Suleman Raja Tropical Ecology and Conservation

Biology
3rd Year

Discuss the diversity and importance of animal-plant interactions in

tropical forest ecosystems

Introduction

Inter-specific symbiotic relationships, beneficial to both the concerned species – also

known as mutualisms, occur across a variety of ecosystems and bear essential
importance to the survival and success of many species. Tropical forest ecosystems
are no exception to this. It is the interactions between plants and animals which
interweave the development and continued biological flow of tropical rainforests in
many different ways. Of course, there are also plant to plant mutualistic relationships,
as well as animal to animal mutualisms which form the mainstay of adaptive life for
some species. Animal-plant relationships differ in that the linked support of life
between plants and animals is what gives rise to the maintenance of a successful and
diverse tropical forest ecosystem.

Being at the root of all food chains, plants form the essential base of rainforest
ecosystems. Perhaps the most important biological process for the survival of a forest
is pollination. In rainforests, animals play a vital role in the facilitation of this process,
as the abiotic factors which would normally allow successful pollination are limited in
rainforests. Such factors mainly being wind facilitated pollination – because of the
humidity and dense foliage in a tropical forest, pollination by wind–carriage is not a
successful means of reproduction for plants. Furthermore, due to the density of
tropical forests, wind pollination would not suffice to carry pollen far enough to
spread and mix genes beyond a certain area. This is where the importance of animal-
plant interactions is imperative. It is the animals in a tropical forest ecosystem which
are the main pollination agents for plants that make up the forest environment. Many
different species are involved in the pollination of plants – helping the plants to
reproduce and diversify genetically, whilst the animals benefit from food sources
provided by plants such as pollen and nectar.

Whilst it is not possible to cover all animal-plant interactions within the constraints of
a written essay, this report will endeavour to provide a comprehensive assessment on
the importance and diversity of animal-plant interactions in rainforests, giving a
selection of different types of interactions / mutualisms and examples of them, drawn
from field experience and reference to learned sources.

Pollinator mutualisms

Insects

The biodiversity of a rainforest can be assessed by the direct diversity of indicator

species such as insects. Rainforests support an incredible number of insect species –
to the extent that a huge many remain undiscovered or unidentified. Insects are
essential to the success and diversity of a tropical forest ecosystem as they act as the
primary pollinators of nearly all plant species in the forest. Vast numbers of insect
species use plants as a place of refuge or for food. Some examples of insects acting as
Suleman Raja Tropical Ecology and Conservation
Biology
3rd Year
pollinators are shown by the following examples of animal-plant interactions in
tropical forests:

Bees – Bees are important pollinators not only in tropical forests but for almost all
flowering plants present in a place where Bees also occur. Typically, Bees collect
nectar from the flowers of flowering plants. In doing this, the body of the Bee must
first brush past the anthers inside a flower, causing pollen to rub off onto the Bee’s
hairy body. The hairs on the body act to keep the pollen there whilst the Bee is in
flight. Therefore, when the Bee visits another flower, the pollen on its body, from
another plant – rubs off onto the stigma or female sexual organ of another flower
while the Bee pushes past this organ in order to obtain the nectar which is normally at
the bottom of a flower. In tropical forests, Bees act in this way to pollinate flowering
plants. Flying insects and animals are what these plants rely heavily on for
pollination, as rainforests do not get exposed to sufficient wind in order to carry
pollen through vast distances. The humidity levels are also too high for pollen to be
successfully blown from plant to plant in the air.

