399:. Plant domatia are formed nesting sites provided by the plant in the form of hollow stems, petioles, thorns, or curled leaves. The production of ant-specialized domatia has been documented in over 100 genera of tropical plants. Beltian bodies provide a high-energy food source to ants in the form of nutritive corpuscles produced on leaflet tips, and they have been described in at least 20 plant families. Extrafloral nectaries (EFNs) are known to occur in at least 66 families of angiosperm plants in both temperate and tropical regions, as well as some ferns, but are absent in all gymnosperms and are most abundant in the tropics. EFNs being outside of the plant flowers are not employed in pollination; their primary purpose is to attract and sustain tending ants. Many plants can control the flow of nectar from the EFNs so that the availability of nectar varies according to daily and seasonal cycles. Because ants can respond quickly to changes in flow rate from EFNs, this may be possible mechanism by which plants can induce greater ant activity during times of peak herbivory, and minimize overall costs of nectar production. The combined nutritional output of EFNs and Beltian bodies can be a significant food source for tending ants, and in some cases can provide the total nutritive needs for an ant colony.
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support the stable persistence of myrmecophily are still unknown. In many cases, variation in external factors can result in interactions that shift along a continuum of mutualism, commensalism, and even parasitism. In almost all mutualisms, the relative costs and benefits of interactions are asymmetrical; that is, one partner experiences greater benefits and/or fewer costs than the other partner. This asymmetry leads to "cheating", in which one partner evolves strategies to receive benefits without providing services in return. As with many other mutualisms, cheating has evolved in interactions between ants and their partners. For example, some lycaenid larvae are taken into ant nests, where they prey on ant brood and offer no services to the ants. Other lycaenids may parasitize ant-plant relationships by feeding on plants that are tended by ants, apparently immune to ant attack because of their own appeasing secretions. Hemipterophagous lycaenids engage in a similar form of parasitism in ant-hemipteran associations. In light of the variability in outcomes of mutualistic interactions, and also the evolution of cheating in many systems, much remains to be learned about the mechanisms that maintain mutualism as an evolutionarily stable interaction.
631:), compared to untended larvae, the percentage of larvae disappearing from plants before late final instar decreased (not statistically significantly, though) and the percentage of larvae infected by parasitoids significantly decreased (from 33% to 9%–12%). These interactions do not come without an energetic cost to the butterfly, however, and ant-tended individuals reach smaller adult sizes than untended individuals due to the costs of appeasing ants during the larval stage. Interactions with ants are not limited to the butterfly's larval stage, and in fact ants can be important partners for butterflies at all stages of their lifecycles. For example, adult females of many lycaenid butterflies, such as
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densities of arthropods, fungi, and plants; determining arthropod species assemblages; and influencing trophic dynamics. 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. 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. Furthermore, by providing associational refugia and habitat amelioration for many species, ants are considered dominant ecosystem engineers.
558:, meaning they can only survive inside ant nests. 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. 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.
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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. 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.
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ants' antennae. Some caterpillars possess specialized receptors that allow them to distinguish between ant antennation and contact from predators and parasites, and others produce acoustic signals that agitate ants, making them more active and likely better defenders of the larvae. As with hemipteran myrmecophiles, ants protect lycaenid larvae from predatory insects (including other ants) and parasitoid wasps, which lay their eggs in the bodies of many species of
Lepidoptera larvae. For example, one study conducted by
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interactions has contributed to the abundance and ecological success of ants, by ensuring a dependable and energy-rich food supply, thus providing a competitive advantage for ants over other invertebrate predators. 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.
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667:) do not possess the morphological or behavioral adaptations to switch to trophobiotic partners. Many ant mutualists can exploit these multispecies interactions to maximize the benefits of myrmecophily. For example, some plants host aphids instead of investing in EFNs, which may be more energetically costly depending on local food availability. The presence of multiple interactors can strongly influence the outcomes of myrmecophily, often in unexpected ways.
