ArticlePDF Available

Food Habits of Asian Elephants Elephas maximus in a Rainforest of Northern Peninsular Malaysia

  • Department of Wildlife & Nat. Parks Peninsular Malaysia

Abstract and Figures

Little is known about the food habits of Asian elephants (Elephas maximus) in tropical rainforests of Southeast Asia. In Peninsular Malaysia, elephant habitat has been extensively modified by human intervention in the past few decades. Most of the primary forest has been logged or given way to plantations, infrastructure, and human inhabitation. Here we compare the food habits of wild elephants in three habitats of Belum-Temengor Forest Complex (BTFC): (1) primary forest, (2) selectively-logged forest, and (3) by the side of a road that bisects the forest complex. We used microhistological fecal analysis to describe elephants' diet. Elephant dung in the primary forest was mainly composed of non-grass monocotyledonous leaves (22%), woody debris (32%), and woody fiber (20%). Those in the logged forest were similar; non-grass monocotyledonous leaves accounted for 33%, woody debris for 24%, and fiber for 26%. At the roadside, elephant dung was dominated by grasses (47%). We conclude that by the road elephants shift their diet into grasses, suggesting that the road acts like a large forest gap, promoting the availability of grasses and other early succession plants. Elephant feeding by the road poses potential conservation conflicts by means of road accidents and increased contact with people.
Content may be subject to copyright.
Mammal Study 41: 155–161 (2016)
© The Mammal Society of Japan Short communication
Food habits of Asian elephants Elephas maximus in a rainforest
of northern Peninsular Malaysia
Shiori Yamamoto-Ebina1, Salman Saaban2, Ahimsa Campos-Arceiz3,* and Seiki Takatsuki1
1 School of Veterinary Medicine, Azabu University, Japan
2 Biodiversity Conservation Division, Department of Wildlife and National Parks Peninsular Malaysia, Malaysia
3 School of Geography, University of Nottingham Malaysia Campus, Malaysia
Abstract. Little is known about the food habits of Asian elephants (Elephas maximus) in tropical
rainforests of Southeast Asia. In Peninsular Malaysia, elephant habitat has been extensively modied
by human intervention in the past few decades. Most of the primary forest has been logged or given
way to plantations, infrastructure, and human inhabitation. Here we compare the food habits of wild
elephants in three habitats of Belum-Temengor Forest Complex (BTFC): (1) primary forest, (2)
selectively-logged forest, and (3) by the side of a road that bisects the forest complex. We used
microhistological fecal analysis to describe elephants’ diet. Elephant dung in the primary forest was
mainly composed of non-grass monocotyledonous leaves (22%), woody debris (32%), and woody ber
(20%). Those in the logged forest were similar; non-grass monocotyledonous leaves accounted for
33%, woody debris for 24%, and ber for 26%. At the roadside, elephant dung was dominated by
grasses (47%). We conclude that by the road elephants shift their diet into grasses, suggesting that the
road acts like a large forest gap, promoting the availability of grasses and other early succession plants.
Elephant feeding by the road poses potential conservation conicts by means of road accidents and
increased contact with people.
Key words: diet, forest fragmentation, infrastructure, megafauna, selectively-logged forest.
Asian elephants (Elephas maximus) are endangered
(Choudhury et al. 2008) with a wild population estimated
in 39,000–47,000 individuals (Fernando and Pastorini
2011) in a range of ca 450,000 km2 in South and South-
east Asia (Sukumar 1989). Within the range, protected
areas cover only 72,000 km2 (~16% of the total) and hence
further population declines are expected (Leimgruber et
al. 2003).
As the largest terrestrial animals, elephants have huge
demands for food, consuming as much as 150 kg of fod-
der per day (Vancuylenberg 1977), and large home ranges
(e.g., Alfred et al. 2012). The need to nd large amounts
of food determines elephants ranging patterns and other
aspects of their behavior. Understanding Asian elephant
feeding ecology and food habits is thus essential for the
conservation of the species, especially in the increasingly
human-modied landscapes they inhabit.
Elephants in Southeast Asia feed on a wide range of
plant types, often showing preference for grasses and
other monocotyledonous plants (Olivier 1978; English
et al. 2014a, 2014b); studies in southern China and
Myanmar showed that elephants feed on nutritious foods
other than grasses (Chen et al. 2006; Campos-Arceiz et al.
2008). Asian elephants are also known to consume a
range of eshy fruits, hence playing an important role as
agents of seed dispersal (Campos-Arceiz and Blake
2011). Nevertheless, information of elephant food habits
in Southeast Asian rainforests is still quite limited, and
further quantitative analyses are required.
Peninsular Malaysia, with a population of ca 1,500 wild
elephants (Saaban et al. 2011) is one of the strongholds of
elephant conservation in Southeast Asia and has experi-
enced a very rapid transformation that led to a decrease in
forest cover from nearly 90% in 1950 (FDTCP 2009) to
less than 40% in 2010 (Miettinen et al. 2011). Elephant
habitats in Malaysia have been replaced by forest planta-
*To whom correspondence should be addressed. E-mail:
156 Mammal Study 41 (2016)
tions, mostly of rubber (Hevea brasiliensis) and oil palm
(Elaeis guineensis), dams, urban areas, and human infra-
structure (Daim 1995). Elephants now often enter into
plantations and raid crops, resulting in the so-called
human-elephant conict—currently the main threat for
Asian elephants in Malaysia (Saaban et al. 2011; Campos-
Arceiz 2013).
Tropical forests such as those in Peninsular Malaysia
have gaps formed after felling of large trees, resulting in
the invasion of bamboos and ginger shrubs. It is thus
expected that forest-dwelling Malaysian elephants can
feed on such gap-specialist plants as found in riverine
habitats of the Kinabatangan river, in Borneo (English et
al. 2014a, 2014b). If primary forests are logged, or a road
or a plantation is created contiguous to the forest, this will
result in open patches and forest edges. These changes
would facilitate the growth of grasses and forbs by
increasing light availability.
The objective of this study is to provide a quantitative
description of the food habits of elephants in a Malaysian
rainforest environment and compare the diet composition
of elephants living in primary forest, selectively logged
forest, and by the side of a road bisecting the forest.
Materials and methods
Study area
This study was conducted in Belum-Temengor Forest
Complex (BTFC), Perak, northern Peninsular Malaysia
(Fig. 1). BTFC is one of the largest blocks of continuous
forests in Peninsular Malaysia; it is contiguous with
Halabala Wildlife Sanctuary and Bang Lang National
Park, in southern Thailand, and forest areas in the Malay-
sian states of Kedah and Kelantan. BTFC is a hilly area
with an altitudinal range of 130–1,500 m above sea level
and vegetation that includes lowland dipterocarp, hill
dipterocarp, and montane forests. Much (152 km2) of
BTFC landscape is submerged under Tasik Temengor, a
large man-made lake dammed in the late 1970s. BTFC is
rich in biodiversity, including over 3,000 species of
owering plants, 185 bird species, and a wide range of
endangered mammals.
