Frugivore

A frugivore /fruːdʒᵿvɔːr/ is a fruit eater. It can be any type of herbivore or omnivore where fruit is a preferred food type. Because approximately 20 percent of all mammalian herbivores also eat fruit, frugivory is considered to be common among mammals.^[1] Since frugivores eat large volumes of fruit, they are highly dependent on the abundance and nutritional composition of fruits. Frugivores can either benefit fruit-producing plants by dispersing seeds, or they can negatively affect plants by digesting seeds along with the fruits. When both the fruit-producing plant and the frugivore species benefit by fruit-eating behavior, their interaction is called mutualism.

Frugivore seed dispersal

Seed dispersal is important for plants because it allows their progeny to move away from their parents over time. The advantages of seed dispersal may have led to the evolution of fleshy fruits, which entice animals to eat the fruits and move the plants seeds from place to place. While many fruit-producing plant species would not disperse far without frugivores, they can usually germinate even if they fall to the ground directly below the parent plant.

Many types of animals are seed dispersers. Mammal and bird species represent the majority of seed-dispersing species. However, frugivorous tortoises, lizards, amphibians, and even fish also disperse seeds.^[2] While frugivores and fruit-producing plant species are worldwide, there is some evidence that tropical forests have more frugivore seed dispersers than the temperate zone.

Plant adaptations to attract dispersers

There are a number of fruit characteristics that seem to be adaptive characteristics to attract frugivores. Many animal-dispersed fruits advertise their palatability to animals with bright colors and attractive smells (mimetic fruits).^[3] Fruit pulp is generally rich in water and carbohydrates and low in protein and lipids. However, the exact nutritional composition of fruits varies widely. The seeds of animal-dispersed fruits are often adapted to survive digestion by frugivores. For example, seeds can become more permeable to water after passage through an animal's gut. This leads to higher germination rates.^[4] Some mistletoe seeds even germinate inside the disperser's intestine.^[4]

Frugivore adaptations for fruit consumption

In order for frugivores to be good seed dispersers, they must digest fruits without consuming a high proportion of the seeds. Many seed-dispersing animals have specialized digestive systems to process fruits, which leave seeds intact. Some bird species have shorter intestines to rapidly pass seeds from fruits, while some frugivorous bat species have longer intestines. Some seed-dispersing frugivores have short gut-retention times, and others can alter intestinal enzyme composition when eating different types of fruits.^[2]

Plant mechanisms to delay or deter frugivory

Further information: Plant defense against herbivory

Plants invest energy into the production of fruits. Plants have evolved to encourage mutualist frugivores to consume their fruit for seed dispersal, but also evolved mechanisms to decrease consumption of fruits when unripe and from non-seed-dispersing predators. Predators and parasites of fruit include seed predators, insects, and microbial frugivores.^[5] Plants have chemical and physical adaptations.

Physical deterrents:^[6]

Cryptic coloration (e.g. green fruits blend in with the plant leaves)
Unpalatable textures (e.g. thick skins made of anti-nutritive substances)
Resins and saps (e.g. prevent animals from swallowing)
Repellent substances, hard outer coats, spines, thorns

Chemical deterrents:

Chemical deterrents in plants are called secondary metabolites. Secondary metabolites are compounds produced by the plant that are not essential for the primary processes, such as growth and reproduction. Toxins might have evolved to prevent consumption by animals that disperse seeds into unsuitable habitats, to prevent too many fruits from being eaten per feeding bout by preventing too many seeds being deposited in one site, or to prevent digestion of the seeds in the gut of the animal.^[7] Secondary chemical defenses are divided into three categories: nitrogen-based, carbon-based terpenes, and carbon-based phenolics.