49:
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524:. Around 4000 species of aphids are described, and they are the most abundant myrmecophilous organisms in the northern temperate zones. Aphids feed on the phloem sap of plants, and as they feed, they excrete honeydew droplets from their anuses. The tending ants ingest these honeydew droplets, then return to their nest to regurgitate the fluid for their nestmates (see
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gain extra protein and ensure efficient resource extraction by maintaining honeydew flow rates that do not exceed the ants' collection capabilities. Even with some predation by ants, aphid colonies can reach larger densities with tending ants than colonies without. Ants have been observed to tend large "herds" of aphids, protecting them from predators and
492:, 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.
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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. Exactly how these associations evolve also remains unclear.
637:, preferentially oviposit on plants where ant partners are present, possibly by using ants' own chemical cues to locate sites where juvenile butterflies will likely be tended by ants. While ant attendance has been widely documented in lycaenid butterflies and to some extent in riodinid butterflies such as
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At both small and large spatiotemporal scales, mutualistic interactions influence patterns of species richness, distribution, and abundance. Myrmecophilous interactions play an important role in determining community structure by influencing inter- and intraspecific competition; regulating population
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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
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Many trophobiotic ants can simultaneously maintain associations with multiple species. Ants that interact with myrmecophilous insects and myrmecophytes are highly associated; species that are adapted to interact with one of these myrmecophiles may switch among them depending on resource availability
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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. The ant guests can have a positive, neutral, or negative effect on the colony. If the infiltrating
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and abundance, variation in nutrient requirements and availability, host plant quality, presence of alternative food sources, abundance and composition of predator and parasitoid species, and abiotic conditions. Because of the large amounts of variation in some of these factors, the mechanisms that
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Mutualisms are geographically ubiquitous, found in all organismic kingdoms, and play a major role in all ecosystems. Combined with the fact that ants are one of the most dominant lifeforms on earth, myrmecophily clearly plays a significant role in the evolution and ecology of diverse organisms, and
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can secrete more honeydew droplets per hour than their body weight) and for some ants, aphids may be their only source of food. In these circumstances, ants may supplement their honeydew intake by preying on the aphids once the aphid populations have reached certain densities. In this way, ants can
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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. Ants
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ants can survive exclusively on these food resources without having to forage elsewhere. For many plants, including the bullhorn acacias, ants can significantly reduce herbivory from both phytophagous insects and larger organisms, such as large grazing mammals. Obligately associated ant species are
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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
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ants, suggesting that the association are better treated as parasitic than mutualistic. Because caterpillars do not automatically pass honeydew, they must be stimulated to secrete droplets, and do so in response to ant antennation, which is the drumming or stroking of the caterpillar's body by the
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in the late 1960s, who provided some of the first experimental evidence that ants significantly reduce herbivory rates of myrmecophytes. Since then many other studies have demonstrated similar results in other systems. In the bullhorn acacia system, in exchange for protection, the acacias provide
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in the tropics. 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. Some evidence shows that the evolution of myrmecophilous
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and hemipterans. About 41% of all ant genera include species that associate with insects. 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. . Interactions between
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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. Finally, current work focusing on ants' role in nutrient supplementation for plants has shown that in many ant-plant
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In addition to leading to coevolution, mutualisms also play an important role in structuring communities. 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
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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
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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
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tree's nitrogen was supplied by ant debris carried to the tree as a result of external foraging. In light of these services, myrmecophily has been considered advantageous in ensuring a plant's survival and ecological success, although the costs to the plant of providing for the ants can be
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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.
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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.
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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. In fact, multiple studies show mites exhibit extreme myrmecophily to numbers far above other myrmecophiles.
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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.
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caterpillars are thought to be more than merely providing nutrition, with components that cause behavior alteration in the ants, with a reduction in the locomotory activity of caterpillar attendants, increased aggression and protectiveness by
528:). Between 90 and 95% of the dry weight of aphid honeydew is various sugars, while the remaining matter includes vitamins, minerals, and amino acids. Aphid honeydew can provide an abundant food source for ants (aphids in the genus
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G. Alvarez, I. Armbrecht, E. Jimenez, H. Armbrecht, and P. Ulloa-Chacon, "Ant-plant
Association in Two Tococa Species From a Primary Rain Forest of Colombian Choco (Hymenoptera: Formicidae)," Sociobiology, vol. 38, 2001, pp.