From a management point of view, BTFC is divided in
two blocks—the Royal Belum State Park in the north
and the Temengor Forest Reserve in the south—bisected
by the East-West highway (Federal Route 4; Fig. 1).
Royal Belum was gazetted in 2007 and with an area of
Fig. 1. Map of Peninsular Malaysia with a close-up of Belum-Temengor Forest Complex, where this study took place. White dots indicate roughly
the sampling areas.
Yamamoto-Ebina et al., Diet of Malaysian elephants 157
1,175 km2. Most of Royal Belum has never been logged.
Temengor on the other hand is a production forest where
logging is allowed and currently ongoing. The East-West
highway is a two-lane road that connects the ~125 km
between the towns of Gerik and Jeli. This road was com-
pleted in 1975 and a band of 3 km to the north and south
of the road was logged in 1970–1995 (MNS 2009). Ele-
phants are frequently seen crossing the road and feeding
in the grasslands on the side of the East-West highway.
Sample collection and analysis
Elephant dung samples were collected in three kinds
of environments within BTFC: (1) in primary forests of
Royal Belum, (2) in selectively logged forests of
Temengor, and (3) by the road side (Fig. 1). We col-
lected a total of 30 independent samples, nine from the
primary forest, 10 from the logged-forest, and 11 from
the roadside. Each sample was collected from a separate
dung pile, from which we collected a spoonful amount
of dung from the internal part of one dung bolus. The sam-
ples were collected in June-July 2013 and preserved in
60% ethanol for subsequent analysis.
We used microhistological fecal analysis (Stewart
1967) and the point-frame method (Chamrad and Box
1964) to produce a quantitative description of the dung
composition. This approach is widely used for ruminants
(e.g., Campos-Arceiz et al. 2004). The principle is consis-
tently usable for elephant dung too. We used a slide glass
gridded with a 1.0 mm aperture and sizing 40 mm by 80
mm. On the slide glass, a frame of 20 × 50 mm was set,
and we put fecal contents into the frame. Because this
method can only be used to analyze small particles and
elephants often defecate large indigested food fragments,
we cut dung contents into smaller pieces with scissors.
They were washed over a sieve of 0.5 mm aperture and
the retained fragments were spread over the slide glass,
and analyzed under a binocular microscope at a magni-
tude of ×40 and ×100. Each fragment covering the cross-
ing point of the grid was identied and counted. We
counted up to 200 points and this process was replicated
three times with each sample and the results averaged
(Takatsuki and Tatewaki 2012).
Initially we categorized plant fragments into 14 groups
but since some of these categories were very rare, we
grouped some of them together, reducing the nal list to
nine categories: (1) grass leaves, (2) monocotyledonous
leaves other than grasses, (3) the culms of grass, bamboo,
and ginger, (4) banana culms, (5) dicotyledonous leaves,
(6) woody debris (small broken debris of wood), (7)
woody ber (i.e. brous material of woody plants other
than leaf veins), (8) others (including fruit parts, seeds,
bark, and root), and (9) unidentied materials.
Data analyses
The fecal composition diversity at each habitat type
was expressed using the Shannon-Wiener’s diversity
index H’:
H’ =
Pi ln Pi
where Pi is the proportion of i in the fecal composition.
Fecal composition similarity across habitats was esti-
mated using Whittaker’s percentage of similarity (PS):
PS = 1
min (Pai, Pbi)
where Pai and Pbi were the proportion of food i at habitat
a, and habitat b, respectively.
Statistical analyses we conducted using R statistical
environment (R Core Team 2016). Differences in diver-
sity (H’) and percentage values of the major food catego-
ries were compared among the three habitats using the
Kruskal-Wallis test. When differences were signicant,
we conducted multiple comparison tests and assessed
signicance using the Dwass, Steel, Critchlow, Flinger W
statistic (function pSDCFlig in R’s NSM3 package with
method = “Asymptotic”; Schneider et al. 2015).
The fecal composition diversity index (H’) ranged from
1.38 to 1.48 and was not different across habitat types
(c2 = 4.6, df = 2, P = 0.098). Fecal composition was very
similar between primary and logged forest (PS = 73.5%)
and quite different between the logged forest and by the
roadside (PS = 36.6%). PS between the primary forest
and the roadside was 51.6%.
Overall, the diet of elephants in BTFC was dominated
by four food categories that accounted for roughly 80% of
the dung composition: grass leaves, the leaves of other
monocots (ginger, palms, and others), woody debris, and
woody ber. Other plant types and parts contributed
small amounts (Fig. 2). The distribution of the different
food categories was, however, very distinct across habitat
types (Fig. 2). Grass leaves, for example, contributed dif-
ferently to elephant diet across habitats, being the domi-
nant food category by the roadside (mean ± SD = 46.6 ±
13.3%) but almost absent in the logged forest (1.1 ± 2.1%;
158 Mammal Study 41 (2016)
Fig. 2). The contribution of grasses in the three habitats
differed signicantly among each other (Table 1). The
contribution of other monocot leaves also differed across
habitats showing the opposite trend to grass leaves—they
were most abundant in the logged forest (33.4 ± 12.9%)
and least by the roadside (3.5 ± 2.7%; Fig. 2). The dif-
ferences were also signicant (Table 1), although the
pair-wise comparisons suggest that only the difference
between logged forest and the roadside are signicant
(Table 1). The contribution of woody debris was also dif-
ferent: they accounted for 32.2 ± 10.7% and 23.5 ± 6.3%
at the primary and logged forest, respectively, but 10.7 ±
6.8% at the roadside (Fig. 2). Woody debris contribution
in the primary forest was greater than that of the logged
forest and the roadside, and in the logged forest greater
than in the roadside (Table 1). This was similar to woody
ber, which accounted for 20.3 ± 10.5% and 26.1 ± 12.6%
in the primary forest and the logged forest, and 13.1 ±
9.6% at the roadside. The value of woody ber by the
roadside was smaller than in the logged forest but there
were no differences between primary and logged forest,
nor between primary forest and the roadside (Table 1).
This is one of the few quantitative analyses of wild
elephant food habits in tropical rainforest environments
of Southeast Asia (see also Olivier 1978; English et al.
2014a, 2014b) and to our knowledge the rst to use dung
microhistological analysis to identify diet contents.