Fig wasp – This is a wonderful example of an extremely specialised pollination

mutualism in tropical ecology whereby both concerned species depend on one another
for their reproductive success and survival.
Figs are plants in the genus Ficus, which have a unique closed inflorescence called a
syconium, typically containing hundreds of flowers (Cook and West, 2005).
Fig trees (genus Ficus) have 755 species worldwide – around 511 of which are within
the Indo-Australasian tropics. (Rossiter, 2008)

The life cycle of the fig wasp starts

inside a fig, where the female deposits
her eggs. Upon hatching, the larvae
develop within the gallad ovary, using
the endosperm tissue of the ovary as a
food source for growth. As adults, they
chew their way out of the galls and
into the fig’s main cavity. Here, the
flightless male wasps fight to mate
with the females that have emerged
from the same clutch – whilst this
occurs, the fruit is in it’s male phase,
thus the bodies and legs of the wasps
get exposed to pollen in the mating stage of the fig wasp lifecycle. After mating,
males and females leave their natal fig – the males die at this stage, whereas the
females fly away to deposit their eggs (oviposition) inside a new fig before they die -
starting the cycle over.
Suleman Raja Tropical Ecology and Conservation
Biology
3rd Year

The morphology of a fig is such, that in order to enter

the fig, fig wasps must enter through the only
opening – called the ostiole. The female wasps collect
pollen from the anthers in its natal plant, before
coming into contact with the female part of a new fig.
This gives rise to very effective pollination because
the wasp gathers sufficient pollen from its natal fig to
pollinate the female reproductive parts of another fig
effectively upon contact with them.
When wasps enter, the fig is unripe and only the
female part is active – thus avoiding self pollination.
The female lays her eggs down the stylet into the
ovary of the fig – in doing this she brushes the pollen
of the natal plant from her legs onto the female part
of the new fig. Adaptations that the fig wasp has for
this lifestyle are a long, flattened and elongated head
and thorax designed for easier entry to the fig via the narrow ostiole. In fact, entering
the fig is still such a squeeze, that female fig wasps often lose their wings and
antennae in doing so.

This animal-plant interaction bears very

important implications to many birds,
primates, bats and other animals as the fig
produces fruit all year round, even when
other plants are not, thus providing a very
important food source to the aforementioned
species. This provision of food at otherwise
tricky times makes the fig a keystone plant
species (Cook and West, 2005), as it is
crucial in maintaining the populations and
diversity of rainforest fauna. The interaction
with the fig wasp keeps the reproductive
cycle of the fig successful, whilst also
providing a very effective means for the fig
wasp to carry out its reproductive cycle also. Both species benefit, as well as other
species – by the maintenance and continued production of figs.

Plants of genus Rafflesia – Plants from this genus have flowers which are known to
emit a repulsive odour of carrion – this is a mechanism which evolved in order to
attract flies to the flower for pollination. A particular species found in the rainforests
of Borneo, Sumatra and Malaysia - Rafflesia arnoldii, produces the largest individual
flower in the world. The flower can grow to be of a diameter of up to three feet and
produces a distinctive, putrid smell of rotting meat. The plant is rare as it grows as a
parasite of the Tetrastigma vine, found only in primary forests. This highly specialised
plant lacks any visible stems, leaves or roots – and grows in a way similar to fungi.
Suleman Raja Tropical Ecology and Conservation
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3rd Year
The smell it omits is used to attract flies – which settle on
the flower and carry pollen from one flower to another –
given the rarity of the plant and the fact that it only lasts
as a flower for a few days – pollination is a rare event,
thus the population of this flower is relatively low. Flies
make good pollinators as they seek out carrion and can
cover large distances. The strength of the odour therefore
plays an integral role in attracting flies for pollination.
This is extremely important for Rafflesia as it relies on
flies for reproductive success.

Birds – Birds also interact with plants as pollinators –

although to a lesser extent than the invertebrate
pollinators. Bird-pollinated flowers are usually tubular,
unscented flowers with a large store of nectar at the base – some birds feed on this
nectar whilst others are attracted to the insects which are attracted to the nectar – in
either instance, birds such as hummingbirds and sunbirds will collect pollen on their
bodies and transfer this to other flowers that they visit.