Examples of secondary chemical defenses in fruit:

Capsaicin is a carbon-based phenolic compound only found in plant genus Capsicum (chili and bell peppers). Capsaicin is responsible for the pungent, "hot" flavor of peppers and inhibits growth of microbes and invertebrates.^[5]
Cyanogenic glycosides are nitrogen-based compounds and are found in 130 plant families, but not necessarily in the fruit of all the plants. It is specifically found in the red berries of the genus Ilex (holly, an evergreen woody plant). It can inhibit electron transport, cellular respiration, induce vomiting, diarrhea, and mild narcosis in animals.^[7]
Emodin is a carbon-based phenolic compound in plants like rhubarb. Emodin can be cathartic or act as a laxative in humans, kills dipteran larvae, inhibits growth of bacteria and fungi, and deters consumption by birds and mice.^[5]

Examples

Birds are a main focus of frugivory research. An article by B.A. Loisell and J.G. Blake, Potential Consequences of Extinction of Frugivorous Birds, discusses the important role frugivorous birds have on ecosystems. The conclusions of their research indicate how the extinction of seed-dispersing species could negatively affect seed removal, seed viability, and plant establishment. This article highlights the importance that seed-dispersing birds have on the deposition of plant species.^[8]

Examples of seed-dispersing birds are the hornbill, the toucan, the aracari, the cotinga (ex. Guianan cock-of-the-rock), and some species of parrots. Frugivores are common in the temperate zone, but mostly found in the tropics. Many frugivorous birds feed mainly on fruits until nesting season, when they incorporate protein-rich insects into their diet.

Many types of animals are seed dispersers. Mammals and bird species represent the majority of seed dispersing species, however, frugivorous fish, tortoises, lizards, and even amphibians also disperse seeds.^[2]

Mammals are considered frugivorous if the seed is dispersed and able to establish. One example of a mammalian frugivore is the maned wolf, or Chrysocyon brachyurus, which is found in South America. A study by J.C. Motta-Junior and K. Martins found that the maned wolf is probably an important seed disperser. The researchers found that 22.5–54.3% of the diet was fruit.^[9]

Another example of mammalian frugivores is the orangutan, whose diet consists of 65% fruit. Orangutans primarily eat fruit, along with young leaves, bark, flowers, honey, insects, and vines. One of their preferred foods is the fruit of the durian tree, which tastes somewhat like sweet, cheesy, garlic custard. They discard the skin, eat the flesh, and spit out the seeds.

Another example of a mammalian frugivore is the gray-bellied night monkey, also known as the owl monkey:^[10]

“Owl monkeys are frugivores and supplement their diet with flowers, insects, nectar, and leaves (Wright 1989; 1994). They prefer small, ripe fruit when available and in order to find these, they forage in large-crown trees (larger than ten meters [32.8 ft]) (Wright 1986). Seasonal availability of fruit varies across environments. Aotus species in tropical forests eat more fruit throughout the year because it is more readily available compared to the dry forests where fruit is limited in the dry season and owl monkeys are more dependent on leaves.^[11]”

Fruit bats are another example of a mammalian frugivore. See the Megabat article about Diet and Importance.

Cassowaries are a keystone species because they spread fruit through digestion, and many seeds will not grow unless they have been digested by a cassowary.

Ecological significance of frugivore seed dispersal

Frugivore seed dispersal is a common phenomenon in many ecosystems. However, it is not a highly specific type of plant–animal interaction. For example, a single species of frugivorous bird may disperse fruits from several species of plants, or a few species of bird may disperse seeds of one plant species.^[12] This lack of specialization could be because fruit availability varies by season and year, which tends to discourage frugivore animals from focusing on just one plant species.^[2] Furthermore, different seed dispersers tend to disperse seeds to different habitats, at different abundances, and distances, depending on their behavior and numbers.^[13]

Conservation

Because seed dispersal allows plant species to disperse to other areas, the loss of frugivores could change plant communities and lead to the local loss of particular plant species. Since frugivore seed dispersal is so important in the tropics, many researchers have studied the loss of frugivores and related it to changed plant population dynamics. Several studies have noted that even the loss of only large frugivores (i.e. monkeys) could have a negative effect, since monkeys are responsible for certain types of long-distance seed dispersal that is not seen with other frugivore types, like birds.^[13] However, plant species whose seeds are dispersed by animals may be less vulnerable to fragmentation than other plant species.^[14] Frugivores can also benefit from the invasion of exotic fruit-producing species and can be vectors of exotic invasion by dispersing non-native seeds.^[15] In short, while anthropogenic habitat loss and change may negatively affect some frugivore species, it may positively affect others.