Forest elephants have the most diverse diets of any wild
herbivores (Blake 2002). Since direct observation of wild
elephant feeding is difcult in tropical rainforest, previ-
ous studies have generally relied on indirect observa-
tion of feeding signs (e.g., English et al. 2014a, 2014b).
Our approach—using microhistological analyses—has
allowed us to identify the relative importance of differ-
ent kinds of plant types as elephant food in the three
different habitats within BTFC. We found that although
food diversity—at the resolution we studied it—was not
different across habitats, the relative contribution of the
different food blocks were considerably different.
The main constituents of the elephant feces in the pri-
mary forest were grass leaves, monocot leaves, woody
debris, and woody ber. Dicot leaves unexpectedly ac-
Fig. 2. Elephant diet composition in three habitat types in Belum-Temengor Forest Complex based on microhistological analyses of elephant
dung. GBG culm = grass, bamboo, and ginger culm; others = fruit parts, seeds, bark, and root. Error bars represent standard errors.
Yamamoto-Ebina et al., Diet of Malaysian elephants 159
counted for a small proportion. It is unclear whether the
elephants did not feed on dicot leaves or digestibility of
them is higher than other food plants. As we expected,
grasses were less important in the forest than by the road
(Fig. 2). However, against our expectations, grasses were
more important in the primary than in the logged forest.
This is probably a sampling effect since the samples in
the primary forest were collected in areas near Temengor
Lake, where grasses are relatively common. It is interest-
ing that in both primary and logged forest habitats, non-
grass monocotyledonous plants including gingers and
palms were important for elephants. These plants grow in
gaps and other disturbed habitats, which again reinforces
the idea of elephants as edge (or gap) specialists (Campos-
Arceiz 2013). Gaps are relatively common in Southeast
Asian dipterocarp forests, where large trees tend to have
small root systems and often fall down producing forest
gaps. Elephants are likely to take advantage of these gaps
and other disturbed habitats such as streams and lake beds.
In the logged forest, monocot leaves and ber repre-
sented a higher proportion of the diet and grass leaves
were almost absent. This seems to reect well the habitat
vegetation where bamboos, gingers, and palms grow
abundantly while grasses are uncommon. The great
similarity between the primary and the logged forest is
attributed to the dominance of monocot leaves, woody
debris, and ber.
By the roadside, elephants were largely reliant on
grasses like Imperata spp., Panicum spp. and other mono-
cot plants that grow abundantly on the sides of the road.
The greater contribution of banana (Musa spp.) was also
unique to this habitat. These are probably wild bananas
that grow abundantly in disturbed habitats. Southeast
Asian rainforests are often described as ‘food deserts’,
where food for large herbivorous and frugivorous mam-
mals is very limited (e.g., Corlett 2007). This must be
especially the case for elephants. In this context, roads
like the East-West highway may be ecologically similar
to a very large gap, where grasses and other early succes-
sion plants grow abundantly. For elephants in BTFC, the
road probably represents a trade-off between abundance
of high quality food (mostly grasses) and high levels of
risk such as trafc and contact with people. Judging from
our casual observations (Campos-Arceiz personal obser-
vation) and data from GPS-telemetry (Campos-Arceiz et
al. unpublished data), some elephants in BTFC spend a
substantial amount of time near the East-West highway,
which brings potential conservation conicts and might
result in changes in the behavior and ecological function
of elephants.
Although our sample size was relatively small and our
sampling did not cover different seasons (e.g., fruiting
periods), this study proves the usefulness of the point-
frame method to study elephant food habits, and the
Table 1. Kruskal-Wallis and Dwass-Steel-Crichlow-Fligner multiple comparisons statistical test outputs for relative contribution of major food
categories to the diet of Asian elephants in three habitats of Belum-Temengor Forest Complex
Kruskal-Wallis test Dwass, Steel, Critchlow, Fligner
Food category chi-sq df P-value Pair W statistic P-value
Grass leaves 41.08 2 1.20E-09 PF-LF –4.63 0.003
PF-Rd 6.39 0
LF-Rd 7.89 0
Other monocot leaves 14.84 2 0.0006 PF-LF –0.021 0.99
PF-Rd –4.73 0.002
LF-Rd –4.95 0.001
Dicot leaves 10.66 20.005 PF-LF 4.54 0.004
PF-Rd 2.2 0.27
LF-Rd –2.64 0.15
Woody debris 34.06 24.02E-08 PF-LF –4.03 0.012
PF-Rd –6.88 < 0.001
LF-Rd –6.36 < 0.001
Woody ber 12.33 2 2.00E-03 PF-LF 1.76 0.43
PF-Rd –2.87 0.11
LF-Rd –4.94 0.001
PF = primary forest; LG = logged forest; and Rd = roadside.
160 Mammal Study 41 (2016)
results showed clear patterns. In our samples there was no
presence of fruit but a few months earlier than our sam-
pling, there was a fruiting episode of Mangifera sp. and
Irvingia malayana and, at that time, many elephant dung
piles contained fruit remains (Campos-Arceiz personal
observations). Longer-term analyses covering fruiting
episodes and wider range analyses covering different
habitat types are needed.
Our results provide interesting hints on one of the
causes of the widespread human-elephant conict that is
threatening the species throughout its range. Since ele-
phants are edge specialists (Campos-Arceiz 2013), they
readily use human-disturbed environments, including
food crops and newly planted rubber and oil palm planta-
tions. Attracted both by the crop and other early succes-
sion plants, elephants come into conict with farmers as
part of their natural optimal foraging strategy. Mitigating
human-elephant conict into human-elephant coexistence
is the most important challenge for Asian elephant con-
servation and will require a sound understanding of the
behavioral and ecological drivers of the conict (Campos-
Arceiz 2013).
In summary, elephants in the two forest types away
from the road fed mainly on monocot leaves other than
grasses and hard dicot material while those at the road-
side fed mainly on grasses. This supports the statements
that Asian elephants can elastically utilize forage plants
depending on habitats (Sukumar 1990; Mohapatra et al.
2013). The presence of roads like the East-West highway,
bisecting otherwise continuous patches of forest, modi-
es elephant food habits and creates conservation con-
icts. In this case, mechanisms must be put in place to
reduce the risk of road accidents and human disturbance
of elephants roaming by the roadside.
Acknowledgments: This study is part of the Manage-
ment & Ecology of Malaysian Elephants (MEME), a joint
research project between the Department of Wildlife and
National Parks (DWNP) Peninsular Malaysia and the
University of Nottingham Malaysia Campus. We are very
grateful to DWNP, and especially to its Director General
Dato’ Abdul Rasid Samsudin, for the permits to conduct
this research and for the continuous support in the eld.
