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THE RAFFLES BULLETIN OF ZOOLOGY 2008
FEEDING PREFERENCES OF PHASMIDS (INSECTA: PHASMIDA)
IN A BORNEAN DIPTEROCARP FOREST
Robert R. Junker
Department of Animal Ecology and Tropical Biology, Biozentrum, University of Würzburg, 97074 Würzburg, Germany
Email: r.junker@biozentrum.uni-wuerzburg.de
Takao Itioka
Graduate School of Human and Environmental Studies, Kyoto University, Sakyo, Kyoto 606-8501, Japan.
Email: itioka@mbox.kudpc.kyoto-u.ac.jp
Phil E. Bragg
8, The Lane, Awsworth, Nottinghamshire, NG16 2QP, U.K.
Nico Blüthgen
Department of Animal Ecology and Tropical Biology, Biozentrum, University of Würzburg, 97074 Würzburg, Germany
Email: bluethgen@biozentrum.uni-wuerzburg.de (Corresponding author)
ABSTRACT. – Stick and leaf insects (Phasmida) representing 19 species (53 individuals) were
collected in a lowland dipterocarp forest (Lambir Hills, Sarawak, Malaysia). Dual-choice tests were
conducted to examine whether phasmids discriminate between young and old leaves of seven plant
species. A second set of tests examined the preferences of phasmids for leaves from Dryobalanops
lanceolata (Dipterocarpaceae) saplings versus leaves from the upper canopy of the same tree species.
Haaniella echinata and other flightless species (Heteropterygidae and Lonchodinae) fed on nearly
all plant species offered and showed significant preferences for old leaves in three plant species.
In contrast, flying phasmids (Ashiphasmatinae and Necrosciinae) rejected leaves from most plants
and did not show consistent leaf age choices. H. echinata and flightless phasmids preferred canopy
leaves from D. lanceolata over leaves from saplings, regardless of leaf age. Our results are consistent
with the hypothesis that young leaves of some plant species are better defended against generalist
herbivores than old leaves and that saplings are better defended than adult trees. Since upper canopy
leaves were highly palatable to understorey phasmids, factors other than chemical defences must
contribute to the low abundance of phasmids in forest canopies.
KEY WORDS. – Dryobalanops lanceolata, forest canopy, Haaniella echinata, herbivory, leaf age,
lowland rainforest.
RAFFLES BULLETIN OF ZOOLOGY 2008 56(2): 445–452
Date of Publication: 31 Aug.2008
© National University of Singapore
INTRODUCTION
In virtually all ecosystems, herbivorous insects constitute a
large proportion of the animal species and consume foliage,
plant sap, or other plant parts from nearly the entire flora. The
diversity of insect herbivores is highest in tropical rainforests
(Novotny et al., 2004). Woody plants in tropical forests are
more damaged by herbivores than those from temperate
forests (Coley & Barone, 1996), but this increased damage is
not necessarily accompanied by a higher density of herbivore
individuals (Novotny et al., 2006). Comparative surveys on
food plant selection of tropical herbivore communities have
begun to unravel their actual host plant ranges and preferences
(Basset, 1996; Barone, 1998; Basset, 1999; Novotny & Basset,
2005; Novotny et al., 2006). While many beetles and butterfly
caterpillars are plant genus specialists, orthopteroids are often
polyphagous (Novotny et al., 2002). However, for most
species the natural host range is unknown. Few studies have
compared the host plant selection of rainforest orthopteroid
communities (e.g. grasshoppers, Rowell, 1978), including
Phasmida (Tay & Seow-Choen, 1996; Novotny et al., 2002;
Blüthgen et al., 2006). Phasmids and other orthopteroids
such as katydids are usually less numerous than several other
tropical insect taxa, particularly in comparison to the large
holometabolous orders (Coleoptera, Diptera, Hymenoptera
and Lepidoptera). In canopy samples from Southeast Asian
rainforests, orthopteroids typically contributed less than 3%
of the arthropod individuals (Stork, 1987; Stork & Brendell,