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Foraging ecology of woodpeckers in lowland Malaysian rain forests

Alison R. Styring and Mohamed Zakaria bin Hussin

Journal of Tropical Ecology / Volume 20 / Issue 05 / September 2004, pp 487 - 494

DOI: 10.1017/S0266467404001579, Published online: 09 August 2004

Link to this article: http://journals.cambridge.org/abstract_S0266467404001579

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Alison R. Styring and Mohamed Zakaria bin Hussin (2004). Foraging ecology of woodpeckers in lowland Malaysian rain
forests. Journal of Tropical Ecology, 20, pp 487-494 doi:10.1017/S0266467404001579

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Journal of Tropical Ecology (2004) 20:487–494. Copyright © 2004 Cambridge University Press
DOI: 10.1017/S0266467404001579 Printed in the United Kingdom

Foraging ecology of woodpeckers in lowland Malaysian rain forests

Alison R. Styring∗1 and Mohamed Zakaria bin Hussin†

∗ Department of Biological Sciences and Museum of Natural Science, Louisiana State University, Baton Rouge, LA 70803, USA
† Department of Forest Management, Faculty of Forestry, Universiti Putra Malaysia, Serdang, Selangor, West Malaysia
(Accepted 29 September 2003)

Abstract: We investigated the foraging ecology of 13 species of woodpecker in logged and unlogged lowland rain forest
at two forest reserves in West Malaysia (Pasoh Forest Reserve and Sungai Lalang Forest Reserve). The parameters
perch diameter and microhabitat/substrate type explained more variation in the data than other parameters, and
effectively divided the guild into two groups: (1) ‘conventional’ – species that excavated frequently, used relatively
large perches, and foraged on snags and patches of dead wood, and (2) ‘novel’ – species that used smaller perches and
microhabitats that are available in tropical forests on a year-round basis (e.g. external, arboreal ant/termite nests and
bamboo). These novel resources may explain, in part, the maintenance of high woodpecker diversity in tropical rain
forests.

Key Words: diversity, forage, Malaysia, woodpecker

INTRODUCTION on foraging were lacking, his observations suggested

that woodpeckers partitioned resources behaviourally
Lowland rain forests of South-East Asia are unique in using a variety of foraging methods, foraging heights
having the highest single-point diversity for woodpeckers and substrates. We approached the hypothesis of
(Picidae) in the world (Short 1978, Styring & Ickes 2001, niche differentiation by making detailed, quantitative
Wells 1999). With up to 16 species in a single patch of observations on the foraging ecology of 13 species at
habitat, woodpeckers comprise the majority of the bark- Pasoh Forest Reserve (PFR) and Sungai Lalang Forest
foraging guild in this region and a larger percentage Reserve (SLFR).
of the total avifauna than do woodpeckers in other
tropical forests (6–8% of total species vs. 3–5% and 2–3%
for tropical rain forests in the Neotropics and Africa res-
pectively; see Styring & Ickes 2001). A variety of genera STUDY AREAS
and a broad range of body sizes are represented in this
South-East Asian guild (Table 1). We conducted research at two forest reserves in West
Woodpeckers are highly adapted for excavating woody Malaysia: Pasoh Forest Reserve (PFR) in the state of
substrates to obtain food and nesting space, and they Negeri Sembilan (2◦ 59 N, 102◦ 17 E; May–July 1998),
form a discrete ecological guild in forests where they and Sungai Lalang Forest Reserve (SLFR) in the state of
occur. Understanding the foraging preferences of so Selangor (3◦ 30 N, 101◦ 53 E; July–Sept 1999, Feb–April
many ecologically similar species will provide insight 2000 and June–Aug 2000) (Figure 1a–c).
into resource use and potential resource partitioning, PFR consists of a core area of c. 600 ha of virgin lowland
and may help us understand how this guild responds dipterocarp forest, surrounded by c. 1400 ha of forest
to disturbance. Short (1978) qualitatively described that was selectively logged in 1955–1956 (Kochummen
the foraging and nesting behaviour of 13 woodpecker et al. 1990, Wong 1985, 1986) (Figure 1b). Additional
species at Pasoh Forest Reserve and several other sites forest management techniques, such as poison-girdling
in Peninsular Malaysia. Although quantitative data of emergent non-commercial timber species and climber
removal, were performed in the selectively logged portion
of Pasoh between 1955 and 1960 (Kochummen et al.
1 Corresponding author. Email: [email protected] 1990). The forest is classified as lowland rain
488 ALISON R. STYRING AND MOHAMED ZAKARIA BIN HUSSIN

