Myrmecophily
Myrmecophily (
The term "myrmecophile" is used mainly for animals that associate with ants. An estimated 10,000 species of ants (Formicidae) are known, with a higher diversity in the tropics.[1] In most terrestrial ecosystems, ants are ecologically and numerically dominant, being the main invertebrate predators. As a result, ants play a key role in controlling arthropod richness, abundance, and community structure.[2] Some evidence shows that the evolution of myrmecophilous interactions has contributed to the abundance and ecological success of ants,[1][3] by ensuring a dependable and energy-rich food supply, thus providing a competitive advantage for ants over other invertebrate predators.[4] Most myrmecophilous associations are opportunistic, unspecialized, and facultative (meaning both species are capable of surviving without the interaction), though obligate mutualisms (those in which one or both species are dependent on the interaction for survival) have also been observed for many species.[5]
As ant nests grow, they are more likely to house more and greater varieties of myrmecophiles. This is partly because larger colonies have greater specializations, so more diversity of ecology within the nests, allowing for more diversity and population sizes among the myrmecophiles.[6][7]
Myrmecophile
A "myrmecophile" is an organism that lives in association with ants.
Myrmecophiles may have various roles in their host ant colony. Many consume waste materials in the nests, such as dead ants, dead larvae, or fungi growing in the nest. Some myrmecophiles, however, feed on the stored food supplies of ants, and a few are predatory on ant eggs, larvae, or pupae. Others benefit the ants by providing a food source for them. Most associations are facultative, benefiting one or both participants, but not being necessary to their survival, but many myrmecophilous relationships are obligate, meaning one or the other participant requires the relationship for survival.
Myrmecophilous associations are best known in butterflies of the family Lycaenidae. Many lycaenid caterpillars produce nectar by specialized organs, and communicate with the ants through sound and vibrations.[8] The association with ants is believed to reduce the parasitisation of the butterfly caterpillars.[9]
Some myrmecophilous beetles are in the families
Ant nests provide environmentally stable environments that are well organized and protected by the host colony. The benefit of ant colonies has resulted in infiltration from a variety of myrmecophiles.[12] The ant guests can have a positive, neutral, or negative effect on the colony. If the infiltrating species' impact is too negative on the colony, they risk discovery; this usually results in relatively small populations of myrmecophiles. Some spider species will use traits such as myrmecomorphy – ant mimicry - and chemical mimicry to infiltrate ant nests, usually to prey on food supplies or the ants themselves.[13] Aribates javensis, a species of oribatid mites, is an obligate myrmecophile that lives in ant nests. These mites are cared for by their ant hosts in exchange for eating litter and bacteria in the nest.[14]
Other myrmecophile groups include:
- Orthoptera, such as the cricket Myrmecophilus acervorum
- Diptera, such as the stratiomyid fly Clitellaria obtusa
- Blattodea, such as the Attaphila cockroaches
- Molluscs, such as Allopeas myrmekophilos[15]
The first major work in cataloguing British myrmecophiles was done by Horace Donisthorpe in his 1927 book The Guests of British Ants.
