Myrmecophily (/mɜːrməˈkɒfəli/ mur-mə-KOF-ə-lee, lit.

 'love of ants') is the term applied to

positive interspecies associations between ants and a variety of other organisms, such as
plants, other arthropods, and fungi. Myrmecophily refers to mutualistic associations with ants,
though in its more general use, the term may also refer to commensal or even parasitic
interactions.

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
ecological within the nests, allowing for more diversity and population sizes among the
myrmecophiles.[6][7]

Contents
 1 Myrmecophile
 2 Ant-plant interactions
 3 Ant-arthropod interactions
 4 Ant-insect interactions

o 4.1 Hemiptera
o 4.2 Lycaenid butterflies
o 4.3 Rove beetles
o 4.4 Multiple levels of myrmecophily

 5 Significance of myrmecophily in ecology

o 5.1 Evolution of positive interactions

o 5.2 Species coexistence
o 5.3 Community structure
o 5.4 Myrmecophily as a model system

 6 See also
 7 References

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 Coccinellidae (e.g. the ladybird Thalassa
saginata), Aphodiidae, Scarabaeidae, Lucanidae, Cholevidae, Pselaphidae, Staphylinidae,
Histeridae, and Ptiliidae (some treated here as subfamilies). In ant-beetle associations, the
myrmecophilous staphylinids are the most diverse of the beetle families.[10] Myrmecophilous
associations are also seen in various other insects, such as aphids and treehoppers, as well as
the hoverfly genus Microdon and several other groups of flies.[11]

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
See also: Nectar and Ant garden

Ant-plant interactions are geographically widespread,[16] with hundreds of species of

myrmecophytic plants in several families, including the Leguminosae, Euphorbiaceae, and
Orchidaceae.[3] 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 myrmecochory), reduced competition from other plants, hygienic services, and/or
nutrient supplementation.[1][17]

Three of the most common and important structural adaptations of ant plants are extrafloral
nectaries, domatia, and (least commonly) Beltian bodies. Plant domatia are formed nesting
sites provided by the plant in the form of hollow stems, petioles, thorns, or curled leaves.[17]
The production of ant-specialized domatia has been documented in over 100 genera of
tropical plants.[17] Beltian bodies provide a high-energy food source to ants in the form of
nutritive corpuscles produced on leaflet tips,[1] and they have been described in at least 20
plant families.[17] 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.[17] 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.[17] 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.

In exchange for nesting sites and food resources, ants protect plants from herbivores. One of
the best-known examples of ant-plant mutualism is in bullhorn acacias (Acacia cornigera)
and their tending Pseudomyrmex ants in Central America.[3][16] This system was studied by
Daniel Janzen in the late 1960s, who provided some of the first experimental evidence that
ants significantly reduce herbivory rates of myrmecophytes.[18][19] Since then many other
studies have demonstrated similar results in other systems.[3][17] In the bullhorn acacia system,
in exchange for protection, the acacias provide domatia, Beltian bodies, and EFNs, and
evidence indicates that the Pseudomyrmex ants can survive exclusively on these food
resources without having to forage elsewhere.[1] For many plants, including the bullhorn
acacias, ants can significantly reduce herbivory from both phytophagous insects and larger
organisms, such as large grazing mammals.[20] Obligately associated ant species are 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.[3]

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 sufficiently high to offsets benefits.[20]