Table 1. List of the 13 woodpecker species commonly found in lowland rain forests in Malaysia. Mean masses taken from Short (1978). Under
‘Location observed’ PFR = Pasoh Forest Reserve, SLFR = Sungai Lalang Forest Reserve, VJR = Virgin Jungle Reserve, 5-y = 5-y-old logged forest,
10-y = 10-y-old logged forest, ul = unlogged forest. Sites listed in parentheses indicate locations where the birds were seen, but no foraging
observations were made.
Mean body Number of foraging
Species Species code mass(g) observations Location observed
Sasia abnormis Temminck, 1825 SA 9 40 SLFR all sites (PFR)
Hemicircus concretus Temminck, 1821 HC 32 38 PFR and SLFR all sites
Meiglyptes tristis Horsfield, 1821 MT 46 126 PFR and SLFR all sites
Meiglyptes tukki Lesson, 1839 MTU 53 31 PFR and SLFR all sites
Celeus brachyurus (Vieillot, 1818) CB 66 22 PFR and SLFR all sites
Blythipicus rubiginosus (Swainson, 1837) BR 84 17 SLFR all sites (PER)
Picus miniaceus Pennant, 1769 PMI 83 4 PFR (SLFR 5-y, ul)
Picus mentalis Temminck, 1825 PM 89 58 PFR and SLFR all sites
Picus puniceus Horsfield, 1821 PP 98 38 PFR and SLFR all sites
Dinopium rafflesii (Vigors & Horsfield, 1830) DR 96 10 PFR and SLFR 5-y, 10-y (VJR)
Reinwardtipicus validus Temminck, 1825 RV 155 54 PFR and SLFR all sites
Dryocopus javensis (Horsfield, 1821) DJ 225 36 PFR and SLFR all sites
Mulleripicus pulverulentus (Temminck, 1826) MP 430 29 SLFR 5-y (PFR and SLFR ul)

(a) West Malaysia

Thailand 0 50 100 150

Kedah km

Pinang Terengganu
Kelantan
Perak North
Straits
South
of
Pahang China
Malacca
Sungai Lalang Sea
Selangor
Forest Reserve
Pasoh Forest Negeri Sembilan
Reserve
Melaka
Johor

Singapore

(b) Pasoh Forest (c) Sungai Lalang

Reserve Forest Reserve
unlogged

5-y-old logged

45-y-old logged
unlogged
0 500 1000 0 1 2
10-y-old logged
m km

Figure 1. Map of (a) West Malaysia with study sites, (b) Pasoh Forest Reserve and (c) Sungai Lalang Forest Reserve.
Woodpecker ecology in Malaysian rain forest 489

Table 2. Description of attack variables used in this study (taken from Remsen & Robinson 1990).
Manoeuvre Purpose
Glean To pick food from a nearby substrate. Can be reached without full extension of legs or
neck without the involvement of acrobatic movements.
Hang To hang, head down in order to reach food not obtainable by any other perched position.
Probe To insert bill into cracks, holes or soft substrates to capture hidden food.
Flake To brush aside loose substrate with a sideways, sweeping motion.
Peck To drive the bill against a substrate to remove some of the exterior of the substrate.
Chisel Like ‘peck’ but the bill is aimed obliquely rather than perpendicularly at the substrate.
Hammer To deliver a series of pecks. Used to excavate deep holes to reach bark or wood-dwelling
insects or sap.