References

↑ Danell, K. and R. Bergstrom. 2002. Mammalian herbivory in terrestrial environments.in C. M. Herrera and O. Pellmyr, editors. Plant-Animal Interactions. Blackwell Publishing, Oxford, UK.
1 2 3 4 Herrera, C. M. 2002. Seed Dispersal by vertebrates.in C. M. Herrera and O. Pellmyr, editors. Plant-Animal Interactions. Blackwell Publishing, Oxford, UK.
↑ Galetti, M. 2002. Seed Dispersal and Mimetic Fruits: Parasitism, Mutualism, Aposematism, or Exaptation?in D. J. Levey, W. R. Silva, and M. Galetti, editors. Seed Dispersal and Frugivory: Ecology, Evolution, and Conservation. CABI Publishing, Oxford, UK.
1 2 Traveset, A., J. Rodriguez-Perez, and B. Pias. 2008. Seed trait changes in dispersers' guts and consequences for germination and seedling growth. Ecology 89:95-106.
1 2 3 Levey DJ, Tewksbury JJ, Izhaki I, Tsahar E, Haak DC. 2007. Evolutionary ecology of secondary compounds in ripe fruit: Case studies with capsaicin and emodin. Wallingford UK: CABI.
↑ Smith, R. L. Ecology and Field Biology. 5th ed. New York: Harper Collins, 1996.
1 2 Barnea, A., J. B. Harborne, and C. Pannell. 1993. What parts of fleshy fruits contain secondary compounds toxic to birds and why? Biochemical Systematics and Ecology. 21 (4): 421-429.
↑ Loiselle, B. A. and J. G. Blake. 2002. Potential consequences of extinction of frugivorous birds for shrubs of a tropical wet forest, in Levey, D. J., Silva, W. R., and M. Galetti, editors. Seed Dispersal and Frugivory: Ecology, Evolution and Conservation. CABI Publishing, Oxford, UK.
↑ Motta-Junior, J. C. and K. Martins. The Frugivorous Diet of the Maned Wolf, Chrysocyon brachyurus in Brazil: Ecology and Conservation, in D. J. Levey, W. R. Silva, and M. Galetti, editors. Seed Dispersal and Frugivory: Ecology, Evolution, and Conservation. CABI Publishing, Oxford, UK.
↑ http://pin.primate.wisc.edu/factsheets/entry/owl_monkey
↑ Cawthon Lang KA. 2005 July 18. Primate Factsheets: Owl monkey (Aotus) Taxonomy, Morphology, & Ecology. <http://pin.primate.wisc.edu/factsheets/entry/owl_monkey>. Accessed 2008 October 28.
↑ Wutherich, D., A. Azocar, C. Garcia-Nunez, and J. F. Silva. 2001. Seed dispersal in Palicourea rigida, a common treelet species from Neotropical savannas. Journal of Tropical Ecology 17:449-458.
1 2 Jordano, P., C. Garcia, J. A. Godoy, and J. L. Garcia-Castano. 2007. Differential contribution of frugivores to complex seed dispersal patterns. Proceedings of the National Academy of Sciences of the United States of America 104:3278-3282.
↑ Montoya, D., M. A. Zavala, M. A. Rodriguez, and D. W. Purves. 2008. Animal versus wind dispersal and the robustness of tree species to deforestation. Science 320:1502-1504.
↑ Buckley, Y. M., S. Anderson, C. P. Catterall, R. T. Corlett, T. Engel, C. R. Gosper, R. Nathan, D. M. Richardson, M. Setter, O. Spiegel, G. Vivian-Smith, F. A. Voigt, J. E. S. Weir, and D. A. Westcott. 2006. Management of plant invasions mediated by frugivore interactions. Journal of Applied Ecology 43:848-857.