Morphological variation of the Diadem Leaf-nosed Bat, Hipposideros diadema, Geoffroy, 1813 (Chiroptera: Hipposideridae) in Caves in West Sumatra, Indonesia

Abstract

We assumed that Bukit Barisan as a physical barrier and its acts to population exchanges in West Sumatra, as well as for
terrestrial animal group. If it does for bats in case for the Diadem Leaf-nosed Bat, Hipposideros diadema, which have superior
dispersal powers to many other terrestrial group, then we might expect to see this some how reflected in morphological
divergence. A total of 58 adult of H. diadema were collected directly using harp traps from several Cave in West Sumatra
(Kalilawa Cave, Padang; Lereng Cave, Pariaman; (western of Bukit Barisan) and Salamaik Cave, Sawahlunto (eastern of Bukit
Barisan)). The samples were collected on January-December 2013. Kruskall Wallis Test, Mann Whitney U Test, Principal
Component Analyses (PCA) and Cluster Analysesdemonstrated that these population could be separated clearly from
one to another. This analyses based on 26 external and 15 skull measurement. The result showed that population of H.
diadema from Salamaik Cave in Sawahlunto differ from H. diadema in Padang and Pariaman. Divergence characters among
three population of H. diadema was found using Kruskall Wallis test. Mann-Whiney U test showed divergence characters
between two different population. The result of PCA was congruence to phenogram obtained by UPGMA that showed close
relationship between population of H. diadema from Kalilawa Cave, Padang to Lereng Cave, Pariaman and different from
Salamaik Cave, Sawahlunto. We conclude that Bukit Barisan barriers could be affected to morphological divergence among
H.diadema in West Sumatra.

Full text

Journal of
Indonesian Natural History
December 2014 Vol.2 No.2

2© University of Andalas / Copenhagen Zoo
Dr. Wilson Novarino
Associate Professor for Biology
Department of Biology
University of Andalas, Indonesia
Email: editorjinh@jinh.org
Journal of Indonesian Natural History
Dr. Carl Traeholt
Programme Director, Southeast Asia
Research and Conservaon Division
Copenhagen Zoo, Denmark
Email: ctraeholt@gmail.com
Dr. Ardinis Arbain
University of Andalas, Indonesia
Indra Arinal
Naonal Park Management, Department of Forestry Indonesia
Dr. Ahimsa Campos-Arceiz
Nongham University Malaysia Campus, Malaysia
Dr. Mads Frost Bertelsen
Research and Conservaon Division, Copenhagen Zoo, Denmark
Dr. Susan Cheyne
Oxford University, Wildlife Research Unit, United Kingdom
Bjorn Dahlen
Green Harvest Environmental Sdn. Bhd, Malaysia
Dr. Niel Furey
Centre for Biodiversity Conservaon, Royal University of Phnom Penh, Cambodia
Dr. Benoit Goossens
Cardi University, United Kingdom
Dr. Djoko Iskandar
Bandung Instute of Technology, Indonesia
Dr. Mahew Linkie
Fauna & Flora Internaonal, Singapore
Dr. Erik Meijaard
People and Nature Consulng Internaonal, Indonesia
Dr. John Payne
Borneo Rhino Alliance, Malaysia
Dr. Ramadhanil Pitopang
Tadulako University, Indonesia
Dr. Lilik Budi Prasetyo
Bogor Instute of Agriculture, Indonesia
Dr. Dewi Malia Prawiradilaga
Indonesia Instute of Science, Indonesia
Dr. Rizaldi
University of Andalas, Indonesia
Dr. Dewi Imelda Roesma
University of Andalas, Indonesia
Dr. Jerine Rovie Ryan
Wildlife Forensics Lab, Dept. of Wildlife and Naonal Parks, Malaysia
Boyd Simpson
Research and Conservaon Division, Copenhagen Zoo, Denmark
Robert B. Stuebing
Herpetology and Conservaon Biology, Indonesia
Dr. Sunarto
WWF-Indonesia
Dr. Jatna Supriatna
University of Indonesia
Dr. Campbell O. Webb
The Arnold Aboretum, Harvard University, USA
Dr. Zainal Z. Zainuddin
Borneo Rhino Alliance, Malaysia
Editorial board
The Journal of Indonesian Natural History is published biannually by the Department of Biology at the Andalas University, Padang, Sumatra
Barat, Indonesia, in collaboraon with Copenhagen Zoo, Denmark. The Department of Biology at Andalas University is dedicated to
educang Indonesian biologists in the study and conservaon of Indonesia’s biodiversity and natural history. Copenhagen Zoo, through its
Research and Conservaon Division, supports in-situ conservaon in Southeast Asia by assisng local organizaons and individuals who
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The Journal of Indonesian Natural History is published by the Department of Biology, Andalas University, Indonesia in collaboraon
with Copenhagen Zoo, Denmark. It is available for free from www.jinh.net
Cover photo: A at-headed cat, Priornailurus planiceps, at a recent kill. The species is listed as “Endangered” on the IUCN
red-list and currently recorded from Southern Thailand, West Malaysia, Borneo and Sumatra. © Carl Traeholt
Editors