Bats – Another vertebrate pollinator. Rainforests are home to a vast array of Bat
species – some of which feed on nectar and pollen. Most Bats that pollinate plants are
from the family Pteropodidae – the fruit bats and flying foxes. It is such Bats, as well
as some of the leaf-nosed bats that pollinate many plants in tropical forests. The
flowers which are pollinated by Bats usually bloom by night, and omit musky or
fruity odours to attract Bats to them. Furthermore, they are often large and strong
enough to support a Bat. The plant and its pollinator often co –evolve to highly
specialised specifics in order to maximise the
mutualistic relationship between them. For example
with bat-pollinated flowers, the flowers as mentioned,
smell fruity and often lack any vivid colouration as
they are nocturnal like the bats are. Therefore they
need not express vivid colouration. In addition to this,
the bats rely more on olfactory senses to locate
flowers – as it is thought that bats lack cones in their
retinas and thus no not rely on colour vision to seek
out food sources.

The importance of animal-plant interaction with regards to pollination are not just
bound to the reproduction and survival of a forest – but also bears great economical
importance for human beings – 1/3 of mouthfuls of food and beverages consumed by
humans come from animal pollination. Moreover, products of insect pollination are
worth $40 billion annually in the US alone. (Rossiter, 2008)
The importance of animal pollination was demonstrated with the example of the West
African oil palm Elaeis guineensis which was introduced to Malaysia. For years the
palm was not able to survive here without hand pollination. Work by botanist R. Syed
in the 1970’s drew the conclusion that this was due to the fact that Malaysia lacked
populations of its specialised pollinator species – the pollinating Weevil Elaeidobius
spp. When this species was introduced and released in Malaysia the oil palms here
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3rd Year
flourished and spread considerably without the need for human intervention for
pollination. The oil palm industry is now a multi-million pound industry in Malaysian
plantations due to the introduction of its pollinator species – this demonstrates how
important one species of animal can be to a plant from its interactions with it.

Conservation of pollinators and plants is an issue of rising concern due to the loss of
wild pollinators. This ‘pollination deficit’ crisis has come about due to the loss of
habitat, spread of disease, pests and deforestation.

“Humanity, for its own sake, must attend to the forgotten pollinators and their
countless dependent plant species” (E.O Wilson)

Seed dispersal mutualisms

Seed dispersal is also a very important process carried out by animals in rainforests.
Following pollination, plants also rely on their animal counterparts to disperse and
spread fertilised seeds. The animals involved in seed dispersal benefit by gaining food
sources in the process – and the plants benefit by successful distribution of their
progeny.

Seed dispersal occurs by invertebrates (ants, beetles, other insects) as well as

vertebrates (birds, bats, rodents, other mammals). Myrmecochery is the term for seed
dispersal by ants. Seeds of many plants contain the accessory organ known as
elaiosomes which are rich in protein and attract ants. This is because ants will remove
and carry the elaiosomes to their nests to feed them to their larvae. Once the larvae
have eaten the elaiosomes, the adult ants will dispose of the inedible seed outside of
the nest or under the ground – where it will germinate. Ants are considered the main
invertebrate dispersers and many tropical plants rely on them to disperse their seeds.

Vertebrate dispersers in tropical rainforests mainly consist of frugivorous birds and

bats. Birds are normally attracted to brightly coloured fruit as a food source. The birds
/ bats that feed on such fruits will ingest its seeds whilst eating the pulp. The seeds are
indigestible and when excreted, become dispersed in the otherwise dense forest.
Orang-utans, gibbons and other primates are also very important seed dispersers in a
similar way within tropical forests. Fruits grow in a variety of shapes and sizes,
specific to different dispersers’ preferences. For instance, larger, bigger-seeded fruits
are consumed by progressively fewer dispersers, and the largest depend on a few
species of mammals and birds which are highly vulnerable to hunting, fragmentation
and habitat loss (Corlett, 1998). This is why conservation of species is hugely
important in the knock-on effect the loss of just a single species can have on other
species including plants.
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Decomposition

The diversity of animal-plant interactions does not stop at pollination and dispersal.
Decomposition is also imperative to the nutrient cycling and success of a forest where
dead organic plant matter inevitably accumulates. Some old living trees also contain
dead wood structures within them and these provide food sources for saproxylic
organisms which rely on dead organic matter in some part of their lifecycles. Aside
from fungi and bacteria, animals which carry out decomposition in rainforest include
Annelids (worms), Arthropods (woodlice), Molluscs (snails) and Insects.

