RESEARCH ARTICLE
Obligate Exudativory Characterizes the Diet of the Pygmy Slow Loris
Nycticebus pygmaeus
CARLY STARR
1
AND K.A.I. NEKARIS
2
*
1
School of Agriculture and Food Science, University of Queensland, Gatton, Queensland, Australia
2
Oxford Brookes University, Nocturnal Primate Research Group, School of Social Sciences and Law, Oxford,
United Kingdom
Few primate species are known to excavate plant sources to procure exudates and other foods via active
gouging. It is now apparent that slow lorises belong to this rare guild of obligate exudativorous primates.
We investigate the diet of the pygmy loris (Nycticebus pygmaeus) in a mixed deciduous forest in the
Seima Protection Forest, Eastern Cambodia, and attempted to determine the importance of this
resource in their diet. Feeding behaviors of six females and seven males were observed using radio‐
tracking to facilitate follows, and nine fecal samples were collected in February–May and January–
March in 2008 and 2009 respectively. We observed 168 feeding bouts, during which the animals ate
exudates (76); fruits (33); arthropods (27); flower parts (21); fungi (3); parts of bamboo culms (7); and
reptiles (1). We filmed 19 bouts of exudativory, and observed animals consuming exudates in an
orthograde posture, or standing quadrupedally over the exudate source. Pygmy lorises also gouged
bamboo to collect lichen and fungi, or broke open dead culms to access invertebrates. Feeding occurred on
terminal tree branches (24), tree trunks (21), bamboo (13), the middle of branches (7), and the
undergrowth (1). The fecal samples contained plant parts, small‐sized arthropods (primarily Coleoptera
and Lepidoptera), reptile scales, animal bones, and animal hairs. Pygmy slow lorises are
morphologically specialized for processing and digesting exudates, displaying small body sizes,
specialized dentitions, elongated, and narrow tongues, large caecums, short duodenums, expanded volar
pads, and modified hindlimbs. These features, combined with the prevalence of exudates in their diet
across seasons, and ill health when exudates are missing from their diet in captivity, points to this
species being an obligate exudativore. Am. J. Primatol. 9999:1–8, 2013. © 2013 Wiley Periodicals, Inc.
Key words: radio‐tracking; slow loris; gummivory; fallback foods
INTRODUCTION
Exudates, including gums, and saps, provide a
carbohydrate‐rich energy source for those primates
able to access and digest them [Heymann &
Smith, 1999]. Although saps, the circulating fluids
in plants, can be reached without excavation, gum
consumption often requires active gouging of the
surface of a tree [Bearder & Martin, 1980b; Vinyard
et al., 2003]. At least 69 primate species are now
known to use exudates, but until recently only five
genera (Callithrix, Mico, Cebuella, Phaner, and
Euoticus) were known to gouge the surface of trees
to obtain exudates [Nash, 1986; Nash & Burrows,
2010; Smith, 2010; Fig. 1]. These taxa are defined as
obligate exudativores and exudates comprise 40% or
more of their feeding time (hereafter referred to as
diet) [Nash & Burrows, 2010]. Morphological special-
izations characterizing these primates include modi-
fied anterior dentition, small body size, a large
caecum and/or colon, a long tongue, and keeled nails
[Hladik, 1979]. Nekaris et al. [2010a] suggest that
Asian lorises (Nycticebus, Loris) should be added to
this list of exudativores given that they possess all
but the last of these adaptations.
Nekaris et al. [2010a] reviewed field studies
of lorises and pottos, and showed that nine species
are now known to include exudates in their diet.
Whereas, pottos are only known to scrape gum, seven
species of Loris and Nycticebus actively gouge for
exudates. Loris clearly seems to use exudates as
Contract grant sponsor: Leverhulme Trust; contract grant
number: RPG‐084.
Conflicts of interest: None.
Ã
Correspondence to: K.A.I. Nekaris, Oxford Brookes University,
Nocturnal Primate Research Group, School of Social Sciences
and Law, Oxford OX3 0BP, United Kingdom.
E‐mail: anekaris@brookes.ac.uk
Received 26 August 2012; revised 29 April 2013; revision
accepted 4 May 2013
DOI: 10.1002/ajp.22171
Published online XX Month Year in Wiley Online Library
(wileyonlinelibrary.com).
American Journal of Primatology 9999:1–8 (2013)
© 2013 Wiley Periodicals, Inc.