8© University of Andalas / Copenhagen Zoo
Ada Chornelia, Djong Hon Tjong and Dewi Imelda Roesma
Department of Biologi, Faculty of Mathemacs and Natural Science, University of Andalas, West Sumatra, Indonesia
Corresponding author: Ada Chorneliaa, email: chorneliaa@yahoo.co.id
Abstrak
Bukit Barisan sebagai barier sik diprediksi berpengaruh terhadap populasi di Sumatera Barat sebagaimana terjadi pada
kelompok hewan terestrial termasuk kelelawar Hipposideros diadema, yang dikenal memiliki kemampuan dispersal yang
nggi. Pengaruh barier ini diharapkan dapat diama pada perbedaan morfologi. Sejumlah 58 individu dewasa H. diadema
dikoleksi dengan menggunakan Harpa trap pada beberapa goa di Sumatera Barat, yang terdiri dari Goa Kalilawa, dan Goa
Lereng di bagian barat Bukit Barisan dan Goa Salamaik di bagian mur Bukit Barisan. Pengkoleksian sampel di lapangan
dilaksanakan pada bulan Januari-Desember 2013. Pengukuran dilakukan terhadap 26 karakter tubuh dan 15 karakter
tengkorak. Uji Mann-Whitney menunjukkan divergensi karakter antar dua populasi yang berbeda. Hasil PCA sesuai dengan
fenogram yang disusun dengan UPGMA yang menunjukkan populasi H. diadema di Goa Salamaik (populasi mur) berbeda
dengan populasi dari Goa Kalilawa dan Goa Lereng (populasi barat). Disimpulkan bahwa barier Bukit Barisan memungkinkan
berpengaruh terhadap divergensi karakter morfologi antara H. diadema di Sumatera Barat.
Abstract
We assumed that Bukit Barisan as a physical barrier and its acts to populaon exchanges in West Sumatra, as well as for
terrestrial animal group. If it does for bats in case for the Diadem Leaf-nosed Bat, Hipposideros diadema, which have superior
dispersal powers to many other terrestrial group, then we might expect to see this some how reected in morphological
divergence. A total of 58 adult of H. diadema were collected directly using harp traps from several Cave in West Sumatra
(Kalilawa Cave, Padang; Lereng Cave, Pariaman; (western of Bukit Barisan) and Salamaik Cave, Sawahlunto (eastern of Bukit
Barisan)). The samples were collected on January-December 2013. Kruskall Wallis Test, Mann Whitney U Test, Principal
Component Analyses (PCA) and Cluster Analysesdemonstrated that these populaon could be separated clearly from
one to another. This analyses based on 26 external and 15 skull measurement. The result showed that populaon of H.
diadema from Salamaik Cave in Sawahlunto dier from H. diadema in Padang and Pariaman. Divergence characters among
three populaon of H. diadema was found using Kruskall Wallis test. Mann-Whiney U test showed divergence characters
between two dierent populaon. The result of PCA was congruence to phenogram obtained by UPGMA that showed close
relaonship between populaon of H. diadema from Kalilawa Cave, Padang to Lereng Cave, Pariaman and dierent from
Salamaik Cave, Sawahlunto. We conclude that Bukit Barisan barriers could be aected to morphological divergence among
H.diadema in West Sumatra.
Keywords: Bukit Barisan, cave, Hipposideros diadema,morphology, variaon.
Morphological variation of the Diadem Leaf-nosed
Bat, Hipposideros diadema

Received 9th December, 2013; First revision 28th April, 2014;
Second revision accepted 14th May, 2015.