SY is grateful to Malaysia’s Economic Planning Unit
(EPU) for granting her the necessary permit (UPE:
40/200/19/3003) to conduct eld research in Malaysia.
Field activities were generously nanced by grants from
Yayasan Sime Darby (grant M0005.54.04), Marinescape
Eco Aquariums (grant M0004.54.04), and Azabu Univer-
sity. Mr. Sampath K. K. Ekanayaka advised and supported
us for analysis; MEME members supported eld sam-
pling and logistics; Ms Kayal Vizi A/P Karupannan
helped SY to obtain research permits, and Y. Yamada
encouraged us during the study—we are very grateful to
all of them for their kind support.
Alfred, R., Ahmad, A. H., Payne, J., Williams, C., Ambu, L. N., How,
P. M. and Goossens, B. 2012. Home range and ranging behaviour
of Bornean elephant (Elephas maximus borneensis) females.
PLOS ONE 7: e31400. DOI: 10.1371/journal.pone.0031400.t007.
Blake, S. 2002. The Ecology of Forest Elephant Distribution and Its
Implications for Conservation. ICAPB, Edinburgh, University of
Edinburgh, 303 pp.
Campos-Arceiz, A. 2013. The dangerous myth of the Noble Beast.
Gajah 38: 5.
Campos-Arceiz, A. and Blake, S. 2011. Megagardeners of the forest—
the role of elephants in seed dispersal. Acta Oecologica 37: 542–
Campos-Arceiz, A., Lin, T. Z., Htun, W., Takatsuki, S. and Leimgruber,
P. 2008. Working with mahouts to explore the diet of work ele-
phants in Myanmar (Burma). Ecological Research 23: 1057–1064.
Campos-Arceiz, A., Takatsuki, S. and Lhagvasuren, B. 2004. Food
overlap between Mongolian gazelles and livestock in Omnogobi,
southern Mongolia. Ecological Research 19: 455–460.
Chamrad, A. D. and Box, W. T. 1964. A point frame for sampling
rumen contents. The Journal of Wildlife Management 28: 473–
Chen, J., Deng, X., Zhang, L. and Bai, Z. 2006. Diet composition and
foraging ecology of Asian elephants in Shangyong, Xishuangbanna,
China. Acta Ecologica Sinica 26: 309–316.
Choudhury, A., Lahiri Choudhury, D. K., Desai, A., Duckworth, J.
W., Easa, P. S., Johnsingh, A. J. T., Fernando, P., Hedges, S.,
Gunawardena, M., Kurt, F., et al. (IUCN SSC Asian Elephant
Specialist Group). 2008. Elephas maximus. In: IUCN (2014).
IUCN Red List of Threatened Species. Version 2014.1. Available
at (Downloaded on 25 October 2014).
Corlett, R. T. 2007. What’s so special about Asian tropical forests?
Current Science 10: 14–20.
Daim, M. S. 1995. Elephant translocation: The Malaysian approach.
Gajah 14: 43–48.
English, M., Ancrenaz, M., Gillespie, G., Goossens, B., Nathan, S. and
Linklater, W. 2014a. Foraging site recursion by forest elephants
Elephas maximus borneensis. Current Zoology 60: 551–559.
English, M., Gillespie, G., Ancrenaz, M., Ismail, S., Goossens, B.,
Nathan, S. and Linklater, W. 2014b. Plant selection and avoidance
by the Bornean elephant (Elephas maximus borneensis) in tropical
forest: does plant recovery rate after herbivory inuence food
choices? Journal of Tropical Ecology 30: 371–379.
FDTCP. 2009. CFS: Master Plan for Ecological Linkages. Final Report,
Federal Department of Town and Country Planning, Peninsular
Malaysia. Unpublished report. Kuala Lumpur, Malaysia.
Fernando, P. and Pastorini, J. 2011. Range-wide status of Asian
elephants. Gajah 35: 15–20.
Leimgruber, P., Gagnon, B., Wemmer, C., Kelly, D. S., Songer, M. A.
and Selig, E. R. 2003. Fragmentation of Asia’s remaining wild
lands: implications for Asian elephant conservation. Animal
Conservation 6: 347–359.
Miettinen, J., Shi, C. and Liew, S. C. 2011. Deforestation rates in
Yamamoto-Ebina et al., Diet of Malaysian elephants 161
insular Southeast Asia between 2000 and 2010. Global Change
Biology 17: 2261–2270.
MNS. 2009. A Study on the Effect of Conversion of Natural Forest to
Plantations along the East-West Highway in the Belum-Temengor
Forest Complex. Kuala Lumpur.
Mohapatra, K. K., Patra, A. K. and Paramanik, D. S. 2013. Food and
feeding behavior of Asiatic elephant (Elephas maximus Linn.) in
Kuldiha Wildlife Sanctuary, Odisha, India. Journal of Environ-
mental Biology 34: 87–92.
Olivier, R. C. D. 1978. On the Ecology of the Asian elephant. Unpub-
lished PhD Thesis, Universal of Cambridge.
R Core Team. 2016. R: A Language and Environment for Statistical
Computing. R Foundation for Statistical Computing, Vienna,
Austria. Available at (Accessed 1 June
Saaban, S., Othman, N., Yasak, M. N. B., Nor, B. M., Zar, A. and
Campos-Arceiz, A. 2011. Current status of Asian elephants in
Peninsular Malaysia. Gajah 35: 67–75.
Schneider, G., Chicken, E. and Becvarik, R. 2015. NSM3: Functions
and Datasets to Accompany Hollander, Wolfe, and Chicken—
Nonparametric Statistical Methods, Third Edition. R package ver-
sion 1.5. Available at
(Accessed 1 June 2016).
Stewart, D. R. M. 1967. Analysis of plant epidermis in feces: a tech-
nique for studying the food preferences of grazing herbivores.
Journal of Applied Ecology 4: 83–111.
Sukumar, R. 1989. The Asian Elephant: Ecology and Management.
Cambridge University Press.
Sukumar, R. 1990. Ecology of the Asian elephant in southern India. II.
Feeding habits and crop raiding patterns. Journal of Tropical
Ecology 6: 33–53.
Takatsuki, S. and Tatewaki, T. 2012. Applicability of the point-frame
method as a food habit analysis method for omnivorous mam-
mals: a case study on medium-sized carnivores. Honyuruikagaku
[Mammalian Science] 52: 167–177 (in Japanese with English
Vancuylenberg, B. W. B. 1977. Feeding behaviour of the Asiatic
elephant in Southeast Sri Lanka in relation to conservation.
Biological Conservation 12: 33–54.
Received 22 December 2015. Accepted 4 June 2016.
Editor was Masaharu Motokawa.