forest (elevation: 75–150 m, average annual rainfall: at PFR were made in primary forest, whereas observations
1900 mm), and the emergent vegetation layer is were collected in both primary and logged forest at SLFR.
dominated by various species in the Dipterocarpaceae, Birds were followed for as long as possible, but only the
primarily in the genera Shorea and Dipterocarpus initial observations were used for statistical analysis to
(Kochummen et al. 1990). avoid problems with non-independent data.
SLFR (Figure 1c) is a large forest reserve (17 722 ha), Multivariate analyses were performed on foraging
consisting of both recently logged (< 5 y ago), mature data to reduce the information into a few, readily
forest logged as long ago as the 1940s, and a few interpretable variables or vectors. Both continuous and
patches of primary forest. The forest is classified as categorical data were collected, and these two types
lowland/hill rain forest (elevation: 100–800 m, average of data were best analysed using different multivariate
annual rainfall: 3115 mm), and, like PFR, the emergent techniques. To make the results as readily understandable
vegetation layer is dominated by species in the genera as possible, continuous variables were converted into
Shorea and Dipterocarpus (Laidlaw 2000). Data were categorical variables using size and height classes. To
collected at three sites within the reserve: Compartment establish meaningful categories, SYSTAT 8.0 (SPSS
18 (260 ha), which was logged from November 1993 Inc. 1998) was used to perform principal components
to September 1995, Compartment 33 (387 ha), which analysis (PCA) on continuous variables (listed 1–4 in
was logged from January 1990 to March 1991, and a the previous paragraph). PCA reduces data by forming
patch of unlogged forest (82 ha) located within a logged linear combinations of variables and is also useful in
compartment (compartment 24 logged from 1960 to classifying variables. Analyses of variance and pair-wise
1963). This unlogged patch of forest was designated as a comparisons of continuous variables were performed
protected area known as a Virgin Jungle Reserve (VJR). to determine natural ‘breaks’ in the data and thus
meaningful categories.
Once continuous variables were converted to
categorical, frequency data, correspondence analysis
METHODS (CA) was performed on the data set using SAS 8.00
(1999). CA is an ordination technique similar to PCA,
Throughout this study, ARS recorded the following data but it is effective at identifying structure in categorical,
using a micro-cassette recorder on foraging birds en- frequency data by detecting correspondence between
countered opportunistically: (1) estimated height above rows and columns of a data matrix. Correspondence
ground, (2) estimated distance from the canopy, (3) dia- analysis was found to explain more variation in foraging
meter at breast height (dbh) of the foraging substrate, data than other multivariate techniques (Miles 1990).
(4) perch diameter, (5) attack manoeuvre (‘hammer’, In order to determine the level at which certain
‘chisel’, ‘flake’, ‘glean’, ‘hang’, ‘peck’, ‘probe’; Table 2), parameters were most important in distinguishing
(6) horizontal position (inner = trunk, middle = large species, dissimilarity matrices were constructed for
branches, outer = branchlets/leaves) and (7) substrate individual foraging parameters (perch, substrate, method,
type (‘ant/termite’ – tunnels and nests found on the height, horizontal position). These indices were combined
external surfaces of tree trunks and branches, ‘bamboo’, to form an additive index of dissimilarity for each pair
‘log’ – a dead tree lying on the ground, ‘liana’ – a woody of species. Using this index, an additive tree was built.
vine, ‘dead patch’ – a section of dead wood on a live Values of dissimilarity were calculated at every branch
tree, ‘live wood’, ‘leaf’, ‘Macaranga’ – a genus of tree that for each parameter so that the parameter accounting
frequently has mutualistic associations with ants, and for the majority of dissimilarity at each branch could be
‘snag’ – a standing dead tree). All observations collected determined.
490 ALISON R. STYRING AND MOHAMED ZAKARIA BIN HUSSIN