West Sumatra has the largest limestone outcrops in
Indonesia. Caves are known as karsts, and West Sumatra
has 114 limestone caves (UKSDA, 1999; Haznan,
2003). Limestone biodiversity consists of three types
of ecosystem, as troglobin, troglophil and trogloxene
(Dunn,1965). A common cave dweller belongs to
a group of bats known as trogloxene (Vermeullen
&Whitten, 1999) in the order of chiroptera (Findley,
1993; Kitchener, 1996; Nowak, 1994). Based on
echolocation calls chiroptera are divided into two sub
order, megachiroptera and microchiroptera (Gunnel &

9
2014 Journal of Indonesian Natural History Vol 2 No 2
Simmons, 2005; Koopman, 1994; Simmons & Geisler,
1998). Based on molecular evidence and evolution
of echolocation in bats, Koopman (1994) proposed
that Megachiroptera (family Pteropodidae) is closely
related to Microchiroptera (Rhinolipids group includes
Rhinolophidae, Megadermatidae, Hipposideridae,
Craseonycteridae, Rhinopomatidae) and grouped them
as Yinpterochiroptera, whereas Jones and Teeling
(2006) grouped Microchiroptera into two infraorders;
Yinochiroptera and Yangochiroptera. This group can
be found in all habitats with some families preferring
caves as roosting sites (Graham, 1994;Vermeullen &
Whitten, 1999).
H. diadema belongs to the family Hipposideridae
(Roundleaf bats), infraorder Yinochiroptera that is
sometimes called Diadem roundleaf bats. Description
of this species is large body size, FA 76-87 mm and
weight 30-47 gram. Fur of upperparts is dark brown
with pale bases, white patches on the shoulders and
sides; underparts greyish-white. In adult females orange
or orange bu often replaces the white. Noseleaf with
3 or 4 lateral leaets; posterior noseleaf large and
rounded (Francis, 2008) (Figure 1). This species has
wide distribution from Burma and Vietnam through
Thailand, Laos, West Malaysia and Indonesia (including
Sumatra, Borneo, and Bali) to New Guinea, Bismarck
Archipelago, Solomon Islands and norteasth Australia;
Philippines; Nicobar Islands (Simmons, 2005).
West Sumatra is separated by Bukit Barisan that
stretches from south to north of Sumatra Island. This
was formed during Miocene when two unequal parts,
the narrow west coast and the wider half of hills and
alluvial areas. The dierent ecological conditions
east and west of Bukit Barisan is likely to have
inuenced the morphology and genetic variation of
the species (Colombijn, 2005; Whitten, 1989). Studies
about morphological traits and genetical variations
associated with ecological conditions suggest that H.
diadema distributed across several small and large
islands (include in Lesser Sunda Islands) belong to 16
dierent subspecies (Kitchener et al., 1992). Rahman
and Abdullah (2010) found that Penthetor lucasi in
three geographical areas of Sarawak (Malaysia) diers
in body size and exhibit strong sexual dimorphism in
certain characters. Benita (2012) studied morphological
variations of Hipposideros larvatus from three caves
in west Sumatra and concluded that the barrier created
by Bukit Barisan mountain range may have lead to
the variation in morphological characters of bats in
Sumatra.
Currently, there is no other published study that focuses
on morphological variation of H. diadema from caves
in West Sumatra. Tate (1941) summarized information
about the subspecies of H.diadema in the Indo-Australian
region and recognized about 16 subspecies. This,
however, does not include subspecies grouping of H.
diadema populations in west Sumatra. We hypothesized
that ecological dierences between east and west Bukit
Barisan may have induced morphological variation
among population of H. diadema. Therefore, the aim of
this study is to investigate the morphological variation
of population H. diadema from three caves in West
Sumatra separated by Bukit Barisan mountain range.
Figure 1. H. diadema collected from Kalilawa Cave, Padang, West Sumatra.
Morphological variation of leaf-nosed bats