... Asian elephants' preferred food are monocot plants, particularly grasses (e.g., [105,106]). During the Holocene, as the grasslands of the Malay Peninsula were replaced by forests, the populations of palms remained stable [21]. ...
... In the Malay Peninsula, elephants eat all four species of cultivated tubers but Olivier's [36] elephant food list does not include observations of any of the wild species. Other studies on the diet of Asian elephants also do not mention wild tubers [106,111,112]. L. Ong (pers. ...
Full-text available
Understanding the relationship between humans and elephants is of particular interest for reducing conflict and encouraging coexistence. This paper reviews the ecological relationship between humans and Asian elephants (Elephas maximus) in the rainforests of the Malay Peninsula, examining the extent of differentiation of spatio-temporal and trophic niches. We highlight the strategies that people and elephants use to partition an overlapping fundamental niche. When elephants are present, forest-dwelling people often build above-the-ground shelters; and when people are present, elephants avoid open areas during the day. People are able to access several foods that are out of reach of elephants or inedible; for example, people use water to leach poisons from tubers of wild yams, use blowpipes to kill arboreal game, and climb trees to access honey. We discuss how the transition to agriculture affected the human–elephant relationship by increasing the potential for competition. We conclude that the traditional foraging cultures of the Malay Peninsula are compatible with wildlife conservation.
... However, in a fragmented landscape, elephants have been found to utilise various types of land cover, from closed canopy to open area (Sitompul et al. 2013a, b;Wadey et al. 2018). Their distribution is mainly driven by the availability of food and water resources, which is what attracts them to agricultural and horticultural land, idle grassland, water bodies and cleared land (Sitompul et al. 2013a, b;Chew et al. 2014;Yamamoto-Ebina et al. 2016;Neupane et al. 2019), while urban areas, settlements and associated non-agricultural areas are usually avoided (Wadey et al. 2018;Sharma et al. 2020). The response curve for type of land use may be misleading due to the correlation between the variables (Phillips et al. 2017). ...
... Although land use is one of the least important variables, habitat loss has been identified as the main driver of elephant displacement (Zahari et al. 2001), which occurs as a result of the conversion of forest to agricultural or urban uses. Moreover, vegetation type also varies according to the type of land use, impacting elephant diet (Yamamoto-Ebina et al. 2016). ...
Full-text available
The Asian elephant has been listed as an endangered species on the International Union for Conservation of Nature Red List since 1986. In Peninsular Malaysia, the elephant population continues to decline, threatened by habitat degradation. Distinguishing their potential habitats is important in mitigating the impacts of forest fragmentation. This study aimed to predict the potential habitats of elephants in the fragmented forest landscape of Ulu Jelai Forest Reserve and to identify its characteristics. Habitat suitability was evaluated using several environmental variables––elevation, slope, land use, lithology, distance to river and distance from urban areas. The results showed that the distribution of suitable habitats mainly occurred in the eastern region of the study area. Elephant habitat prediction, using maximum entropy modelling, indicated that their most suitable habitat was located in a forested area that was greatly influenced by altitude and river proximity, while urban and agricultural areas cause forest fragmentation and restricted suitability. The prediction of wildlife habitats using maximum entropy modelling is a favourable and effective tool for improving the management and conservation planning of wildlife in the region.
... The continued expansion of agricultural land and infrastructure across the country appears to significantly disturb elephant habitat, resulting in increasing HEC incidents in the vicinity of LULCC (Chen et HEC generally. We found that HEC and LULCC mostly occurred within a 1 km radius of a recent LULCC event (Table 1, Table 2) (Yamamoto-Ebina et al., 2016). ...
Full-text available
Human–elephant conflict (HEC) is a key environmental issue in number of Asian countries, including Sri Lanka. Incidents of HEC have significantly increased in Sri Lanka between 1991 and 2018, with 1734 human deaths reported in this period (281% increase), 4837 elephant deaths (1172% increase), 1053 human injuries (140% increase) and more than 23,000 property damage reports (1406% increase). In this study we present a Sri Lanka wide analysis to explore the role of land use and land cover change (LULCC) in relation to HEC, using official government data and a land cover change dataset (1993–2018) recently developed by the authors using satellite imagery from the Landsat archive. We investigated rates of HEC over time and compared these to rates of LULCC over the same period. We also present spatial analytics of HEC and LULCC, as well as determining hotspots of HEC and LULCC using a kernel density estimator. Annual HEC incidents were found to broadly increase in line with land use change events (r = 0.43, p < 0.05). Human deaths, elephant deaths, human injuries and property damage hotspots show distinct spatial patterns: human deaths and injuries being more concentrated in the North West, Polonnaruwa and Ampara, wildlife regions; while elephant deaths are spread throughout the HEC region and property damage is high in the Central, Polonnaruwa Anuradhapura, North West, and Southern wildlife regions. We found a strong negative correlation between HEC location and distance to LULCC events. In total, 98% HEC occurred within 1 km of an area that experienced recent LULCC Since 2017, the primary HEC hotspots have shifted to the south and east of the country in concert with LULCC. These countrywide perspectives could help inform HEC mitigation strategies in Sri Lanka and other countries facing similar human-wildlife challenges.
... Forest patch adjacent agricultural land or crop field is the amazing source of food for elephant. Elephant seeks to eat secondary vegetation, i.e. crop products such as paddy than natural foods (Yamamoto-Ebina et al, 2016). So the amount of agricultural land is another important factor for choosing suitable habitat by elephant. ...
Full-text available
Diversified landscape functions have deep connection with habitat preference of elephant in several elephant habitats including Mayurjharna Elephant Reserve (MER). The main objective of the present study was to find out the elephant habitat suitability zone or areas considering geo-spatial components of landscape. For that reason, the study taken landuse factors, i.e. forest cover, forest core, forest fragmentation, forest edge, built-up area, agri-cultural land and road pattern, to fulfil the objective of the study. The selected landuse factors were separated from a classified landuse landcover (LULC) map of 2018 and inter-sected in a grid framework. From these grids, information factor density maps were pre-pared in ArcGIS 10.3 version software. Each factor has specific and different significance on habitat preference. After that, factors of significant level or percentage-wise ranks were calculated using analytical hierarchy process (AHP) on the basis of their ecological func-tion. Chosen factors were categorized and placed according to their significant percentage in Weighted Overlay tool in ArcGIS 10.3 software for getting the final outcome. After the analysis, it was found that 87.18 areas out of total area in MER are very suitable for elephant habitat. Reasonably suitable areas are 306.74 and areas, which are very less suitable for elephants in this reserve. 147.66 areas in MER are not suitable for elephant where basic habitat requirements are very insufficient. The well suitable areas are mostly covered with the high-density forest and large forest core areas in this reserve.