–1.5
Hemicircus concretus
Upperstorey

upperstorey
–1.0 Mulleripicus pulverulentus

Meiglyptes tristis Celeus brachyurus

–0.5
Sasia abnormis
Dryocopus javensis
vertical placement

0.0
Factor 2 (30%)

Picus puniceus Midstorey

Picus mentalis
0.5

1.0 Blythipicus rubiginosus

Meiglyptes tukki
understorey

1.5
Understorey
2.0 Reinwardtipicus validus

–2.0 –1.5 –1.0 –0.5 0.0 0.5 1.0 1.5 2.0

Factor 1 (52%)
small substrate diameter large

Figure 2. Principal components analysis of continuous foraging variables (perch diameter, dbh of foraging substrate, height above ground and
distance from the canopy). Circles indicate groups of species that did not differ significantly (probability of 0.05 or less) for any continuous variable
when ANOVA was performed and pairwise comparisons made. The y-axis was inverted to portray foraging height from the bottom of the figure
to the top (so that understorey-foraging woodpeckers appear at the bottom of the figure and upperstorey-foraging woodpeckers appear at the top
rather than vice versa).

RESULTS categories: (1) understorey – 5 m or lower; (2) midstorey –

higher than 5 m and lower than 5 m from the canopy; and
Over the course of the study, a total of 503 independent (3) upperstorey – 5 m from the canopy or higher.
foraging observations were recorded. With the exception Although broad interpretations of the raw categorical
of Meiglyptes tristis, the number of foraging observations data are not easily made, a few results were apparent
for individual species was rather low. Because there were from the frequency data (Appendix 1). Variation was
only four independent observations of Picus miniaceus, this considerable among substrates and attack methods.
species was not included in any analyses. With only 10 However, all species used dead and live wood to some
independent observations, Dinopium rafflesii was included extent. The substrate variables ‘Macaranga’, and ‘leaf’,
in some of the multivariate analyses, but not in analyses and the method ‘hang’ were used by mostly one species
where probabilities were important. each. Meiglyptes tristis used Macaranga extensively, and it
Principal components analysis revealed two principal often foraged on leaves and stipules.
components that explained 82% of the variation in Correspondence analysis of foraging variables resulted
the data (Figure 2). Principal component one, which in three dimensions that explained 52% of the variation
explained over 50% of the variation in the data set, was in the data (Figure 3). Similar to the PCA, Dimension
weighted heavily on perch diameter. Principal component one was weighted heavily toward perch diameter (41%
two (explaining 30% of the variation in the data) was of the variation in Dimension 1) followed by microhabitat
a combination of the variables ‘height above ground’ (31% of the variation in Dimension 1). Dimension
and ‘distance from the canopy’ and can be thought of as two was weighted heavily on the parameter substrate
vertical placement. Based on this PCA (Figure 2), ‘perch (followed by height). Meiglyptes tristis which foraged
diameter’ was divided into five categories: XS – perch is primarily on leaves, and Blythipicus rubiginosus which
less than 5 cm diameter; S – perch is 5 cm or greater was restricted to dense understorey, clearly separated
and less than 10 cm; M – perch is 10 cm or greater from the other species. Also, the group containing the
and less than 20 cm; L – perch is 20 cm or greater and species Meiglyptes tukki, Celeus brachyurus, Sasia abnormis
less than 35 cm; and XL – perch is greater than 35 cm. and Hemicircus concretus (all of which used relatively
Likewise, ‘vertical placement’ was categorized into three small perches) was distinct from the group containing
Woodpecker ecology in Malaysian rain forest 491

–0.5
Meiglyptes tristis Dryocopus javensis

upperstorey
–0.3 Celeus brachyurus Mulleripicus pulverulentus

leaf Reinwardtipicus validus

–0.1 Hemicircus concretus
Dimension 2 (17%) Picus puniceus
microhabitat (39%)
height (31%)

0.1 Picus mentalis

Meiglyptes tukki
0.3

Sasia abnormis
0.5

0.7
understorey

Novel foragers Conventional foragers

log

0.9
Blythipicus rubiginosus

1 0.5 0 –0.5 –1
Dimension 1 (32%)
small perch diameter (41%) large
leaf microhabitat (31%) snag

Figure 3. Correspondence plot of foraging variables. Axes were inverted so that woodpeckers using small perches appear on the left side of the figure
and those using larger perches appear on the right (rather than vice versa), and so that understorey log-foraging woodpeckers appear at the bottom
and upperstorey leaf-foraging woodpeckers appear at the top (rather than vice versa).