10 © University of Andalas / Copenhagen Zoo

Bats were captured from three caves in West Sumatra:
Kalilawa Cave, Padang (00o56’51.1S, 100o29’50.2E);
Lereng Cave, Pariaman (00o92’95.8S, 100o33’89.4E)
and Salamaik Cave, Sawahlunto (00o40’11.6S,
100o44’24.1E) (Figure 2). Bats were captured using
harp traps (Francis, 1989) setup in entrances of the caves.
The traps were deployed in the afternoon and checked
in the evening and in the early morning. For each bat
were captured, we recorded the age (adult or young)
and sex. Presence of growth bands at the nger joints
of H. diadema were also recorded (Anthony, 1988).
Bat trapping took place during January-February 2013
and the specimens were deposited in the Zoological
Museum at the University of Andalas (MZUA).
Thirty six characters were measured; twenty one
external characters following Rahman and Abdullah
(2010), and fteen skull character following Kitchener
and Maryanto (1993). These external characters
measurements were as follows, with abbreviations
in parentheses; ear length (E), head and body length
(HB), tail to ventral length (TV), Forearm length (FA),
tibia length (TB), rst digit length (PIB), hind foot
length (HF), second digit metacarpal (D2MCL), third
digit metacarpal (D3MCL), fourth digit metacarpal
(D4MCL), fth digit metacarpal (D5MCL), third digit
rst (D3P1L) and second phalank length (D3P2L),
fourth digit rst (D4P1L) and second phalank length
(D4P2L), fth digit rst (D5P1L) and second phalank
length (D5P2L), antitragus high (TA), eye diameters
(DM), posterior nose leaf breadth (LDP) and anterior
nose leaf width (LDA). The skull characters measured
were the great skull length (GSL), cranial length (PIL),
least interorbital width (LI), zygomatic width (LTP),
width across caninus to another caninus from outer basal
face (CCB), palatal bridge length (PBL), width across
molar to another molar from outer mass face (MMB),
tymphanic bulla length (TBL), tymphanic bulla width
(TBB), cochlea width (CW), cranial heigh (CH), rostrum
heigh (RH), rostrum length (RL), lower tooth row length
(IML) and dentary length (DL) (Figure 3).
Figure 2. Locality of H. diadema specimens used in this study
in West Sumatra (insert), Sumatra Island, Indonesia.
Figure 1. A diadem leaf-nosed bat, H. diadema, from Kalilawa cave, Padang, West Sumatra.
Chornelia et al.

11
2014 Journal of Indonesian Natural History Vol 2 No 2
The data measurements were divided by forearm
(external measurements) and great skull length
(skull measurements) to standardised body size for
all specimens. Morphological variations among the
populations were tested using Kruskall-Wallis Test and
possible dierences between populations were tested
using Mann Whitney U Test at a signicancy level of
5% using SPSS® software. All data were transformed
to log10 values before Principal Component Analyses
(PCA) and Cluster analysis shown up by UPGMA
(Unweighted Pair Group Method Arithmatic Average)
using MVSP 3.1 and NTSyspc Ver 2.0.2i software.

A total of 58 adult H. diadema consisting of 31 males
and 27 females were collected and measured. The
number of specimens collected from each cave was:
Kalilawa cave (14 male and 4 female), Lereng cave (3
male and 11 female) and Salamaik cave (14 male and
12 female).
Morphological characters
The morphological characters of male and female
specimens from Salamaik Cave population are relative
B
Figure 3. Twentyone external characters (a,b) (modied from Rahman and Abdulllah, 2010) and een skull
characters (a;dorsal, b; ventral, c; lateral) (modied from Kitchener and Maryanto, 1993) were used in this study to
measure morphological dierences.
A
CD E
Morphological variation of leaf-nosed bats