Termites - Prolific decomposers of dead wood and other plant matter such as roots
and leaf litter are termites (order Isoptera) – these insects are found in huge numbers
throughout tropical forests and play a vital role in the ‘tidying up’ of dead plant matter
in the forest, on which they rely for food.
The importance of termites and their interactions with plants is vital for keeping a
forest free of dead matter, particularly large logs of wood which would normally take
considerably long to decompose without the termites feeding on it.

Defence and shelter

An example drawn from the field exhibiting an

exclusive and highly specialised mutualism
between plant and animal is that of the Bornean
plants in the genus Macaranga. The mutualism
between the Macaranga plants and their ants is
known as a Myrmecosymbiosis. The ants live
inside the structures on the stems known as
stipules, and in the hollow shoots of the plant,
where the plant provides them with so-called
food bodies rich in lipids, carbohydrates and
proteins (Linsenmair, 2001). These are
extrafloral nectaries from which the ants feed. In
return for this energetically costly provision, the
ants provide biotic defence / protection to the
plant from herbivorous predictors such as caterpillars and other insects which may
graze on the leaves of the Macaranga. The ants will patrol the plant and defend their
biotic host by killing any intruders they come across on the plant. Some macaranga
plants are specialised so as to host just one particular species of ant, by having waxy
stems that are not climbable by other species due to their slippery texture. This
relationship is very important for the concerned species, and studies show that ant
colony size on a macaranga plant is indeed negatively correlated with leaf damage
caused by herbivory (Linsenmair, 2001). The importance of this relationship to both
species is clear - ants gain food and shelter, whilst the plant gains defence and thus
better survival. This example further highlights how diverse and specialised animal-
plant interactions in tropical forests can be.
Suleman Raja Tropical Ecology and Conservation
Biology
3rd Year
Less mutualistic animal-plant interactions

Some of the interactions between plant and animal are not beneficial to both parties
concerned and thus not all animal-plant interactions should be considered as
mutualisms. For example, rainforests are home to many species of predatory plants –
which feed on insects by trapping and killing them. There are around 600 species of
predatory plants (Rossiter, 2008) which divide into three sub-categories by trapping
mechanism. These are the pitfall traps, snap traps, and flypaper traps.

Flypaper traps – As the name suggests, these plants operate in a way similar to fly
paper – they produce a sticky mucilage secretion which acts to entrap insects which
either walk over the plant or land on it from flight. They are often found growing in
bogs where nutrient levels are low and derive their nutrients from consuming and
digesting the insects they trap.
The multiple, independent evolution of carnivory in diverse plant families suggests
that it is an adaptation to the low nutrient, bright, waterlogged habitats in which
carnivorous plants occur (Ellison and Gotelli, 2001)

These plants have evolved independently in five dicot families (Lentibulariaceae,

Roridulaceae, Biblidaceae, Droceraceae and Dioncophyllaceae) The Droceraceae
consists of three genera (Drosera, Aldrovanda and Dionaea)
(Ellison and Gotelli, 2001)
The Drosera genera are known commonly as the sundews and contain the most
species worldwide (around 150 spp). The other two genera Aldrovanda and Dionaea
are Snap traps - A very well known example of this is the Venus fly trap Dionaea
muscipula.

These plants have tiny receptor hairs inside the cavity of the trap, which when
triggered by the presence of an insect, cause the two discs forming the trap to snap
shut – enclosing the insect and sealing the edges with interlocking spikes to secure the
prey. Once closed, the Venus fly trap will consume and digest the prey by enzymatic
breakdown and absorption before re-opening.
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Pitfall traps – Finally, pitfall traps are also aptly named by what they do – the plant
structure is like a pitfall – when insects fall into the fluid-filled cavity of the plant,
they cannot escape due to the waxy inside walls of the
pitfall – they therefore drown or perish within the
bounds of the plant and are subsequently consumed
by the plant.

