Symphiles are insects or other organisms which live as welcome guests in the nest of a social insect (such as the ant, myrmecophily, or termite, termitophily) by which they are fed and guarded. The relationship between the symphile and host may be symbiotic, inquiline or parasitic. [1]
This is a selection of taxa exhibiting symphilia, not a complete list.
Fibularhizoctonia , sometimes referred to as cuckoo fungus due to their adaptation to mimic termite eggs, employ chemical and morphological mimicry to benefit from the defense termites provide their brood. If termite workers are present to care for a brood which contains cuckoo fungus, the sclerotia, or "termite balls", are unlikely to germinate and their presence will increase the survival rate of the termite eggs. When worker termites were experimentally removed from brood that contained slerotia, the fungus germinated by exploiting the termite eggs. This means the termitophilic relationship between termites and Fibularhizoctonia can be parasitic or mutualistic. [2]
The large blue butterfly, Phengaris arion (formerly Maculinea arion), exhibits a unique parasitic relationship with a single species of red ant, Myrmica sabuleti . [3]
Cuckoo bumblebees, members of the subgenus Psithyrus in the genus Bombus, are obligate brood parasites; they must use colonies of true bumblebees to rear their young. A Psithyrus female will kill or subdue the host colony's queen and then use pheromones and/or physical attacks to force the host colony to feed her and raise her brood. [4]
Many species of Staphylinidae (commonly known as “Rove Beetles”) have developed complex interspecies relationships with ants. Ant associations range from near free-living species which prey only on ants, to obligate inquilines of ants, which exhibit extreme morphological and chemical adaptations to the harsh environments of ant nests. Some species are fully integrated into the host colony, and are cleaned and fed by ants. Many of these, including species in tribe Clavigerini, are myrmecophagous, placating their hosts with glandular secretions while eating the brood. [5]
Staphylinidae is currently considered to be the largest family of beetles, with over 58,000 species described. As such, many myrmecophilous species are unknown. The majority of studied myrmecophilous Rove Beetles belong to the subfamily Aleocharinae, including the commonly studied genera Pella , Dinarda , Tetradonia , Ecitomorpha , Ecitophya , Atemeles , and Limechusa , and to the subfamily Pselaphinae, which includes Claviger and Adranes . There are also representatives of Scydmaenidae, which includes 117 myrmecophilous species in 20 genera [6] The Aleocharinae possess defensive glands on their abdomens, which are used in myrmecophilous species to prevent attacks by their host ant and in more extreme cases to integrate completely into the colony. Many Pselephinae species have trichomes, tufts of hairs which hold placating pheromones. Pselephines have evolved trichomes independently at least four times, most notably in all members of Clavigerini, but also in Attapsenius and Songius genera. [7]
Due to their large number and diversity, myrmecophilous Rove Beetles occupy an array of behaviors. Myrmecophilous interactions can be generalized into categories, in three of which Staphylinids can be found. The synecthrans, or “persecuted guests,” the synoeketes, or “tolerated guests,” and the symphiles, or “true guests.” [8]
Synecthran insects live on the periphery of the host colony and are not accepted into the colony. [9]
Synoeketetic insects live in close contact with their host ants but are not integrated into the colony. These species may be further categorized as neutral, mimetic, loricate, and symphiloid synoeketes.
Symphilic insects have been fully integrated into the host’s society. Symphilic species have undergone complex morphological adaptations, many gaining the appearance of their host's species. Most have developed trichomes, which secrete appeasement pheromones. The most extreme adaptations, found in members of tribe Clavigerini, include the reduction of mouthparts for trophallaxis and the fusing of many body and antennal segments. While most symphiles use antennal contact to stimulate food giving from their host, at least one member of Clavigerini, Claviger testaceus, secretes a chemical to induce regurgitation from its host ant Lasius flavus . [10] Symphiles typically take on many roles in the colony, raising young, feeding and grooming adults, and helping transport food and larvae. Many Staphylinids are capable of following ant pheromone trails, although they are not limited to following trails laid by their host ant. This allows symphiles of army ants to migrate with the colony. [11] Most species are trophallactic, being fed by other members of the colony. Almost all species have also been observed feeding on the brood, making them obligate parasites.
Once the larvae of the large blue butterfly (Phengaris arion) is brought into a Myrmica sabuleti colony, it will mimic the sounds a queen Myrmica larva would make, increasing the chances that the host ant colony will prefer to care for it over their own larvae. The caterpillar feeds on the ant grubs and is a predacious symphile.
Chemical mimicry refers to the production of one species’ chemical signals by another species. Many myrmecophilous Staphylinids have evolved chemical mimicry to deter or placate ants. For Staphylinids accepted into the host colony, chemical mimicry is used for camouflage. The majority of the chemical signals used are cuticular hydrocarbons, which are produced in the cuticle of the host ant at certain concentrations and are palpated to determine the identity of an ant. Species in close contact with their host ants are able to pick up the host’s hydrocarbons and imitate the ant’s hydrocarbon pattern, thus appearing in scent at least to be the same species as the host ant. As hydrocarbon patterns are specific to an individual colony, the rove beetles are generally restricted to one nest. The production of a new hydrocarbon pattern takes time, during which the beetle is vulnerable to detection and attack. Some species, such as Zyras comes , produce volatile pheromones as well as cuticular hydrocarbons, which may provide it more protection than contact based pheromones while traveling with its host in foraging trails. [12]
The army ants that rove beetles prey on are blind, so it is important that the rove beetles feel similar to their host species. Physical adaptation to resemble ants has evolved in rove beetles on at least twelve separate occasions. [13]
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,000 species have been classified. They are easily identified by their geniculate (elbowed) antennae and the distinctive node-like structure that forms their slender waists.