12 © University of Andalas / Copenhagen Zoo
Characters Kalilawa cave
(N=14)
Lereng Cave
(N=3)
Salamaik cave
(N=14) Kruskall-Wallis test
E 31.48 ± 2.36 31.5 ± 3.07 28.44 ± 1.56 H = 10.09; p = 0.006*
HB 100.92 ± 6.9 101.75 ± 6.83 102.66 ± 3.17 H = 0.142; p = 0.9313 ns
TV 56.98 ± 3.34 57.87 ± 3.48 52.40 ± 3.22 H = 10.66; p = 0.005*
FA 78.37 ± 2.48 78.35 ± 2.07 75.92 ± 2.39 H = 8.385; p = 0.015*
TB 43.43 ±2.62 41.09-1.68 40.53 ± 2.32 H = 10.8; p = 0.005*
PIB 12.84 ±1.16 11.60 ± 1.15 10.81 ± 0.86 H = 17.37; p = 0.000*
HF 15.39 ± 1.69 14.65 ±1.19 15.80 ± 1.56 H = 0.431; p = 0.806 ns
D2MCL 77.58 ± 3.27 77.64 ± 3.92 71.03 ± 2.68 H = 18.39; p = 0.000*
D3MCL 74.81 ±1.77 73.79 ±1.26 73.92 ± 2.02 H = 1.273; p = 0.529 ns
D4MCL 72.97 ±3.06 72.35 ±2.46 73.04 ± 2.45 H = 0.264; p = 0.876 ns
D5MCL 66.27 ± 2.10 65.77 ± 2.28 61.53 ± 2.17 H = 16.27; p = 0.000*
D3P1L 33.75 ± 3.01 33.43 ±1.21 27.79 ± 15.85 H = 7.861; p = 0.019*
D4P1L 24.47 ± 2.22 23.59 ±3.03 24.09 ± 2.33 H = 0.188; p = 0.910 ns
D5P1L 26.35 ± 1.93 26.09 ±1.85 26.37 ± 1.17 H = 0.136; p = 0.934 ns
D3P2L 36.33 ± 2.20 34.87 ±1.68 33.45 ± 1.35 H = 11.08; p = 0.003*
D4P2L 21.90 ±7.95 17.21 ±1.71 17.70 ± 1.17 H = 5.768; p = 0.056 ns
D5P2L 20.28 ± 1.96 19.74 ±0.07 18.34 ± 1.26 H = 7.004; p = 0.030*
TA 6.71 ± 1.22 5.27 ±0.31 5.71 ± 0.67 H = 8.149; p = 0.017*
DM 3.12 ± 0.46 2.97 ±0.53 2.40 ±0.34 H = 15.64; p = 0.000*
LDP 14.71 ± 1.41 14.47 ±1.64 16.16 ± 0.54 H = 13.07; p = 0.001*
LDA 14.36 ± 0.86 13.49 ±0.40 14.93 ± 0.60 H = 5.74; p = 0.057ns
GSL 31.57 ± 0.71 31.97 ± 1.49 30.56 ± 0.54 H = 12.77; p = 0.002*
PIL 68.25 ± 6.71 66.03 ± 3.85 69.02 ± 7.75 H = 0.195; p = 0.907 ns
LI 12.66 ± 1.08 12.33 ± 1.97 12.63 ± 1.66 H = 0.160; p = 0.923 ns
LTP 53.43 ± 8.20 54.35 ± 2.10 55.78 ± 1.99 H = 2.452; p = 0.294 ns
CCB 25.56 ± 1.51 24.82 ±1.89 24.71 ± 1.89 H = 1.147; p = 0.564 ns
PBL 30.04 ± 0.88 31.19 ± 2.92 30.23 ± 2.83 H = 1.449; p = 0.485 ns
MMB 39.48 ± 1.84 38.31 ± 3.26 39.98 ± 1.71 H = 1.917; p = 0.384 ns
TBL 13.79 ± 1.58 12.86 ± 0.88 13.62 ± 1.47 H = 1.571; p = 0.456 ns
TBB 45.04 ± 0.94 44.83 ± 2.84 45.68 ± 1.75 H = 0.5819; p = 0.747 ns
CW 10.63 ± 1.49 10.95 ± 0.92 10.17 ± 1.20 H = 0.927; p = 0.629 ns
CH 38.06 ± 2.61 35.69 ± 2.28 36.59 ± 20.41 H = 3.555; p = 0.169 ns
RH 30.62 ± 1.19 30.3 ± 1.44 29.72 ± 1.38 H = 2.059; p = 0.357 ns
RL 14.61 ± 1.88 16.03 ± 1.85 13.28 ± 1.17 H = 9.872; p = 0.007*
IML 45.63 ± 2.63 44.42 ± 2.81 46.45 ± 2.05 H = 2.061; p = 0.357 ns
DL 68.66 ± 2.02 64.74 ± 5.83 69.23 ± 2.21 H = 1.831; p = 0.400 ns
Table 1. Measurement of male samples in millimeters, for (a) external character measurements (b) Skull character
measurement. For each samples, mean ± standard deviaon, df=2, N=sample size, p-value, H=Kruskall-Wallis value.
*=signicance level p≤0.05, ns=not signicant)
B) Skull measurements
A) External characters measurements
Chornelia et al.