Meiglyptes tukki
(substrate) 0.38 (height)
Sasia abnormis
1.51
Hemicircus concretus
Novel 1.79 (method) Meiglyptes tristis
foragers 14.63 0.36 (perch/substrate)
Celeus brachyurus
(perch)

0.23 (method)
Picus puniceus
Conventional Picus mentalis
foragers 2.28 (substrate)
Reinwardtipicus validus
0.27
2.19 0.82 (height) Dryocopus javensis
(substrate) (method)
Mulleripicus pulverulentus
Blythipicus rubiginosus
Figure 4. Additive ecological tree of foraging variables. Numbers beside branches indicate the overall dissimilarity between groups (see Appendix 2),
and the foraging variables that accounted for most of the dissimilarity at each branch are indicated in parentheses.

Reinwardtipicus validus, Dryocopus javensis, Mulleripicus birds that foraged on dead-wood patches and snags
pulverulentus, Picus mentalis and Picus puniceus (which (large-perch birds) and birds that used resources such
used larger perches). as ant/termite nests and bamboo (small-perch birds).
The additive ecological tree built from the dissimilarity At the next set of branches, substrate preference was
matrix revealed that perch diameter was important at important in subdividing each group into more specific
the base of the tree and divided birds that used small groups based on substrate preferences such as those that
perches from those that used large ones (Figure 4 and used woody, live substrates such as bamboo/small live
Appendix 2). Consequently, this division also represents branches and Macaranga trees (Meiglyptes tristis, Celeus
492 ALISON R. STYRING AND MOHAMED ZAKARIA BIN HUSSIN

