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Pollination and Reproductive Ecology of
Three Lowland Tropical Pitcher Plants (Nepenthes) and Their Hybrids
Christopher K. Frazier
Biology Department University of New Mexico
Albuquerque, NM 87131 USA
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Tropical pitcher plants have been the subject of substantial
scientific scrutiny for their prey digestion (Frazier, 2000), but other
aspects of their biology, particularly their reproductive ecology have
garnered much less attention. Species in the genus Nepenthes are
interfertile under greenhouse conditions, yet in the field demonstrate
a high degree of sympatry. This suggests that differences in reproductive
strategies may play an important role in reducing hybridization and maintaining
species distinction under natural conditions. I studied the three species
of Nepenthes present in Johore, Malaysia and Singapore, N. gracilis,
N. ampullaria and N. rafflesiana, to determine if there
are significant differences among species in phenology, floral and nectar
characteristics and pollinators. I also addressed the fitness of naturally
occuring hybrids. Differences in flowering period represents at best a
partial barrier to cross-pollination among species. Each species has three
flowering periods between January and July, but the timing of these episodes
is offset among species such that at any given time one, two or all three
species are in flower. Floral presentation differs among species in such
characters as the number of flowers per inflorescence, flower size and
color and sepal shape (Frazier, in press). N. ampullaria flowers
present pollen during the day, while the other two species open their
flowers in the evening. Nectar is produced nocturnally in all three species,
but they differ in sugar composition and quantity of nectar produced.
These differences suggest the species are adapted to attract different
suites of pollinators. All species are pollinated by moths and small Diptera,
but to differing degrees. N. rafflesiana is pollinated almost entirely
by moths and all of its pollination is nocturnal. N. gracilis is
pollinated almost equally by moths and small Diptera at night and also
by wasps during the night and day. By day, N. ampullaria flowers
have a foul odour and are pollinated by large flies, wasps and butterflies.
The flowers produce a sweet odour at night and are then visited by moths.
These differences are significant between species, but still not sufficient
to constitute a complete barrier to cross-pollination.
Low hybrid fitness may also act as a barrier to gene
flow between the parent species. I found that hybrids in the wild abort
significantly more flowers, produce fewer seeds per inflorescence and
have a lower efficiency of seed production (seeds produced per gram of
inflorescence) than the parent taxa. Nevertheless, some hybrid inflorescences
demonstrated zero flower abortion and produced seed quantities on par
with the parent taxa, suggesting that ecological phenomena and not intrinsic
sterility barriers are the main factors limiting hybrid fitness.
My results show that many factors interact to limit,
but not preclude hybrid production and that lower hybrid fitness comprises
a further, though incomplete barrier to gene flow. I suggest that the
combination of such partial barriers allows sympatric species to retain
species distinction without intrinsic sterility barriers, although some
degree of interspecific gene flow is still likely.
References
Frazier, C.K. (in press) Pollination and reproductive
ecology of Nepenthes. In: Clarke, C., Nepenthes of Sumatra and
Peninsular Malaysia, Natural History Publications, Kota Kinabalu, Malaysia.
Frazier, C.K. 2000 The enduring controversies concerning
the process of protein digestion in Nepenthes (Nepenthaceae).
Carniv. Pl. Newslett., 29:2, 56.
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