191:. Many lycaenid caterpillars produce nectar by specialized organs, and communicate with the ants through sound and vibrations. The association with ants is believed to reduce the parasitisation of the butterfly caterpillars.
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and quality. Of the ant genera that include species that associate with ant plants, 94% also include species that associate with trophobionts. In contrast, ants that are adapted to cultivate fungus (leaf cutter ants, tribe
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N.E. Pierce, M.F. Braby, A. Heath, D.J. Lohman, J. Mathew, D.B. Rand, and M.A. Travassos, "The ecology and evolution of ant association in the
Lycaenidae (Lepidoptera)," Annual Review of Entomology, vol. 47, 2002, pp.
539:. Aphid species that are associated with ants often have reduced structural and behavioral defense mechanisms, and are less able to defend themselves from attack than aphid species that are not associated with ants.
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A.A. Agrawal and J.A. Fordyce, "Induced indirect defence in a lycaenid-ant association: the regulation of a resource in a mutualism ," Proceedings of the Royal
Society of London, vol. 267, 2000, pp. 1857-1861.
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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.
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M. Heil and D. McKey, "Protective ant-plant interactions as model systems in ecological and evolutionary research," Annual Review of
Ecology, Evolution, and Systematics, vol. 34, 2003, pp. 425-453.
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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
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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.
1412:"Taste-enhancing Effects of Glycine on the Sweetness of Glucosea Gustatory Aspect of Symbiosis between the Ant, Camponotus japonicus, and the Larvae of the Lycaenid Butterfly, Niphanda fusca"
277:, 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.
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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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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
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T.H. Oliver, S.R. Leather, and J.M. Cook, "Macroevolutionary patterns in the origin of mutualisms involving ants," Journal of
Evolutionary Biology, vol. 21, Nov. 2008, pp. 1597-1608.
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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".
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feed and appease ants. The secretions are a mixture of sugar and amino acids, which in synergy is more attractive to the ants than either component in its own. The secretions of
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some of most aggressive ants in the world, and can defend a plant against herbivory by large mammals by repeatedly biting their attacker and spraying formic acid into the wound.
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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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K. Fiedler, Systematic, evolutionary, and ecological implication of myrmecophily within the Lycaenidae, UND Museum Alexander Koenig: Bonner Zoologische Monographien, 1991.
380:. In general, myrmecophytes (or ant plants) usually provide some form of shelter and food in exchange for ant "tending", which may include protection, seed dispersal (see
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J. Hoeksema and E. Bruna, "Pursuing the big questions about interspecific mutualism: a review of theoretical approaches," Oecologia, vol. 125, 2000, pp. 321-330.
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necessary to their survival, but many myrmecophilous relationships are obligate, meaning one or the other participant requires the relationship for survival.
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Stanton, M. L. and T. M. Palmer (2011). "The high cost of mutualism: effects of four species of East African ant symbionts on their myrmecophyte host tree".
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K. Fiedler, B. Holldobler, and P. Seufert, "Butterflies and ants: The communicative domain," Cellular and molecular life sciences, vol. 52, 1996, pp. 14-24.
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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
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Ito, Fuminori (September 2013). "Evaluation of the benefits of a myrmecophilous oribatid mite, Aribates javensis, to a myrmicine ant, Myrmecina sp".
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Myrmecophiles may have various roles in their host ant colony. Many consume waste materials in the nests, such as dead ants, dead larvae, or
554:), and most of these interaction are facultative and opportunistic with some cases of obligate associations, such as hemipterans that are
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Ant-plant interactions are geographically widespread, with hundreds of species of myrmecophytic plants in several families, including the
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V. Rico-Gray and P. Oliveira, The Ecology and Evolution of Ant-Plant Interactions, Chicago and London: University of Chicago Press, 2007.