brachyurus, Sasia abnormis and Hemicircus concretus) from (Meiglyptes tristis) and Macaranga trees (Meiglyptes tristis
those who used arboreal ant nests (Meiglyptes tukki); and and Celeus brachyurus to a lesser extent).
dead liana/rattan foragers (Blythipicus rubiginosus) from Of the subguild of birds that foraged on these
dead wood patch (Picus mentalis and P. puniceus) and characteristically tropical substrates, Meiglyptes tristis
snag foragers (Reinwardtipicus validus, Dryocopus javensis, displayed the most noteworthy foraging preferences.
Mulleripicus pulverulentus). At the terminal branches, This species foraged primarily on leaves, quite often on
behavioural variables such as vertical placement and pioneer trees in logged forest, small trees in large treefall
foraging method were useful in separating individual gaps and even on bamboo leaves. However, M. tristis
species. The only exception to this is the species pair strongly preferred the leaves of Macaranga trees. Tree
Celeus brachyurus and Meiglyptes tristis. This is because, species in the genus Macaranga are prevalent pioneer
although more similar to one another than to other species in Malaysian forests, and some are known to
species, these two species exhibited numerous differences have mutualistic associations with ants in the genus
in their substrate preferences. Crematogaster. Meiglyptes tristis foraged frequently on
The most ecologically similar species, Picus mentalis both the leaves and stipules of Macaranga trees. ARS
and P. puniceus, differed significantly only for the disturbed these stipules on numerous occasions, and
parameter Attack Manoeuvre (chi-square = 16.5, df = 5, inevitably, small ants would run out to investigate. Tiny
P = 0.006), whereas other similar species groups differed ant nests were also found at the base of bamboo leaves.
significantly among at least three parameters (horizontal Since ants are the only known food item of M. tristis
placement (P < 0.0001), vertical placement (P < 0.0001) (Smythies 1981), it is likely that it was eating ants
and perch diameter (P = 0.001) for Sasia abnormis while foraging on leaves. When not foraging on leaves,
and Hemicircus concretus; horizontal position (P = 0.02), this species foraged on ant and termite tunnels on tree
vertical placement (P = 0.0004) and attack manoeuvre branches and also on ball-shaped ant nests suspended in
(P = 0.006) for Reinwardtipicus validus and Dryocopus trees. ARS confirmed that the nests were ant nests by
javensis; and horizontal position (P = 0.001), attack observing the organisms swarming out of and around
manoeuvre (P = 0.02), perch diameter (P < 0.0001) these nests while woodpeckers were foraging on them
and substrate (P = 0.0002) for Celeus brachyurus and (they were always ants).
Meiglyptes tristis). This study supports Short’s (1978) suggestions that
Malaysian woodpeckers employ a broad array of foraging
strategies, and suggests that the additional woodpecker
DISCUSSION diversity seen in tropical rain forests is to some degree
related to their specialization on microhabitats not
Analyses of foraging behaviour revealed that this diverse available in temperate forests.
guild of woodpeckers exhibited a great range in resource
use. Woodpeckers partitioned foraging resources spatially
by using different-sized perches (for woodpeckers, the ACKNOWLEDGEMENTS
perch is frequently also its foraging substrate) and by
foraging at different levels in the forest. These birds We thank The Ministry of Science, Technology and the
also exhibited a variety of microhabitat preferences Environment for IRPA funding for the fauna conservation
ranging from typical substrate use of live and dead project. We also thank Dr Abdul Razak, Director General
wood of various types to foraging primarily on ant or of the Forestry Research Institute of Malaysia, for
termite nests, bamboo and even leaves. The variables permission to conduct research at Pasoh Forest Reserve.
‘perch diameter’ and ‘substrate’ in particular were The Wildlife Group at UPM generously provided ARS
important in dividing this group into two primary with help and friendship (B. S. Fatt, N. F. b. Haneda,
subguilds: (1) ‘conventional’ woodpeckers – the larger- R. M. Lomoljo, S. Phirasack, S. A. Silang, S. H. b. M.
bodied woodpeckers that foraged either on dead-wood Afandi, T. b. Yussof and Z. b. Rosli). K. Ickes and J. Light
patches (Blythipicus rubiginosus, Picus puniceus and provided field assistance. J. V. Remsen, M. Wong and two
P. mentalis), or on snags (Reinwardtipicus validus, anonymous reviewers made constructive comments on
Dryocopus javensis and Mulleripicus pulverulentus), and the manuscript. ARS was funded in part by the Frank M.
(2) ‘novel’ woodpeckers – the smaller-bodied woodpeckers Chapman Memorial Fund from the American Museum
that foraged on substrates that do not typically occur of Natural History, the Charles M. Fugler Fellowship
in temperate forests such as bamboo (Sasia abnormis in Tropical Vertebrate Biology from the Louisiana State
and Hemicircus concretus to a lesser extent), external University Museum of Natural Science, the LSU Museum
arboreal ant nests (Meiglyptes tristis, M. tukki and of Natural Science Tropical Bird Research Fund and the
Celeus brachyurus), evergreen broadleaved vegetation LSU Museum of Natural Science Research Award.
Woodpecker ecology in Malaysian rain forest 493

