003; Maryanto et al. 2011; Maryanto et al. 2012). B. bidens noted generally
found in upper mountain area at 1800-2000 m asl and caves (Maryanto et al.
2011). Bergmans and Rozendaal (1988) recorded B.bidens in lowland forests,
between 200 and 500 m asl of Tambun and Lembeh Island. In this study, this

16
species were found at 1453 m asl, 2000 m asl and 2200 m asl in caves with large
colony.
T. suhaniahae and T. nigrescens were generally found in mixed garden
such as cacao and coffee, secondary and primary forest and lower forest area
(Maryanto et al. 2012). T. nigrescens and T. suhaniahae simpatrically found in
different habitat types such mixed garden, pine forest and primary forest with
river flow (1400-2000 m asl). This suggested that both species are widely
distributed in different habitat types. Rousettus genera that were found are R.
celebensis and R. amplexicaudatus. Both species were found in mixed garden, but
R. celebensis was found in secondary and primary forest with river flow (12002000 m asl). Previous study noted that R. celebensis related to garden location and
cave (Bergmans and Rozendall 1988) and lower lowland forest with 1200 m asl
(Maryanto et al. 2011).
Species that were specialized nectarivorus, E. spelaea and R.
amplexicaudatus were found in mixed garden (1453 m asl), both species were
play major in pollination (Maryati et al. 2008; Soegiharto et al. 2011). This
species were commonly found in agricultural with flowers availability, but some
studies were found this species in some caves, including karst caves with high and
low light intensity (Maryanto and Maharadatunkamsi 1991; Kunz and Fenton
2003; Ruczynski et al. 2010; Wijayanti 2011). Altringham (1996) reported that R.
amplexicaudatus and E. spelaea were able to used echolocation, that cases suggest
that both species were adapted in hight and low condition (Wijayanti 2011). In
this study both species were suspected used the caves in Bawakaraeng mountain
region as roosting habitat. In addition, E. spelaea and R. amplexicaudatus has a
widely foraging area about 38 km, this ability was important in seeds dispersal
(Suyanto 2001; Kunz and Fenton 2003). Another species that were found in
mixed garden (1453 m asl) are D. exoleta, D. viridis and S. wallacei. Esselsstyn
(2007) were found Styloctenium near guava trees in Phillipines, while Maryanto et
al. (2011) recorded S. wallacei at 600-1800 m asl in lowlands area. D. exoleta and
D. viridis were recorded found in lowland forest at 600-1200 m asl (Maryanto et
al. 2011).
Fruit Bat Individuals Number Captured In Relation To Moon Phases
The behaviour and activity of nocturnal animals affected with changing
light conditions in relation with lunar cycle (Lang et al. 2005). Morrison (1978)
reported that lunar phobia to explain this behaviour, some animals forage less or
avoid the bright moon phases in order to reduce predation risk. Lunar phobia have
been documented in some animals such as Rodent (Clarke 1983) and Kanggoro
rats (Dipodomys spectabilis) (Daly et al. 1992). Bats are classified as nocturnal
animals where the activity at night is affected by moon light cycle or moon
phases (Lunar phobia) (Barlow 1999; Lang et al. 2005; Bork 2006; Mello et al.
2013). This behavior suggests that flying in moonlight may significantly increase
risk of predation by visually oriented predators such as snake and owl (Lang et al.
2005).
Lunar phobia have been reported in some bats in Neotropical areas
(Elangovan and marimuthu 2001; Lang et al. 2005; Bork 2006; Mello et al. 2013).
Several studies have documented that captured rates of both insectivorous and

17
frugivores bats decrease when full moon or bright moonlight (Morrisson 1978;
Lang et al. 2005; Bork 2006; Mello et al. 2013).
All species in this study mostly found in new moon, waxing crescent, third
quarter and waxing gibbous, except for B. bidens, that the only species was found
in full moon phases (Table 2). Lang et al. (2005) reported that foraging activity
bat, Lophostoma silvicolum were higher in new moon phases than full moon
phases. Some studies reported that the number individuals captured of Noctilio
leporinus (Bork 2006); Artibeus lituratus, Carollia perspicillata, Sturnira lilium
(Mello et al. 2013) were found higher in dark moon phases, they are new moon,
waning gibbous and waxing gibbous than full moon phases. Bats that were found
in full moon phases were more adapted in light moon intensity than other bats
(Bork 2006). Mello et al. (2013) reported that during full moon bats move to
darker parts to forage B. bidens that were found in full moon phases showed this
species was more adapted to the bright moonlight intensity, suspected this species
were found in degraded habitat, such as agricultural (Bergmans and Rozendaal
1988).
4 CONCLUSIONS
The highest individual fruit bats captured in Bawakaraeng mountain were
Boneia bidens (60%) and Thoopterus suhaniahae (18.86%), while composition of
the fruit bats captured by sex are 121 females (45.66%) and 144 males (54.33%).
In this study, the most age catagorized were caught are adult (reproductive). The
highest diversity of fruit bats in Bawakaraeng Mountain were in mixed garden
(1453 m asl), primary forest (2000 m asl) and secondary forest with stream (1200
m asl). Fruit bats abundance in this study were found mixed garden (1453 m asl)
and primary forest (2000 m asl) were tightly associated with food availability.
Moon phases affected the number of bats individual captured where the number of
individual bats captured in the dark moon phase was higher than full moon
phases.
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