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J.J. Stachowicz, "Mutualism, Facilitation, and the Structure of Ecological Communities," BioScience, vol. 51, Mar. 2001, pp. 235-246.
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B. Holldobler and E.O. Wilson, Journey to the Ants, Cambridge, Massachusetts: The Belknap Press of Harvard University Press, 1994.
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D. Janzen, "Coevolution of mutualism between ants and acacias in Central America," Evolution, vol. 20, 1966, pp. 249-275.
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are the most diverse of the beetle families. Myrmecophilous associations are also seen in various other insects, such as
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B. Holldobler and E.O. Wilson, The Ants, Cambridge, Massachusetts: The Belknap Press of Harvard University Press, 1990.
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relationships, nutrient flow is bidirectional. One study has estimated that while 80% of the carbon in the bodies of
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B. Stadler and T. Dixon, Mutualism: Ants and their insect partners, Cambridge: Cambridge University Press, 2008.
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1548:. Kitching, R. L. (Roger Laurence), 1945-, CSIRO (Australia). Collingwood, VIC, Australia: CSIRO Pub. 1999.
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Rettenmeyer, Carl W. (1962). "Notes on Host Specificity and Behavior of Myrmecophilous Macrochelid Mites".
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1081:"Spider-Ant Associations: An Updated Review of Myrmecomorphy, Myrmecophily, and Myrmecophagy in Spiders"
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Lapeva-Gjonova, Albena (April 2013). “Ant-Associated Beetle Fauna in Bulgaria: A Review and New Data”.
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1167:(Gastropoda, Pulmonata), the first myrmecophilous mollusc living in colonies of the ponerine army ant
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and a variety of other organisms, such as plants, other arthropods, and fungi. Myrmecophily refers to
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1593:"Biology of Immature Eurybia elvina (Lepidoptera: Riodinidae), a Myrmecophilous Metalmark Butterfly"
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Meigen, with a phylogenetic review of the myrmecophila species-group [Diptera, Milichiidae]"
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D. Janzen, "Interaction of the bull's-horn acacia (Acacia cornigera L. ) with an ant inhabitant (
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Ants engage in associations with other honeydew-producing hemipterans, such as scale insects (
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Some of the best-studied myrmecophilous interactions involve ants and hemipterans, especially
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Many species of arthropods are dependent on ant species and live amongst them in their nests.
384:), reduced competition from other plants, hygienic services, and/or nutrient supplementation.
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KISTNER, DAVID H. (1979), "Social and Evolutionary Significance of Social Insect Symbionts",
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643:, many other lepidopteran species are known to associate with ants, including many moths.
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Wada, Ayako; Isobe, Yu; Yamaguchi, Susumu; Yamaoka, Ryohei; Ozaki, Mamiko (2001-10-01).
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F. Smith) in Eastern Mexico," Univ. Kansas Sci. Bull. , vol. 47, 1967, pp. 315-558.
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associations with ants, though in its more general use, the term may also refer to
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to infiltrate ant nests, usually to prey on food supplies or the ants themselves.
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406:. One of the best-known examples of ant-plant mutualism is in bullhorn acacias (
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Three of the most common and important structural adaptations of ant plants are
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Vantaux, Amélie; Roux, Olivier; Magro, Alexandra; Orivel, Jérôme (2012-01-27).
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In exchange for nesting sites and food resources, ants protect plants from
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Horvitz, Carol C.; Turnbull, Christine; Harvey, Donald J. (1987-07-01).
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Myrmecophilous associations are best known in butterflies of the family
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and colleagues in Colorado experimentally found that for the larvae of
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239:(some treated here as subfamilies). In ant-beetle associations, the
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domatia, Beltian bodies, and EFNs, and evidence indicates that the
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Positive interspecies associations between ants and other organisms
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912:. Cambridge, Mass.: Belknap Press of Harvard University Press.