13
2014 Journal of Indonesian Natural History Vol 2 No 2
Characters Kalilawa cave
(N=14)
Lereng Cave
(N=3)
Salamaik cave
(N=14) Kruskall-Wallis test
E 33.75 ± 2.17 31.99 ±1.83 29.15 ± 1.92 H = 13.87; p = 0.000*
HB 104.45 ± 4.19 105.73 ± 2.91 104.38 ± 4.11 H = 0.8009; p = 0.67 ns
TV 58.44 ± 0.40 57.48 ± 4 49 54.20 ± 3.68 H = 4.325; p = 0.115 ns
FA 75.03 ± 0.77 78.39 ± 1.79 78.33 ± 1.91 H = 8.62; p = 0.013*
TB 44.40 ± 0.79 42.22 ± 2.47 39.40 ± 0.93 H = 14.18; p = 0.000*
PIB 12.95 ± 1.76 12.67 ± 1.55 11.63 ± 1.35 H = 3.819; p = 0.148 ns
HF 18.57 ± 0.43 16.76 ± 1.52 15.41 ± 1.23 H = 11.43; p = 0.003*
D2MCL 82.79 ± 2.30 79.1 ± 2.91 82.69 ± 3.40 H = 15.51; p = 0.000*
D3MCL 77.18 ± 1.38 75.16 ± 2.67 73.66 ± 1.43 H = 7.528; p = 0.023*
D4MCL 73.69 ± 0.99 74.01 ± 2.23 71.18 ± 1.68 H = 11.67; p = 0.003*
D5MCL 69.19 ± 2.08 68.72 ± 3.29 60.85 ± 1.39 H = 16.09; p = 0.000*
D3P1L 34.66 ± 0.26 33.29 ± 11.12 31.31 ± 1.95 H = 15.62; p = 0.000*
D4P1L 23.85 ± 1.84 26.56 ± 4.03 24.95 ± 0.88 H = 3.155; p = 0.206 ns
D5P1L 27.02 ± 0.54 25.93 ± 1.96 26.81 ± 1.28 H = 1.194; p = 0.551 ns
D3P2L 38.25 ± 0.18 37.07 ± 5.81 32.25 ± 1.40 H = 15.73; p = 0.000*
D4P2L 17.60 ± 0.75 20.92 ± 9.12 17.20 ± 0.86 H = 4.216; p = 0.122 ns
D5P2L 21.03 ± 0.15 19.79 ± 1.87 18.88 ± 8.73 H = 12.94; p = 0.002*
TA 7.14 ± 0.26 7.16 ± 3.61 5.49 ± 1.17 H = 4.283; p =0.118 ns
DM 3.33 ± 0.05 3.28 ± 1.17 2.50 ± 0.39 H = 10.13; p = 0.006*
LDP 16.60 ± 0.99 14.06 ± 0.67 15.98 ± 0.75 H = 17.33; p = 0.000*
LDA 14.57 ± 0.98 14.50 ± 5.12 14.95 ± 0.54 H = 4.109; p = 0.128 ns
GSL 32.13 ± 0.51 31.99 ± 0.44 31.75 ± 0.58 H = 1.869; p = 0.393 ns
PIL/GSL 64.42 ± 1.21 66.02 ± 7.32 63.83 ± 1.49 H = 4.242; p = 0.119 ns
LI/GSL 12.59 ± 0.60 13.12 ± 2.36 11.15 ± 1.46 H = 5.717; p = 0.057 ns
LTP/GSL 53.72 ± 2.27 55.32 ± 2.15 52.66 ± 1.53 H = 7.225; p = 0.027*
CCB/GSL 24.69 ± 0.35 25.99 ± 1.84 23.19 ± 1.53 H = 14.61; p = 0.000*
PBL/GSL 29.23 ± 0.80 31.03 ± 1.67 27.76 ± 1.26 H = 15.15; p = 0.000*
MMB/GSL 39.29 ± 0.42 38.81 ± 1.99 37.71 ± 1.20 H = 2.165; p = 0.339 ns
TBL/GSL 11.71 ± 0.50 13.47 ± 2.39 9.94 ± 1.00 H = 17.2; p = 0.000*
TBB/GSL 44.32 ± 0.78 45.15 ± 2.68 43.92 ± 1.63 H = 1.022; p = 0.600 ns
CW/GSL 12.14 ± 1.44 11.26 ± 2.88 11.31 ± 7.02 H = 10.52; p = 0.005*
CH/GSL 35.16 ± 1.70 38.21 ± 2.80 34.21 ± 2.19 H = 13.44; p = 0.001*
RH/GSL 30.46 ± 0.53 30.99 ± 2.18 26.73 ± 4.82 H = 14.5; p = 0.000*
RL/GSL 12.80 ± 1.77 29.56 ± 14.81 12.80 ± 1.22 H = 13.75; p = 0.001*
IML/GSL 41.97 ± 4.49 47.58 ± 2.52 40.73 ± 0.70 H = 15.61; p = 0.000*
DL/GSL 66.13 ± 0.60 66.17 ± 4.06 65.70 ± 1.55 H = 0.914; p = 0.6343 ns
Table 2. Measurement of female samples in millimeters for (a) external character measurements (b) Skull character
measurement. For each samples, mean ± standard deviaon, df=2, N=sample size, p-value, H=Kruskall-Wallis value.
*=signicance level p≤0.05, ns=not signicant)
B) Skull measurements
A) External characters measurements
Morphological variation of leaf-nosed bats