LITERATURE CITED SHORT, L. L. 1978. Sympatry in woodpeckers of lowland Malayan

forest. Biotropica 10:122–133.
KOCHUMMEN, K. M., LAFRANKIE, J. V. & MANOKARAN, N. 1990. SMYTHIES, B. E. 1981. The birds of Borneo. (Third edition). The Earl of
Floristic composition of Pasoh Forest Reserve, a lowland rain forest Cranbrook (ed.). Sabah Society and Malayan Nature Society. Kuala
in Peninsular Malaysia. Journal of Tropical Forest Science 3:1– Lumpur. 593 pp.
13. SPSS Inc. 1998. SYSTAT 8.0. Chicago.
LAIDLAW, K. R. 2000. Effects of habitat disturbance and protected STYRING, A. R. & ICKES, K. 2001. Woodpecker abundance in a
areas on mammals of Peninsular Malaysia. Conservation Biology logged (40 years ago) vs. unlogged lowland dipterocarp forest in
14:1639–1648. Peninsular Malaysia. Journal of Tropical Ecology 17:261–268.
MILES, D. B. 1990. A comparison of three multivariate techniques WELLS, D. R. 1999. The birds of the Thai-Malay Peninsula. Academic
for the analysis of avian foraging data. Studies in Avian Biology Press, London. 647 pp.
13:295–308. WONG, M. 1985. Understory birds as indicators of regeneration in a
REMSEN, J. V. & ROBINSON, S. K. 1990. A classification scheme for patch of selectively logged West Malaysian rainforest. Pp. 249–266
foraging behavior of birds in terrestrial habitats. Studies in Avian in Diamond, A. W. & Lovejoy, T. E. (eds). Conservation of tropical forest
Biology 13:144–160. birds. International Council for Bird Preservation, Cambridge. 318 pp.
SAS Institute Inc. 1999. SAS version 8.00 for Windows. Cary, WONG, M. 1986. Trophic organization of understory birds in a
North Carolina. Malaysian dipterocarp forest. Auk 103:100–116.

Appendix 1 a–c. Ecological variables used to perform correspondence analysis (CA). Data are written as proportions. For abbreviations of species
names, see Table 1.
(a) Spatial variables. XS = perch < 5 cm diameter; S = perch ≥ 5 cm and < 10 cm; M = perch ≥ 10 cm and < 20 cm; L = perch is ≥ 20 cm and
< 35 cm; and XL = perch is > 35 cm. Understorey = height ≤ 5 m from the ground, Midstorey = height > 5 m from the ground and < 5 m from the
canopy, and Upperstorey ≥ 5 m from the canopy.
Height
Perch diameter Horizontal position
Under- Mid- Upper-
Species XS S M L XL storey storey storey Inner Middle Outer N
SA 0.70 0.30 0.00 0.00 0.00 0.53 0.20 0.28 0.70 0.25 0.05 40
HC 0.26 0.69 0.03 0.03 0.00 0.03 0.21 0.76 0.21 0.63 0.16 38
MT 0.28 0.58 0.10 0.00 0.04 0.16 0.42 0.42 0.14 0.29 0.57 126
MTU 0.32 0.42 0.13 0.10 0.03 0.39 0.32 0.29 0.65 0.35 0.00 31
CB 0.05 0.32 0.55 0.00 0.09 0.05 0.41 0.55 0.41 0.41 0.18 22
BR 0.00 0.29 0.47 0.06 0.18 0.65 0.29 0.06 0.88 0.12 0.00 17
PM 0.02 0.14 0.43 0.18 0.24 0.06 0.71 0.23 0.76 0.19 0.05 58
PMI 0.00 0.25 0.75 0.00 0.00 1.00 0.00 0.00 1.00 0.00 0.00 4
PP 0.00 0.09 0.38 0.41 0.12 0.11 0.51 0.38 0.68 0.32 0.00 38
DR 0.00 0.33 0.67 0.00 0.00 0.10 0.70 0.20 0.30 0.70 0.00 10
RV 0.00 0.06 0.12 0.33 0.49 0.15 0.64 0.21 0.83 0.13 0.04 54
DJ 0.00 0.00 0.03 0.36 0.61 0.03 0.36 0.61 0.64 0.36 0.00 36
MP 0.00 0.00 0.45 0.03 0.52 0.00 0.41 0.59 0.45 0.52 0.03 29

(b) Microhabitat variables (see text for definitions).