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A "myrmecophile" is an organism that lives in association with
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V. Witte; R. Janssen; A. Eppenstein; U. Maschwitz (2002). "
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in the community structure of many terrestrial ecosystems.
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Hojo, Masaru K.; Pierce, Naomi E.; Tsuji, Kazuki (2015).
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Ants tend a wide variety of insect species, most notably
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1379:"Evolutionary Perspectives on Myrmecophily in Ladybirds"
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Kronauer, Daniel J. C.; Pierce, Naomi E. (2011-03-22).
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honeydew-producing insects and ants is often called
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ants in Central America. This system was studied by
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194:Some myrmecophilous beetles are in the families
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132:) is the term applied to positive interspecies
1597:Annals of the Entomological Society of America
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1328:Journal of the Kansas Entomological Society
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993:de Meijere: a junior subjective synonym of
625:that were tended by a certain ant species (
587:stimulated by the ants. Here they are on a
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977:(Cairo: Hindawi Pub. Co.) 2013: 1-14
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516:A leaf-hopper nymph tended by an ant
508:Ant obtaining honeydew from an aphid
338:The first major work in cataloguing
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280:Other myrmecophile groups include:
261:and several other groups of flies.
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1114:Experimental and Applied Acarology
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680:Evolution of positive interactions
570:Ant tending a Lycaenid caterpillar
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41:For organisms that eat ants, see
1079:Cushing, Paula E. (2012-04-22).
65:
1584:
1538:
1452:
1403:
1383:Psyche: A Journal of Entomology
1370:
1319:
1156:
1105:
1085:Psyche: A Journal of Entomology
1072:
1023:
980:
967:
714:
658:Multiple levels of myrmecophily
646:
167:
946:
901:
880:, Elsevier, pp. 339–413,
13:
1:
751:
653:Myrmecophily in Staphylinidae
1002:Tijdschrift voor Entomologie
499:
7:
723:
241:myrmecophilous staphylinids
10:
1707:
1171:(Formicidae, Ponerinae)".
908:Wilson, Edward O. (1971).
650:
462:Ant-arthropod interactions
357:
348:The Guests of British Ants
342:myrmecophiles was done by
40:
29:
1478:10.1016/j.cub.2015.07.016
1126:10.1007/s10493-013-9678-6
1049:10.1016/j.cub.2011.01.050
1016:10.1163/22119434-99900017
30:Not to be confused with
1429:10.1093/chemse/26.8.983
1266:Pseudomyrmex ferruginea
1169:Leptogenys distinguenda
671:Significance in ecology
475:Ant-insect interactions
395:, and (least commonly)
293:Myrmecophilus acervorum
1576:: CS1 maint: others (
1165:Allopeas myrmekophilos
596:
571:
517:
509:
354:Ant-plant interactions
57:
1609:10.1093/aesa/80.4.513
956:Zoologischer Anzeiger
651:Further information:
622:Glaucopsyche lygdamus
585:extrafloral nectaries
577:
569:
515:
507:
389:extrafloral nectaries
51:
910:The insect societies
562:Lycaenid butterflies
550:), and treehoppers (
412:) and their tending
269:– ant mimicry - and
56:being tended by ants
1691:Mutualism (biology)
1396:10.1155/2012/591570
1098:10.1155/2012/151989
706:Community structure
697:Species coexistence
690:species composition
1187:10.1007/PL00012646
603:Narathura japonica
597:
583:eats nectar from
572:
518:
510:
453:spp.), 90% of the
344:Horace Donisthorpe
308:Clitellaria obtusa
58:
1471:(17): 2260–2264.
1295:10.1890/10-1239.1
987:I. Brake (1999).