... little contribution of fruits(Yamamoto-Ebina et al., 2016). Indeed, elephants in BTFC seem to retain a good body condition throughout the year and over the years, not showing seasonal changes in body condition (Campos-Arceiz, personal observation).Our observations on the temporal patterns of seed dispersal by Asian elephants in Malaysia are congruent with the stance ofFredriksson et al. (2006) on the importance of long-term frugivory studies to account for the complex cycles and interannual masting episodes of trees in the forests of Sundaland. ...
Full-text available
Asian elephants (Elephas maximus) have inhabited almost all forests in tropical Asia until recently, yet little is known about their role in ecological processes, particularly in the Sundaic forests of Southeast Asia. These forests are peculiar in their phenology, with supra-annual and highly irregular episodes of mast fruiting. Here we present a long-term (six-year) monitoring of the seeds dispersed by elephants in dipterocarp forests of northern Peninsular Malaysia. We conducted monthly dung surveys at two mineral licks (11.3 km apart) frequently visited by elephants. Additionally, we recorded haphazard observations of seeds and seedlings in elephant dung at other locations. We recorded a minimum of 48 morphospecies from at least 25 plant families dispersed by elephants. Elephant seed dispersal was very heterogenous in space, with only 30.3 % of the morphospecies dispersed at both sites (Jaccard dissimilarity index = 0.48). Temporally, elephants dispersed seeds in sporadic pulses of abundance and diversity, without any apparent seasonality (seeds appeared in 19.1 % of 1,284 dung piles and 57.1 % of the 63 months in which we found dung) and with long periods without any seed being dispersed. Nearly half (48 %) of the plants dispersed by elephants belong to a megafaunal dispersal syndrome. Our long-term approach allowed us to unravel an important aspect of Asian elephants’ role and effectiveness in the seed dispersal cycle. Sundaland’s forests are undergoing a rapid loss of their not-long-ago common megaherbivores (rhinos and elephants), with profound and long-term consequences for ecosystem functioning.
... A 0.5-1 mm piece from one randomly chosen bird dropping from each plot from each collection date was placed on a microscope slide, dissolved in a few drops of water, and was examined under a compound microscope at 25-100× for the presence of leaf, seed, and arthropod fragments. To understand bird diets, I compared unique anatomical features (i.e., leaf and fruit surface trichomes, stomatal characteristics, seed surface characteristics, and leaf margin spines) of plant fragments in the bird feces with those of cover crops and weeds from the field; this approach is often used to study animal diets (Baumgartner and Martin, 1939;Martin, 1955;Stewart, 1967;Vaughan, 1967;Jennings and Barkham, 1975;Stevens et al., 1987;Yamamoto-Ebina et al., 2016;Takatsuki and Morinaga, 2020). Signs of folivory on cover crops and weeds were noted when droppings were collected. ...
Full-text available
Agriculture in many regions of the world has reduced bird habitat and abundance, and altered avian community structure. A study was conducted on an organic research farm over two winters (Oct to Mar) in an intensive agricultural region of Salinas Valley, CA to determine how cover crop variety and planting density influenced birds. Cover crops were rye (Secale cereale), a mixture of rye and legumes (Vicia spp., Pisum sativum), and a mustard mixture (Brassica juncea, Sinapis alba). White-crowned Sparrows (Zonotrichia leucophyrs), Savannah Sparrows (Passerculus sandwichensis) and Song Sparrows (Melospiza melodia) were observed both years in the study field. Bird droppings in cover crops were quantified and dissected to determine dietary preferences, and sparrow movement when flushed was determined. Dropping number and weight per m2 were at least 10 times greater in mustard than in rye and in the legume-rye mixture. Droppings were dominated by leaf tissue in mustard vs. arthropod tissue in rye and legume-rye. Within cover crop variety, plant density did not have a clear or consistent effect on sparrows. Sparrows flushed from cover crops usually settled in mustard. The White-crowned Sparrow fed on mustard leaves and apparently on weed foliage under mustard. The arthropod-dominated droppings in rye and legume-rye cover crops were consistent with the food preferences of Song and Savannah Sparrows. The White-crowned Sparrow's clear preference for mustard cover crops is likely due in part to their high dietary needs for sulfur-rich amino acids during the prenuptual molt. This paper provides novel information to help farmers and others understand the cover crop preferences of sparrows, and ways that farmers might use mustard cover crops as trap crops to reduce White-crowned Sparrow feeding damage on winter and spring vegetable crops. It also provides evidence of ecosystems services that these sparrows provide by feeding on weed tissue in winter cover crops.
... Asian Elephants (Elephas maximus) are edge specialists (Yamamoto-Ebina et al., 2016), and they readily use human-disturbed environments near to forest, including agriculture areas and newly planted rubber and oil palm plantations. Being a mega-herbivore, an adult elephant weighs 1,000-5,000 kg, and needs to eat approximately 10% of its body weight every day (Fernando, 2015). ...
Full-text available
The research provides a scientific synthesis of information on HEC encountered by SDPB Malaysia operations for the duration of 2011-2018. This research suggested that elephant depredation mostly occurs when the oil palm trees are below five years old, and the most damage takes place when the tree is one year old. The spatial distribution of highest HEC intensity and damage frequency occurred mostly in the area of entry point at estate borders and some were reduced with the application of mitigation. The temporal pattern of HEC in SDPB suggested that some estates showed a clear reduction in HEC when comparing HEC incidents before and after the year of electric fencing is in place but not for all. This concurred that an electric fence is useful when applied in the right conditions, but it may not be a solution for all HEC. Further research and observation are needed at respective estates of SDPB. The HEC pattern is not correlated with monthly rainfall. The total economic loss for the 8 years duration is RM24 million.
... The comparison between ethnoveterinary medicine and Asian elephant diet also utilized two datasets, one of elephant use reports provided by Karen knowledge holders, and the other of literature on Asian elephant diet in Thailand (Kroutnoi et al., 2017;Kitamura et al., 2007) and throughout their range (Suba et al., 2018;Alahakoon et al., 2017;Koirala et al., 2016;Yamamoto-Ebina et al., 2016;English et al. 2014English et al. , 2015Roy and Chowdhury, 2014;Mohapatra et al., 2013;Sitompul et al., 2013;Baskaran, 2010;Pradhan et al., 2008;Campos-Arceiz et al., 2008;Varma et al., 2008;Samansiri and Weerakoon, 2007;Chen et al., 2006;Himmelsbach et al., 2006;Hettiarchchi et al., 2005;Steinheim et al., 2005). This literature was further supplemented by unpublished data provided by Global Volunteers International and Mahouts Elephant Foundation, the organizations running compassionate conservation programs in two of the villages. ...