Pitcher plants of the genus Nepenthes are examples of

tropical pitfall traps. They occur mainly in Borneo
and the genus contains around 117 species (Rossiter,
2008). Locally, these plants are known as ‘Monkey
cups’ due to their cup-like structure which holds
water. Indeed, it has been observed that Monkeys will
sometimes pick these plants and drink from them like
a cup. However, the main interaction these plants
have with animals is that of their prey species. Insects
are attracted to nectaries located along the entrance to
the pitcher. When they settle on the rim the small
insects usually slip and fall in – where they are met
by water and very waxy walls to prevent escape. Most prey insects are ants, termites
or midges which drown and accumulate in the fluid.
Furthermore, pitcher plants also play a part in mutualistic interactions with insects as
well as the predator-prey interaction. Some invertebrates live as ‘infauna’ and inhabit
the pitcher without dieing. These include mosquito larvae as aquatic filter feeders and
predators, as well as midge larvae as detritus feeders. By inhabiting the pitcher,
infauna species help with the break down of prey and thus provide a mutualism by
providing the plant with nutrients as well as obtaining food and shelter for itself in the
pitcher. Moreover, some pitcher species are colonised by swimming ants which
inhabit the plant’s tendrils. These ants also form a mutualism with the plant in that
they remove prey items from full pitchers – this causes the plants with ants to have
fewer putrid pitchers which prevents the death of infauna by the prevalence of anoxic
conditions in the presence of too many dead insects inside the pitcher.

Some species of pitcher such as the massive Nepenthes rajah prey upon vertebrates
such as birds, lizards, frogs and rodents
such as mice and rats. (Rossiter, 2008).
This is a rare animal-plant interaction
whereby vertebrates are preyed upon by a
plant. This further highlights how diverse
animal-plant interactions can be within an
ecosystem.
Suleman Raja Tropical Ecology and Conservation
Biology
3rd Year

Conclusion

The aforementioned animal-plant interactions are just a few examples of such

relationships in tropical rainforest ecology. The purpose was to provide an all round
guide on the diversity and importance of these interactions as both a factor on the
survival of the forests as well as implications to human beings. The types of
relationships covered included pollinators, dispersers, decomposers, defence and
shelter relationships and plant predation. It is stressed that these subheadings only
touch on a selected number of types of animal-plant interactions in order to outline the
diversity of animal-plant interactions and their importance. In tropical forests, there
are countless mutualisms and animal-plant interactions which are very diverse and
also serve huge importance to ecosystem structure and function.

References

 Rossiter, S, 2008. Animal-plant interactions (Lecture, Brunei)

 Bascompte, J and P Jordano, 2007. Plant-Animal Mutualistic Networks: The

Architecture of Biodiversity: Annual Review of Ecology, Evolution, and
Systematics. 38: 567-593

 Boucher, DH et al, 1982. The Ecology of mutualism. 13: 315-347

 Knudsen, J. T. & L. Tollsten. 1995. Floral scent in bat-pollinated plants: a

case of convergent evolution. Bot. J. Linn. Soc. 119: 45-57.

 Cook, J. M and S West, 2005. Figs and fig wasps: Current Biology Vol 15: 24

 Corlett, R.T, 1998. Frugivory and seed dispersal by vertebrates in

the Oriental (Indomalayan) Region: Biological review., 73: 413-448

 Linsenmair, K. E et al., 2001. Adaptations to biotic and abiotic stress:

Macaranga-ant plants optimise investment in biotic defence. Journal of
Experimental Botany, 52(363), pp.2057-2065

 Ellison, A.M and J Gotelli, 2001. Evolutionary ecology of carnivorous plants.

Trends in ecology and evolution., 16 (11)