895:978-0-12-342201-9
628:Formica podzolica
593:yellow crazy ants
589:Philippine orchid
346:in his 1927 book
275:Aribates javensis
251:, as well as the
204:Thalassa saginata
16:(Redirected from
1698:
1661:
1658:
1649:
1646:
1637:
1634:
1621:
1620:
1588:
1582:
1581:
1575:
1567:
1542:
1536:
1532:
1523:
1520:
1514:
1511:
1500:
1497:
1491:
1490:
1480:
1456:
1450:
1449:
1431:
1407:
1401:
1400:
1398:
1374:
1368:
1365:
1352:
1351:
1323:
1317:
1313:
1307:
1306:
1289:(5): 1073–1082.
1278:
1269:
1262:
1256:
1253:
1247:
1244:
1213:
1210:
1199:
1198:
1174:Insectes Sociaux
1160:
1154:
1153:
1109:
1103:
1102:
1100:
1076:
1070:
1069:
1051:
1042:(6): R208–R209.
1027:
1021:
1020:
1018:
991:Prosaetomilichia
984:
978:
971:
965:
964:
950:
944:
938:
932:
931:
905:
899:
898:
873:
862:
859:
836:
833:
827:
824:
793:
790:
777:
774:
438:parasitic plants
409:Acacia cornigera
271:chemical mimicry
131:
128:
125:
122:
119:
112:
106:
105:
102:
101:
98:
95:
92:
89:
86:
83:
80:
77:
74:
71:
21:
1706:
1705:
1701:
1700:
1699:
1697:
1696:
1695:
1666:
1665:
1664:
1659:
1652:
1647:
1640:
1635:
1624:
1589:
1585:
1569:
1568:
1556:
1544:
1543:
1539:
1533:
1526:
1521:
1517:
1512:
1503:
1498:
1494:
1465:Current Biology
1457:
1453:
1416:Chemical Senses
1408:
1404:
1375:
1371:
1366:
1355:
1324:
1320:
1314:
1310:
1279:
1272:
1263:
1259:
1254:
1250:
1245:
1216:
1211:
1202:
1161:
1157:
1110:
1106:
1077:
1073:
1036:Current Biology
1032:"Myrmecophiles"
1028:
1024:
985:
981:
972:
968:
951:
947:
939:
935:
920:
906:
902:
896:
874:
865:
860:
839:
834:
830:
825:
796:
791:
780:
775:
758:
754:
726:
717:
708:
699:
682:
673:
660:
655:
649:
564:
502:
477:
464:
432:commonly prune
366:
356:
170:
129:
126:
123:
120:
110:
68:
64:
52:Myrmecophilous
46:
39:
28:
23:
22:
15:
12:
11:
5:
1704:
1694:
1693:
1688:
1683:
1678:
1663:
1662:
1650:
1638:
1622:
1603:(4): 513–519.
1583:
1554:
1537:
1524:
1515:
1501:
1492:
1451:
1422:(8): 983–992.
1402:
1369:
1353:
1334:(4): 358–360.
1318:
1308:
1270:
1257:
1248:
1214:
1200:
1181:(4): 301–305.
1155:
1104:
1071:
1022:
979:
966:
945:
933:
918:
900:
894:
878:Social Insects
863:
837:
828:
794:
778:
755:
753:
750:
749:
748:
742:
737:
732:
725:
722:
716:
713:
707:
704:
698:
695:
681:
678:
672:
669:
659:
656:
648:
645:
640:Eurybia elvina
591:bud with some
580:Loxura atymnus
563:
560:
548:Pseudococcidae
546:), mealybugs (
501:
498:
476:
473:
463:
460:
397:Beltian bodies
355:
352:
336:
335:
323:
316:, such as the
311:
301:, such as the
296:
287:, such as the
169:
166:
152:interactions.