Ant-plant interactions
Ant-plant interactions are geographically widespread,
Three of the most common and important structural adaptations of ant plants are
In exchange for nesting sites and food resources, ants protect plants from
Myrmecophily is considered a form of indirect plant defense against herbivory, though ants often provide other services in addition to protection. Some ants provide hygienic services to keep leaf surfaces clean and deter disease, and defense against fungal pathogens has also been demonstrated.[17] Ants commonly prune epiphytes, vines, and parasitic plants from their host plant, and they sometimes thin the shoots of neighboring plants, as well. In doing so, ants reduce plant-plant competition for space, light, nutrients, and water.[1] Finally, current work focusing on ants' role in nutrient supplementation for plants has shown that in many ant-plant relationships, nutrient flow is bidirectional. One study has estimated that while 80% of the carbon in the bodies of Azteca spp. workers is supplied by the host tree (Cecropia spp.), 90% of the Cecropia tree's nitrogen was supplied by ant debris carried to the tree as a result of external foraging.[21] In light of these services, myrmecophily has been considered advantageous in ensuring a plant's survival and ecological success,[17] although the costs to the plant of providing for the ants can be sufficiently high to offsets benefits.[20]
Ant-arthropod interactions
Many species of arthropods are dependent on ant species and live amongst them in their nests. Mites are particularly adept at being myrmecophiles, being that they are small enough to enter nests easily and to not be evicted from the homes and bodies of ants.[6] In fact, multiple studies show mites exhibit extreme myrmecophily to numbers far above other myrmecophiles.[22][6]
Ant-insect interactions
Ants tend a wide variety of insect species, most notably lycaenid butterfly caterpillars and hemipterans.[5] About 41% of all ant genera include species that associate with insects.[23] These types of ant-insect interactions involve the ant providing some service in exchange for nutrients in the form of honeydew, a sugary fluid excreted by many phytophagous insects. .[5] Interactions between honeydew-producing insects and ants is often called trophobiosis, a term which merges notions of trophic relationships with symbioses between ants and insects. This term has been criticized, however, on the basis that myrmecophilous interactions are often more complex than simple trophic interactions, and the use of symbiosis is inappropriate for describing interactions among free-living organisms.[5]
Insects may also form adaptations to contend with ant aggression, resulting in either mutualistic or parasitic bonds with ant colonies. Some beetles from the family Coccinellidae have developed behaviors, body shapes, and chemical mimicry to prey on ant-tended aphids.[24]
Hemiptera
Some of the best-studied myrmecophilous interactions involve ants and hemipterans, especially
Ants engage in associations with other honeydew-producing hemipterans, such as scale insects (Coccidae), mealybugs (Pseudococcidae), and treehoppers (Membracidae), and most of these interaction are facultative and opportunistic with some cases of obligate associations, such as hemipterans that are inquiline, meaning they can only survive inside ant nests.[5] In addition to protection, ants may provide other services in exchange for hemipteran honeydew. Some ants bring hemipteran larvae into the ant nests and rear them along with their own ant brood.[3] Additionally, ants may actively aid in hemipteran dispersal; queen ants have been observed transporting aphids during their dispersive flights to establish a new colony, and worker ants often carry aphids to a new nesting site if the previous ant nest has been disturbed. Ants may also carry hemipterans to different parts of a plant or to different plants to ensure a fresh food source and/or adequate protection for the herd.
Lycaenid butterflies
Myrmecophily among lycaenid caterpillars differs from the associations of hemipterans because caterpillars feed on plant tissues, not phloem sap, and therefore do not continually excrete honeydew. Caterpillars of lycaenid butterflies have therefore evolved specialized organs that secrete chemicals to feed and appease ants.
Rove beetles
Multiple levels of myrmecophily
Many trophobiotic ants can simultaneously maintain associations with multiple species.
Significance in ecology
Mutualisms are geographically ubiquitous, found in all organismic kingdoms, and play a major role in all ecosystems.[33][34] Combined with the fact that ants are one of the most dominant lifeforms on earth,[16] myrmecophily clearly plays a significant role in the evolution and ecology of diverse organisms, and in the community structure of many terrestrial ecosystems.
Evolution of positive interactions
Questions of how and why species coevolve are of great interest and significance. In many myrmecophilous organisms, ant associations have been influential in the ecological success, diversity, and persistence of species. Analyses of phylogenetic information for myrmecophilous organisms, as well as ant lineages, have demonstrated that myrmecophily has arisen independently in most groups several times. Because multiple gains (and perhaps losses) of myrmecophilous adaptations have happened, the evolutionary sequence of events in most lineages is unknown.[30] Exactly how these associations evolve also remains unclear.