Ethnopharmacological relevance Ethnoveterinary medicine is often assumed to be a subset of human medicinal knowledge. Here we investigate the possibility that some ethnoveterinary medicine rather originates from observations of animal self-medication. We document and analyze the ethnoveterinary medicine used by Karen mahouts for elephant care and attempt to determine whether this knowledge originated from humans or elephants. Materials and methods Elephant camp owners and mahouts in four communities in northern Thailand were interviewed about their knowledge and use of plants for ethnoveterinary elephant care. For each ethnoveterinary plant, data were collected on Karen human medicinal uses and whether elephants independently consume them. Based on overlaps between ethnoveterinary use, human medicinal use and elephant dietary use, plants were classified into three categories: those that originated from Karen human medicine, those that originated from Asian elephant self-medication, and those which were present in both human and elephant knowledge traditions. Results The use of 34 plants (32 identified at least to genus) and two additional non-plant remedies (salt and human urine) were reported to be used in ethnoveterinary elephant medicine. A total of 44 treatments in 11 use categories were recorded: tonic, wounds, compress, eye problems, indigestion, broken bones, galactagogue, snakebite, fatigue, skin and musth regulation. Of the ethnoveterinary plants, 55% had the same use in human medicine, 43% had different uses and 2% had no use. Elephants consume 84% of the ethnoveterinary plants as part of their natural diet. Discussion Analysis indicates that 32% of plant uses likely originated from Karen human medicine, 60% of plant uses likely existed independently in both human and elephant knowledge systems, and 8% of plant uses likely originated from elephant self-medicating behavior. The tonic use category shows the strongest evidence of influence from observations of elephant self-medication. The use of tonic medicines appears to be increasing as a way to mitigate the unnaturally limited diet of elephants in tourist camps. Conclusion Ethnoveterinary medicine for elephant care is influenced by both human medicinal knowledge and elephant knowledge of plants for self-medication. The ethnoveterinary knowledge domain appears to be the result of an interactive process linked to convergent evolution or co-evolution between humans and Asian elephants.
... Finally, the food source segregation can be observed through the difference in diet selection by the full utilisers in which 92% of diet selection by smooth-coated otter is dominated by fish [25] while wild boar diet is dominated by 90% of plant material include rhizome and roots [26] and Asian elephant is a herbivore that feeds on monocot leaves, hard dicot material and grasses [27]. The clear food source segregations by all three species make it possible for them to live together in Puah Reservoir. ...
Impoundment is the main phase of dam development that significantly destroys mammals and its habitats. Elephant rescue activity in Kenyir Reservoir post-impoundment suggests the ability of mammals to swim to the remaining terrestrial habitats in search of shelters. Due to the lack of data on wildlife survival in the reservoirs of Malaysia, this study aimed to establish a list of mammal species surviving in Puah Reservoir, Terengganu after two years of inundation, to assess the developed adaptive behaviour and to describe their movement pattern in the reservoir. Four camera traps were placed on each of the seven selected land-bridge islands for 12 months. All captured images were analysed through camtrapR package in R-3.5.0 software. A total of 11 mammal species were recorded surviving in Puah Reservoir. From these species, Asian palm civet and long-tailed macaque have adapted to permanently inhabit an island while smooth-coated otter, Asian elephant and wild boar are well adapted to fully utilise the reservoir. The primates or solitary mammals such as Sumatran serow, barking deer, marbled cat, Malay tapir, dusky leaf monkey and pale-thighed langur have adapted to be the occasional utilisers of the reservoir. The second group has developed a movement strategy where their presence on each island are not overlapping with each other. This study concludes that mammals can survive in a reservoir by adapting to habitat changes and develop a survival strategy. This study will be among the earlier study in Malaysia which documents the survival data of mammals in reservoirs.
Full-text available
In the Malay Peninsula, people have lived alongside Asian elephants (Elephas maximus) for around 55,000 years but our expansion now endangers the species. With the aim of gaining knowledge on how to we can live together in future, I reviewed the ecology, history, and management of human-elephant relations in the Peninsula. I found that indigenous people (Orang Asli) occupied many of the same landscapes as elephants and, despite a degree of ecological overlap, managed to enjoy a convivial coexistence by following the pathways elephants created through the rainforest, and by subsisting off wild yams. Around 6500 years ago, a swidden-farming culture arrived and crop-raiding elephants were killed and occasionally eaten. Around 2500 years ago, new settlers arrived and elephants came to be sought for ivory, to be captured, tamed, and even exported. Aspects of the traditional forager and swiddener cultures remain in Belum-Temengor, a priority elephant conservation site in the north of the Peninsula. Here, I surveyed 37 villages to examine beliefs, attitudes, and behaviour towards elephants. I found that tough elephants were the main source of human-wildlife conflict, most respondents considered the animals to be worthy of respect. Thre were some indications that younger respondents tended to have less tolerant attitudes. To get a clearer idea on how to manage elephants in this landscape, I mapped the villages and monitored the movement of four elephants using satellite collars. I found that governement-sponsored rubber plantations, exposed villagers to elephant raids despite the construction of electric fences. Based on these findings I propose a five-phase strategic intervention approach to elephant conservation: (i) land-use planning; (ii) barriers to protect people (including electric fences); (iii) compensation for losses; (iv) education and engagements; and (v) removal (killing or capture).
Full-text available
Tropical forests developed in isolation in five main areas during the Tertiary: Asia, New Guinea, Africa, Madagascar and the Neotropics. Asian forests share taxa with New Guinea and Africa, but there are also unique features. Most emphasis has been on the dominance of lowland moist forests by the Dipterocarpaceae, but the importance of the Fagaceae in lowland forests is also unique. Among the vertebrates, gibbons, tree shrews, forest rhinoceroses and lowland bears are unique to Asia, as is the diversity of squirrels, babblers and gliding vertebrates. Honeybees are shared with Africa, but only Asian forests support several coexisting species.
Full-text available
The plant vigour hypothesis proposes that herbivores should favour feeding on more vigorously growing plants or plant modules. Similarly, we would expect herbivores to favour plants that regrow vigorously after herbivory. Larger animals, like elephants, may also select plant species relative to their availability and prefer species with larger growth forms in order to meet their intake requirements. The food preferences of the Bornean elephant (Elephas maximus borneensis) in the Lower Kinabatangan Wildlife Sanctuary, Sabah, Malaysia, were investigated along 12 transects in areas where elephants were recently sighted feeding. One hundred and eighty-two plants were eaten and 185 plants were measured for species availability along transects. Species vigour was determined by the monthly regrowth in new shoot length after elephant feeding and the number of new shoots produced on each plant. Measurements were carried out on each plant for 9 months, or until the new shoot was eaten. Plant sizes were determined from their basal diameter. The Bornean elephant did not prefer more vigorous species or species with larger growth forms. New shoots did not grow longer on preferred than avoided species. Additionally, unlike other elephants that live in a forest environment, the Bornean elephant preferred species from the Poaceae (specifically Phragmites karka and Dinochloa scabrida) over other plant types including gingers, palms, lianas and woody trees.