26:
9:
6:
4:
3:
2:
1703:
1692:
1689:
1687:
1684:
1682:
1679:
1677:
1674:
1673:
1671:
1657:
1655:
1645:
1643:
1633:
1631:
1629:
1627:
1618:
1614:
1610:
1606:
1602:
1598:
1594:
1587:
1579:
1573:
1565:
1561:
1557:
1551:
1547:
1541:
1531:
1529:
1519:
1510:
1508:
1506:
1496:
1488:
1484:
1479:
1474:
1470:
1466:
1462:
1455:
1447:
1443:
1439:
1435:
1430:
1425:
1421:
1417:
1413:
1406:
1397:
1392:
1388:
1384:
1380:
1373:
1364:
1362:
1360:
1358:
1349:
1345:
1341:
1337:
1333:
1329:
1322:
1312:
1304:
1300:
1296:
1292:
1288:
1284:
1277:
1275:
1267:
1261:
1252:
1243:
1241:
1239:
1237:
1235:
1233:
1231:
1229:
1227:
1225:
1223:
1221:
1219:
1209:
1207:
1205:
1196:
1192:
1188:
1184:
1180:
1176:
1175:
1170:
1166:
1159:
1151:
1147:
1143:
1139:
1135:
1131:
1127:
1123:
1119:
1115:
1108:
1099:
1094:
1090:
1086:
1082:
1075:
1067:
1063:
1059:
1055:
1050:
1045:
1041:
1037:
1033:
1026:
1017:
1012:
1008:
1004:
1003:
998:
996:
992:
983:
976:
970:
962:
958:
957:
949:
943:
937:
929:
925:
921:
919:0-674-45490-1
915:
911:
904:
897:
891:
887:
883:
879:
872:
870:
868:
858:
856:
854:
852:
850:
848:
846:
844:
842:
832:
823:
821:
819:
817:
815:
813:
811:
809:
807:
805:
803:
801:
799:
789:
787:
785:
783:
773:
771:
769:
767:
765:
763:
761:
756:
747:(ant mimicry)
746:
745:Myrmecomorphy
743:
741:
740:Myrmecotrophy
738:
736:
733:
731:
728:
727:
721:
712:
703:
694:
691:
686:
677:
668:
666:
654:
644:
642:
641:
636:
635:
630:
629:
624:
623:
618:
613:
611:
605:
604:
594:
590:
586:
582:
581:
576:
568:
559:
557:
553:
549:
545:
540:
538:
533:
532:
531:Tuberolachnus
527:
523:
514:
506:
497:
493:
491:
486:
482:
472:
469:
459:
456:
452:
451:
446:
445:
439:
436:, vines, and
435:
429:
426:
421:
420:Daniel Janzen
417:
416:
411:
410:
405:
400:
398:
394:
390:
385:
383:
379:
375:
374:Euphorbiaceae
371:
365:
361:
351:
349:
345:
341:
334:
333:myrmekophilos
332:
327:
324:
321:
320:
315:
312:
310:
309:
304:
300:
297:
295:
294:
290:
286:
283:
282:
281:
278:
276:
272:
268:
267:myrmecomorphy
262:
260:
259:
254:
250:
246:
242:
238:
234:
230:
229:Staphylinidae
226:
222:
218:
214:
210:
206:
205:
201:
197:
196:Coccinellidae
192:
190:
185:
182:
177:
175:
165:
161:
158:
153:
151:
147:
143:
139:
135:
115:
114:
104:
62:
55:
50:
44:
37:
36:Myrmecophilus
33:
19:
1600:
1596:
1586:
1545:
1540:
1518:
1495:
1468:
1464:
1454:
1419:
1415:
1405:
1386:
1382:
1372:
1331:
1327:
1321:
1311:
1286:
1282:
1265:
1260:
1251:
1178:
1172:
1168:
1164:
1158:
1120:(1): 79–85.
1117:
1113:
1107:
1088:
1084:
1074:
1039:
1035:
1025:
1009:(1): 31–36.