In studying the coevolution of myrmecophilous organisms, many researchers have addressed the relative costs and benefits of mutualistic interactions, which can vary drastically according to local
Species coexistence
In addition to leading to coevolution, mutualisms also play an important role in structuring communities.[33] One of the most obvious ways in which myrmecophily influences community structure is by allowing for the coexistence of species that might otherwise be antagonists or competitors. For many myrmecophiles, engaging in ant associations is first and foremost a method of avoiding predation by ants. For example, the caterpillars of lycaenid butterflies are an ideal source of food for ants: they are slow-moving, soft-bodied, and highly nutritious, yet they have evolved complex structures to not only appease ant aggression, but also to elicit protective services from the ants.[2] To explain why ants cooperate with other species as opposed to preying on them, two related hypotheses have been proposed; cooperation either provides ants with resources that are otherwise difficult to find, or it ensures the long-term availability of those resources.[5]
Community structure
At both small and large spatiotemporal scales, mutualistic interactions influence patterns of species richness, distribution, and abundance.[35] Myrmecophilous interactions play an important role in determining community structure by influencing inter- and intraspecific competition; regulating population densities of arthropods, fungi, and plants; determining arthropod species assemblages; and influencing trophic dynamics.[5] Recent work in tropical forests has shown that ant mutualisms may play key roles in structuring food webs, as ants can control entire communities of arthropods in forest canopies.[17] Myrmecophily has also been key in the ecological success of ants. Ant biomass and abundance in many ecosystems exceeds that of their potential prey, suggesting a strong role of myrmecophily in supporting larger populations of ants than would otherwise be possible.[17] Furthermore, by providing associational refugia and habitat amelioration for many species, ants are considered dominant ecosystem engineers.[3][35]
Model system
Myrmecophilous interactions provide an important model system for exploring ecological and evolutionary questions regarding coevolution, plant defense theory, food web structure, species coexistence, and evolutionarily stable strategies. Because many myrmecophilous relationships are easily manipulated and tractable, they allow for testing and experimentation that may not be possible in other interactions. Therefore, they provide ideal model systems in which to explore the magnitude, dynamics, and frequency of mutualism in nature.[17]
See also
- Myrmecochory
- Myrmecophyte
- Myrmecotrophy
- Myrmecomorphy (ant mimicry)
References
- ^ a b c d e f g B. Holldobler and E.O. Wilson, The Ants, Cambridge, Massachusetts: The Belknap Press of Harvard University Press, 1990.
- ^ a b c d K. Fiedler, B. Holldobler, and P. Seufert, "Butterflies and ants: The communicative domain," Cellular and molecular life sciences, vol. 52, 1996, pp. 14-24.
- ^ a b c d e f g h i j k l m B. Holldobler and E.O. Wilson, Journey to the Ants, Cambridge, Massachusetts: The Belknap Press of Harvard University Press, 1994.
- ^ N. Bluthgen, N.E. Stork, and K. Fiedler, "Bottom-up control and co-occurrence in complex communities: honeydew and nectar determine a rainforest ant mosaic," Oikos, vol. 106, 2004, pp. 344-358.
- ^ a b c d e f g h i B. Stadler and T. Dixon, Mutualism: Ants and their insect partners, Cambridge: Cambridge University Press, 2008.
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- ^ Pierce NE, Braby MF, Heath A, Lohman DJ, Mathew J, Rand DB, Travassos MA. 2002. The ecology and evolution of ant association in the Lycaenidae (Lepidoptera.) Annual Review of Entomology 47: 733-771. PDF
- ^ H. T. Baumgarten & K. Fiedler (1998). "Parasitoids of lycaenid butterfly caterpillars: different patterns in resource use and their impact on the hosts' symbiosis with ants". Zoologischer Anzeiger. 236: 167–180.
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- ^ D. Janzen, "Coevolution of mutualism between ants and acacias in Central America," Evolution, vol. 20, 1966, pp. 249-275.
- ^ D. Janzen, "Interaction of the bull's-horn acacia (Acacia cornigera L. ) with an ant inhabitant (Pseudomyrmex ferruginea F. Smith) in Eastern Mexico," Univ. Kansas Sci. Bull. , vol. 47, 1967, pp. 315-558.
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- ^ a b c K. Fiedler, Systematic, evolutionary, and ecological implication of myrmecophily within the Lycaenidae, UND Museum Alexander Koenig: Bonner Zoologische Monographien, 1991.
- ^ A.M. Fraser, A.H. Axen, and N.E. Pierce, "Assessing the Quality of Different Ant Species as Partners of a Myrmecophilous Butterfly," Oecologia, vol. 129, Nov. 2001, pp. 452-460.
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- ^ a b M. Doebeli and N. Knowlton, "The evolution of interspecific mutualisms," Proceedings of the National Academy of Sciences of the United States of America, vol. 95, 1998, pp. 8676-8680.
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