14 © University of Andalas / Copenhagen Zoo
small compared to the specimens from Kalilawa cave
and Lereng cave. The sample size, mean, standard
deviation, maximum and minimum values for all
characters measurements, including external and skull
characters measurements of H. diadema are presented
in Table 1 and 2. Comparison of adult specimens among
the tree population showed signicant dierences
among them in 15 morphometric characters among male
consist of 13 external characters and 2 skull characters,
and 21 morphometric character measurements of
female, consist of 13 external characters and 9 skull
characters. We recorded signicant dierences in male
externall characters for TV, FA, TB, PIB, D2MCL,
D5MCL, D3P1L, D3P2L, D5P2L, TA, DM, LDP,
and for skull characters GSL and RL. For females we
recorded signicant dierences for D2MCL, E, FA, TB,
HF, D3MCL, D5MCL, D3P1L, D3P2L, D5P2L, DM,
LDP, and for skull characters LTP, CCB, TBL, CW, CH,
RH, RH, RL, and IML. Signicant dierences in both
characters measurements indicates high divergence of
external and skull characters of H. diadema between
the three populations (Kalilawa cave, Lereng cave, and
Salamaik cave).
A Mann Whitney U-test was used to compare remaining
characters. The H. diadema populations from Lereng and
Salamaik caves diers signicantly for both males and
females, This was also the case between the Kalilawa and
Lereng cave populations, and Kalilawa and Salamaik
cave populations. The males from Kalilawa cave and
Lereng cave only diered signicantly on one character,
whereas females diered on two 2 characters. Males
and females from Kalilawa and Salamaik caves diered
signcantly on two characters. The male populations of
Lereng and Salamaik caves diered signicantly on three
characters and in ve characters for females (Table 3).
Unweighted Pair Group Method Arithmatic Average and
Principall Component Analysis
Euclidian distances showed up in a UPGMA analysis
as clusters between three population with males and
females analysed separately. PCA of 36 characters
revealed a clear separation between the three dierent
cave populations (Fig. 4).
UPGMA analysis revealed a close relationship between
Padang and Pariaman populations (0.25 (male) and
0.14 (female)), and PCA showed that of H. diadema
populations from Padang were closely related to the
Pariaman populations, and clearly distinct from the
Sawahlunto populations.