1006:
1000:
994:
990:
982:
974:
969:
960:
954:
948:
936:
909:
903:
877:
831:
735:Myrmecophyte
730:Myrmecochory
718:
715:Model system
709:
700:
687:
683:
674:
661:
647:Rove beetles
638:
632:
626:
620:
610:Pristomyrmex
608:
601:
598:
578:
541:
529:
526:trophallaxis
519:
494:
490:trophobiosis
485:caterpillars
478:
465:
454:
448:
442:
430:
425:Pseudomyrmex
424:
415:Pseudomyrmex
413:
407:
401:
386:
382:myrmecochory
367:
347:
337:
329:
317:
306:
291:
279:
274:
263:
256:
213:Scarabaeidae
202:
193:
186:
178:
171:
168:Myrmecophile
162:
154:
134:associations
127:love of ants
61:Myrmecophily
60:
59:
43:Myrmecophagy
32:Myrmecophila
18:Myrmecophile
1681:Myrmecology
634:J. evagoras
552:Membracidae
537:parasitoids
378:Orchidaceae
370:Leguminosae
322:cockroaches
303:stratiomyid
249:treehoppers
225:Pselaphidae
142:mutualistic
1670:Categories
1555:0643050272
963:: 167–180.
752:References
483:butterfly
404:herbivores
364:Ant garden
358:See also:
328:, such as
285:Orthoptera
233:Histeridae
221:Cholevidae
209:Aphodiidae
198:(e.g. the
189:Lycaenidae
1617:0013-8746
1572:cite book
1438:0379-864X
1340:0022-8567
1134:0168-8162
1058:0960-9822
612:punctatus
556:inquiline
500:Hemiptera
434:epiphytes
319:Attaphila
314:Blattodea
237:Ptiliidae
217:Lucanidae
157:diversity
150:parasitic
146:commensal
1564:40792921
1535:733-771.
1487:26234210
1446:11595675
1348:25083278
1316:558-602.
1303:21661568
1195:27145464
1150:17193861
1142:23423426
1091:: 1–23.
1066:21419982
995:Milichia
724:See also
544:Coccidae
481:lycaenid
455:Cecropia
450:Cecropia
331:Allopeas
326:Molluscs
258:Microdon
253:hoverfly
200:ladybird
148:or even
136:between
1676:Ecology
1389:: 1–7.
1283:Ecology
393:domatia
340:British
299:Diptera
289:cricket
121:
109:mur-mə-
1615:
1562:
1552:
1485:
1444:
1436:
1346:
1338:
1301:
1193:
1148:
1140:
1132:
1064:
1056:
975:Psyche
928:199513
926:
916:
892:
665:Attini
617:Pierce
522:aphids
444:Azteca
376:, and
360:Nectar
255:genus
245:aphids
235:, and
113:-ə-lee
54:aphids
1344:JSTOR
1191:S2CID
1146:S2CID
468:Mites
181:fungi
1686:Ants
1613:ISSN
1578:link
1560:OCLC
1550:ISBN
1483:PMID
1442:PMID
1434:ISSN
1387:2012
1336:ISSN
1299:PMID
1138:PMID
1130:ISSN
1089:2012
1062:PMID
1054:ISSN
924:OCLC
914:ISBN
890:ISBN
362:and
305:fly
247:and
174:ants
138:ants
118:lit.
1605:doi
1473:doi
1424:doi
1391:doi
1291:doi
1183:doi
1122:doi
1093:doi
1044:doi
1011:doi
1007:142
961:236
942:PDF
882:doi
207:),
111:KOF
73:ɜːr
34:or
1672::
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1641:^
1625:^
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1570:{{
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1504:^
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1469:25
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1356:^
1342:.
1332:35
1330:.
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1287:92
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1273:^
1217:^
1203:^
1189:.
1179:49
1177:.
1144:.
1136:.
1128:.
1118:61
1116:.
1087:.
1083:.
1060:.
1052:.
1040:21
1038:.
1034:.
1005:.
999:.
959:.
922:.
888:,
866:^
840:^
797:^
781:^
759:^
391:,
372:,
350:.
231:,
227:,
223:,
219:,
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1475::
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1393::
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1305:.
1293::
1197:.
1185::
1152:.
1124::
1101:.
1095::
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1046::
1019:.
1013::
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930:.
884::
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130:'
124:'
103:/
100:i
97:l
94:ə
91:f
88:ɒ
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82:ˈ
79:ə
76:m
70:m
67:/
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45:.
38:.
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Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