Microhabitat
Species Ant/Termite Bamboo Log Liana Dead patch Live wood Leaf Macaranga Snag Misc. N
SA 0.03 0.23 0.00 0.03 0.18 0.50 0.05 0.00 0.00 0.00 40
HC 0.00 0.05 0.03 0.00 0.26 0.58 0.00 0.00 0.00 0.08 38
MT 0.16 0.03 0.00 0.02 0.06 0.15 0.13 0.42 0.02 0.00 126
MTU 0.42 0.00 0.00 0.00 0.13 0.29 0.00 0.03 0.10 0.03 31
CB 0.18 0.18 0.00 0.00 0.14 0.27 0.00 0.09 0.14 0.00 22
BR 0.00 0.00 0.35 0.12 0.06 0.24 0.00 0.00 0.18 0.06 17
PM 0.03 0.02 0.00 0.02 0.57 0.29 0.00 0.00 0.05 0.02 58
PMI 0.25 0.00 0.00 0.00 0.25 0.00 0.00 0.00 0.50 0.00 4
PP 0.00 0.00 0.00 0.00 0.58 0.34 0.00 0.00 0.08 0.00 38
DR 0.00 0.10 0.00 0.00 0.50 0.30 0.00 0.00 0.10 0.00 10
RV 0.00 0.00 0.00 0.04 0.12 0.04 0.00 0.00 0.79 0.02 54
DJ 0.00 0.00 0.00 0.00 0.14 0.06 0.00 0.00 0.81 0.00 36
MP 0.00 0.00 0.00 0.00 0.24 0.34 0.00 0.00 0.41 0.00 29
494 ALISON R. STYRING AND MOHAMED ZAKARIA BIN HUSSIN

(c) Attack manoeuvre (see text for definitions).

Attack manoeuvre
Species Chisel Flake Glean Hammer Hang Peck Probe N
SA 0.00 0.00 0.20 0.50 0.00 0.30 0.00 40
HC 0.00 0.00 0.26 0.50 0.08 0.08 0.08 38
MT 0.01 0.17 0.24 0.10 0.10 0.12 0.25 126
MTU 0.03 0.00 0.23 0.26 0.00 0.48 0.00 31
CB 0.05 0.14 0.55 0.05 0.05 0.18 0.00 22
BR 0.00 0.06 0.00 0.35 0.00 0.18 0.41 17
PM 0.02 0.02 0.19 0.21 0.00 0.17 0.40 58
PMI 0.00 0.00 0.00 0.00 0.00 0.50 0.50 4
PP 0.08 0.03 0.08 0.32 0.00 0.39 0.11 38
DR 0.00 0.00 0.00 0.50 0.00 0.50 0.00 10
RV 0.33 0.04 0.00 0.35 0.00 0.24 0.04 54
DJ 0.03 0.11 0.03 0.61 0.00 0.19 0.03 36
MP 0.03 0.03 0.00 0.17 0.00 0.62 0.14 29

Appendix 2. Additive dissimilarity matrix of 11 woodpecker species (Dinopium rafflesii and Picus miniaceus not included due to small sample
size). For species abbreviations, see Table 1.
BR CB DJ HC MP MT MTU PM PP RV SA
BR 0.00
CB 0.46 0.00
DJ 0.51 0.44 0.00
HC 0.57 0.42 0.51 0.00
MP 0.51 0.41 0.37 0.52 0.00
MT 0.57 0.36 0.59 0.43 0.61 0.00
MTU 0.41 0.38 0.46 0.41 0.47 0.40 0.00
PM 0.34 0.39 0.47 0.51 0.42 0.49 0.38 0.00
PP 0.39 0.37 0.34 0.45 0.35 0.53 0.32 0.23 0.00
RV 0.44 0.56 0.27 0.64 0.45 0.60 0.44 0.35 0.35 0.00
SA 0.43 0.44 0.47 0.38 0.59 0.50 0.32 0.47 0.42 0.55 0.00