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in eusocial insects such as ants, wasps, bees, and termites.
The rove beetles are a family (Staphylinidae) of beetles, primarily distinguished by their short elytra that typically leave more than half of their abdominal segments exposed. With over 66,000 species in thousands of genera, the group is the largest family in the beetle order, and one of the largest families of organisms. It is an ancient group, with fossilized rove beetles known from the Triassic, 200 million years ago, and possibly even earlier if the genus Leehermania proves to be a member of this family. They are an ecologically and morphologically diverse group of beetles, and commonly encountered in terrestrial ecosystems.
The large blue is a species of butterfly in the family Lycaenidae. The species was first defined in 1758 and first recorded in Britain in 1795. In 1979 the species became mostly extinct in Britain but has been successfully reintroduced with new conservation methods. The species is classified as "near threatened" on the IUCN Red List of Threatened Species. Today P. arion can be found in Europe, the Caucasus, Armenia, western Siberia, Altai, north-western Kazakhstan and Sichuan.
The name army ant (or legionary ant or marabunta) is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limited area.
Phengaris alcon, the Alcon blue or Alcon large blue, is a butterfly of the family Lycaenidae and is found in Europe and across the Palearctic to Siberia and Mongolia.
Ant mimicry or myrmecomorphy is mimicry of ants by other organisms; it has evolved over 70 times. Ants are abundant all over the world, and potential predators that rely on vision to identify their prey, such as birds and wasps, normally avoid them, because they are either unpalatable or aggressive. Some arthropods mimic ants to escape predation, while some predators of ants, especially spiders, mimic them anatomically and behaviourally in aggressive mimicry. Ant mimicry has existed almost as long as ants themselves; the earliest ant mimics in the fossil record appear in the mid-Cretaceous alongside the earliest ants.
Myrmica is a genus of ants within the subfamily Myrmicinae. It is widespread throughout the temperate regions of the Holarctic and high mountains in Southeast Asia.
Myrmecophily 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.
Phengaris rebeli, common name mountain Alcon blue, is a species of butterfly in the family Lycaenidae. It was first found and described in Styria, Austria, on Mount Hochschwab around 1700. Although it was initially classified as a subspecies of P. alcon, a European researcher, Lucien A. Berger, designated it as a separate species in 1946. Genetic similarities between P. rebeli and P. alcon have led many researchers to argue that the two are the same species and differences are due to intraspecific variation.
Necrophoresis is a sanitation behavior found in social insects – such as ants, bees, wasps, and termites – in which they carry away the dead bodies of members of their colony from the nest or hive area. The term was introduced in 1958 by E.O. Wilson and his colleagues. The behaviour was however known from before with Pliny making the claim that ants were the only animals other than humans to bury their dead.
Chemical mimicry is a type of biological mimicry involving the use of chemicals to dupe an operator.
Insects have a wide variety of predators, including birds, reptiles, amphibians, mammals, carnivorous plants, and other arthropods. The great majority (80–99.99%) of individuals born do not survive to reproductive age, with perhaps 50% of this mortality rate attributed to predation. In order to deal with this ongoing escapist battle, insects have evolved a wide range of defense mechanisms. The only restraint on these adaptations is that their cost, in terms of time and energy, does not exceed the benefit that they provide to the organism. The further that a feature tips the balance towards beneficial, the more likely that selection will act upon the trait, passing it down to further generations. The opposite also holds true; defenses that are too costly will have a little chance of being passed down. Examples of defenses that have withstood the test of time include hiding, escape by flight or running, and firmly holding ground to fight as well as producing chemicals and social structures that help prevent predation.
Physogastrism or physogastry is a characteristic of certain arthropods, where the abdomen is greatly enlarged and membranous. The most common examples are the "queens" of certain species of eusocial insects such as termites, bees and ants, in which the abdomen swells in order to hold enlarged ovaries, thus increasing fecundity. This means that the queen has the ability to hold more and produce more eggs at one time. Physogastric queens produce an enormous number of eggs which can account for a significant amount of their body weight. In the termite species Macrotermes subhyalinus, eggs can make up a third of their body weight, and a 15-gram queen can produce up to 30 eggs per minute. The physogastric queens' egg production is supported by oocyte proteins supplied by the "queen body fat."
Myrmica schencki is a species of ant in the genus Myrmica.
Claviger is a genus of beetles in the family Staphylinidae, subfamily Pselaphinae. About 40 species and subspecies are described, divided into two subgenera, Claviger (Claviger) and Claviger (Clavifer). Claviger displays unusual biological adaptations to myrmecophily. This pselaphid is of palearctic distribution.
This is a glossary of terms used in the descriptions of ants.
Many species of Staphylinidae have developed complex interspecies relationships with ants, known as myrmecophily. Rove beetles are among the most rich and diverse families of myrmecophilous beetles, with a wide variety of relationships with ants. Ant associations range from near free-living species which prey only on ants, to obligate inquilines of ants, which exhibit extreme morphological and chemical adaptations to the harsh environments of ant nests. Some species are fully integrated into the host colony, and are cleaned and fed by ants. Many of these, including species in tribe Clavigerini, are myrmecophagous, placating their hosts with glandular secretions while eating the brood.
Chemical communication in insects is social signalling between insects of the same or different species, using chemicals. These chemicals may be volatile, to be detected at a distance by other insects' sense of smell, or non-volatile, to be detected on an insect's cuticle by other insects' sense of taste. Many of these chemicals are pheromones, acting like hormones outside the body.
Aenictoteratini is a myrmecophilous tribe of rove beetles in the subfamily Aleocharinae which contains 10 genera, 7 of which are monotypic. In total, there are 17 species currently listed as Aenictoteratini.