Full-text available
Recursion by herbivores is the repeated use of the same site or plants. Recursion by wild animals is rarely investi-gated but may be ubiquitous. Optimal foraging theory predicts site recursion as a function of the quality of the site, extent of its last use, and time since its last use because these influence site resource status and recovery. We used GPS collars, behaviour and site sampling to investigate recursion to foraging sites for two elephant Elephas maximus borneensis herds in the Lower Kinaba-tangan Wildlife Sanctuary, Borneo, over a 12 month period. Recursion occurred to 48 out of 87 foraging sites and was most common within 48 hours or between 151–250 days, indicating two different types of recursion. Recursion was more likely to oc-cur if the site had previously been occupied for longer. Moreover, the time spent at a site at recursion was the same as the time spent at the site on the first occasion. The number of days that had passed between the first visit and recursion was also positively correlated with how much time was spent at the site at recursion. Habitat type also influenced the intensity of site-use, with more time spent at recursion within riverine/open grass areas along forest margins compared to other habitat types. Recursion is a common behaviour used by the elephants and its pattern suggests it may be a foraging strategy for revisiting areas of greater value. The qualities of recursion sites might usefully be incorporated into landscape management strategies for elephant conservation in the area [Current Zoology 60 (4): 551–559, 2014].
Full-text available
I recently read a student's paper saying 'as more forests are cleared and fragmented, elephants have no choice but to encroach into plantations in their search for food, water, and mates' and the myth of le bon sauvage immediately came to mind. The myth of the noble savage, popular in the 18 th and 19 th centuries, has to some extent impregnated conservation biology's philosophy in recent decades. This myth is an idealization of people living in traditional societies, attributing them a noble spirit and behavior. Accordingly, conservation biologists have often assumed that human traditional societies always live sustainably and in harmony with their environment. Now we know, however, that conservation by indigenous people is uncommon (Raymond 2007) and that, for example, stone-age societies wiped out over 90% and 70% of large mammal species when they reached Australia and the Americas (Barnosky et al. 2004). The myth of the noble beast is very similar and also widely spread in conservation biology. Large herbivores and carnivores often come into conflict with people not because they have no other option but as part of their optimal foraging strategy (Stephens 1986). For example, food for elephants can be very limited in a pristine tropical rainforest yet abundant in nearby plantations and human-dominated landscapes (crops and early succession plants are excellent elephant forage). In this situation, elephants may choose to come out of the forest we consider their legitimate habitat and 'encroach' into human areas, resulting in the well-known human-elephant conflict. The key driver of conflict in this scenario is not the amount of forest available for the elephant but the amount of interface between forest and crops.
This paper describes the adaptation of the point analysis method, a standard vegetational sampling technique, to rumen analyses. The sampling device consisted of a frame containing five hatpins placed at a 45° angle through a wooden bar. The device was tested by two investigators on several combinations of artificially constructed populations of plant fragments and on actual rumen contents of white-tailed deer (Odocoileus virginianus). There were no significant differences between the ability of two investigators to estimate the volumetric composition of the samples, nor was there a difference in the ability of the individual investigator to repeat his estimates. The point analyzer gave reliable results provided (1) the rumen sample was adequately mixed and (2) there were no large items with unusual surface texture in the mixture.
The Asian elephant has had a unique cultural association with people. Unfortunately, elephants and people have also been in conflict, resulting in the decline in elephants throughout their former range in Southern Asia. This book provides an ecological analysis of elephant human interaction and its implications for the conservation of elephants. The foraging habits of elephants and their impact on vegetation are considered, along with the interactions that occur between elephants and humans. The ecological data provide the basis for recommendations on elephant conservation and management, keeping in view the socioeconomic imperatives of the Asian region.This first comprehensive account of Asian elephant ecology will be of particular interest to conservation biologists and mammalogists.
1. Most methods of studying the food preferences of grazing herbivores are difficult or impossible to apply under conditions such as those encountered on the East African plains, which have an outstanding large-mammal fauna and a rich herbaceous flora. There are therefore few data regarding the preferences of these mammals. 2. A method involving the identification of plant epidermis in faeces avoids some of the limitations of other techniques. It has been used elsewhere both with large mammals and with other groups, but usually where the numbers of animal and plant species are fewer; the data obtained have usually been qualitative only. 3. The present study examines the qualitative and quantitative potential of the method under East African conditions. It has been limited to grasses, and to their leaf epidermis. It has involved feeding experiments with seven animal species (six ruminants and one non-ruminant) and, for quantitative purposes, eight species of grasses, the number of the latter being limited by practical considerations. 4. The results indicate that perennial grasses forming more than 5~%, by fresh weight, of a constant diet can be identified in the faeces. No information was obtained regarding species which form a smaller, regular part of the diet, or on ephemerals regularly forming more than 5%. 5. With a changing diet perennials temporarily forming a major part will be identified within a period governed by the times of throughput and elimination (see 6 below). Grasses eaten in occasional small quantities may not be identified; the evidence was insufficient to indicate whether ephemeral species are less likely to be recorded. 6. In both the ruminants and the non-ruminant the period between the first ingestion of a grass and its first appearance in the faeces (time of throughput) was 20-30 h; the period between the last ingestion and last appearance in the faeces (time of elimination) was about 3 days in the non-ruminant and 5-6 days in the ruminants. 7. Since the major constituents of the diet can be identified in the faeces, quantitative data on a frequency basis, indicating the relative importance of different grasses in the diet, can be obtained. 8. The proportions of epidermis from different grasses can be estimated in the faeces by measuring the area of fragments or by using point quadrats; counts of fragments are invalid for this purpose since different grasses break into fragments differing significantly in size. The proportions so measured do not necessarily represent those ingested, however, since different grasses may be digested to different extents. The fact that epidermis from both leaf surfaces survives digestion in some species and from only one in others does not account for all these differences. There are also significant differences between animals in the extent to which particular grasses are digested. 9. Although for intensive studies involving a few plant and animal species it might be possible to establish correction factors to allow for variations in digestibility, this and the subsequent analyses would be extremely time-consuming; it must therefore be accepted that most studies must be limited to obtaining quantitative data on a frequency basis.