Ch. Schulze et al., Understorey versus canopy: patterns of vertical stratification and diversity among Lepidoptera in a Bornean rain forest, PLANT ECOL, 153(1-2), 2001, pp. 133-152
We studied the vertical distribution of Lepidoptera from a canopy walkway w
ithin a dipterocarp rain forest at Kinabalu Park (Borneo) using three diffe
rent methods: (1) Bait traps to survey fruit-feeding nymphalid butterflies,
(2) standardized counts for predominantly flower-visiting butterflies and
their potential predators, aerial-hawking birds, and (3) attraction by blac
klight for hawk- and tiger moths. There was a distinct decrease in the abun
dance of fruit-feeding nymphalids towards the canopy, probably due to a red
uced and less predictable availability of rotting fruits in higher strata.
These constraints might also be responsible for a higher abundance variatio
n in the canopy, and a significant shift in size from larger species in the
understorey to smaller ones in the canopy. Changes of microclimate and the
conspicuous increase of insectivorous aerial-hawking birds from ground to
canopy layer may be responsible for the prominent change in species composi
tion of fruit-feeding nymphalids between 20 and 30 m. Nectar-feeding Lepido
ptera showed a reversed abundance pattern. One main factor contributing to
the much higher abundance of flower-visiting butterflies and moth taxa in t
he canopy, such as Sphingidae and some Arctiinae, might be the increase of
nectar resources available in upper vegetation layers. A distinctly higher
diversity in hawkmoths was also found in the canopy. A higher abundance of
insectivorous aerial-hawking birds in the canopy might contribute to the sh
ift in body design of fruit-feeding nymphalids from more slender bodies at
lower vegetation layers to stouter ones (i.e. species which are stronger on
the wing) in the canopy. Larval resources could play an additional role in
specialisation on but a small part of the vertical gradient. This may expl
ain stratification pattern of the nymphalid subfamilies Morphinae and Satyr
inae. Monocotyledoneous larval food plants of both taxa, whose flight activ
ity is largely restricted to the understorey, occur mostly in lower vegetat
ion layers. Our observations on a wide taxonomic and ecological range of bu
tterflies and moths indicate that tropical forest canopies hold a distinct
and unique Lepidoptera fauna, whose species richness and abundance patterns
differ from lower strata. However, the notion of tropical forest canopies
as peaks of terrestrial diversity does not hold uniformly for all taxa or g
uilds.