H. diadema populations from three caves separated from
each other by Bukit Barisan in West Sumatra revealed
variations and morphological character divergences. Our
data suggest that the individuals from Kalilawa cave are
more closely related to individuals from the Lereng cave,
whereas it diered from Salamaik cave. From 21 external
and 15 skull characters used in this analysis the H.
diadema population from Salamaik cave are signicantly
smaller than conspecics from Kalilawa and Lereng
caves. Ecological circumstances related with breeding,
foraging, crowding and resources avaibility may dier
between the three populations due to their separation
by the Bukit Barisan range. Kalilawa and Lereng caves
are located on the western side of Bukit Barisan and at
a lower altitude than the Salamaik cave on the eastern
side. The ecological dierent conditions, combined
with a lower inter-population migration, may have
required dierent behavioural adaptation and resulting
morphological variations. Euclidian distance among H.
diadema populations from the three study sites showed
that geographic distance is reected in the relationship
distance. Kalilawa cave is closer geographically and in
relationship distance to Lereng cave than to Salamaik
cave.
Rahman and Abdullah (2010) reported morphological
variations between geographical separated populations
of Penthetor lucasi in Sarawak and suggested that
ecological conditions as the likely main cause of the
dierentiation. Kitchener and Suyanto (1996) suggest
that the Pleistocene- modern time island arrangement
have caused relatively recent morphological changes.
Kitchener, Konishi and Suyanto (1996) assumed
that longitude was the most important variable when
predicting overall skull and body size. In contrast Whitten
(1987) argued that Bukit Barisan was formed already
during Miocene, and therefore separated populations
of H. diadema in West Sumatra at a much earlier stage.
Kitchener et al., (1992) noted that H. diadema in Lesser
Sunda Island was divided into three phenetic grouping
based on external and skull measurements: H.d. diadema,
H.d. reginae and H.d. masoni in one group; H.d. griseus
and H.d. oceanitis in a second group, and H.d.nobilis in
a separate cluster. The study indicated that the eastern
form of H. d. diadema is smaller than the western
form, suggesting dierent ecological conditions had
required dierent adaptational strategies and eventually
morphological unique forms.
Chornelia et al.

15
2014 Journal of Indonesian Natural History Vol 2 No 2
Whereas morphological variation could give rise
to speciation, we were unable to determine from
morphological characters alone that individuals of H.
diadema from the three dierent study sites belong
to dierent subspecies. Further studies on the genetic
variation of H. diadema in West Sumatra is needed to
conrm if there are indeed three dierent subspecies.

We gratefully acknowledge the support of all teachers at
Biology Department, Faculty of Mathematic and Natural
Science, Andalas University especially to Genetics and
Cytology Laboratory, Biologi Department and Museum
Zoology Universitas Andalas, to Dr. Syaifullah, Dr.
Rizaldi, Dr. Wilson Novarino and all assistants. Thank
you to our colleagues who assisted us in the eld, KCA-
LH Raesia FMIPA UNAND, particularly Fajri, Beny
Ramdani, Rezi Rahmi Amolia, Vivi Martinsyah, Riki
Novtian Burlis, Reki Kardiman, Heru Handika, Kedhy
Lavandino and all members. Thanks a lot to all friends
who supported this research; Jiji, Rahma, Widia, Nova,
Putri, Fitri, Anita, Wita, Ami, Icha and Nurul.
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