SEED DISPERSAL

Herbivores can be important agents of dispersal for plants, even if they are not primarily seed or fruit eaters. Animal dung provides an ideal environment for the germination of seeds and the growth of seedlings. The fibrous nature of undigested plant material means that it has good moisture retention qualities. It is rich in nitrogen, as well as organic matter which is easily broken down to inorganic nutrients by microorganisms. Many seeds germinate better if they pass through a digestive tract, and some will only germinate after a hard protective layer has been partly digested away.

One of the best places for a germinating seed to find itself is in a patch of elephant dung, and elephants may be important agents of dispersal for many plants. Elephants love the seed pods of the apple-ring acacia (Acacia albida), a tree which is particularly important in their diet in some areas for it has green leaves during the dry season when most other plants are desiccated and of low quality. The only natural seedlings of this tree I have ever seen were growing out of elephant dung. Grasses and other herbs are also frequently seen germinating in patches of elephant dung. Elephants have a passion for the northern ilala palm (Hyphane benguellensis), which they often knock down just to chew the sweet bases of the leaves. But all of the benefit doesn't go to the elephant, for they also eat the fruit, and in the process may help to disperse the seeds. It is not uncommon to find northern ilala palms growing in single file along major routes of elephant movement.

I have also seen seedlings of many plant species growing out of the dung of giraffe, buffalo, kudu, eland, and to a lesser extent that or the smaller antelope such as impala and waterbuck. Although baboons are not herbivores, being omnivores they eat a considerable amount of plant material, including seeds and fruits. They are highly mobile, and so can be important agents of dispersal.

While it is true that herbivores in general are often important agents of dispersal, it is the specialist fruit eaters which provide some of the more interesting cases. The production of fleshy fruits represents a considerable drain on a plant's energy supplies. The loss of energy which occurs when a fruit is eaten represents an energetic cost to the plant. This cost must be more than balanced by a greater benefit, otherwise fleshy fruits could not evolve. The benefit to the plant is of course the dispersal of its seeds. Edible fleshy fruits exist only because they are a plant's means of tricking animals, including humans, into dispersing their seeds. One can argue that many of them have been highly successful, for many fruit-bearing plants have succeeded in getting us to plant and tend them in large stands, which we call orchards.

Figs must be one of the most fascinating groups of frugivore dispersed plants, with the strangler figs being the most unusual. The strangler figs, which can become very large trees, often start life when a seed is deposited in the branches of another tree within the droppings of a frugivorous bird or mammal. The seed germinates, and aerial roots wind their way down the trunk of the host tree, eventually reaching the soil and developing into normal underground roots. The aerial parts of these roots thicken and form the stem of the fig tree, and with time the host tree is strangled. These plants can also grow as normal trees if the seeds are dropped on the soil.

The size to which fruits develop on ripening is probably determined evolutionarily by the mouth sizes of the suite of frugivores which are available to eat them. Producing a large fruit is obviously more costly to the plant than producing a small fruit, so there is a low likelihood that large fruits will evolve unless this gives some increase in survival value. Specialist frugivores are uncommon at high latitudes, and most edible fruits are probably consumed by small birds. Not surprisingly, large fleshy fruits are rare at high latitudes. In the tropics, where there are many large frugivorous birds, as well as fruit-eating omnivores such as monkeys, baboons, humans and other primates, larger edible fruits are more common.

Frugivorous birds, such as the go-way bird (upper) and louries (lower), consume fruits and in doing so they help to disperse the seeds.
Edible fleshy fruits exist only because they are a plant's means of tricking animals, including humans, into dispersing their seeds (partridgeberry, Canada).

Granivores may also function as agents of dispersal, in spite of the fact that they actually eat and digest seeds. Often granivores remove seeds from the site of production to one or more caches, where they may germinate if conditions are favourable.

One of the more interesting cases of dispersal of seeds by animals occurs in the Fynbos Biome, where some seeds are dispersed by ants. The plants provide an edible structure, called an eliasome, on the seed and this encourages the ants to take the seeds to their nests. The ants eat the eliasomes, leaving the seeds free to germinate under the right circumstances.