Myrmecophiles

Myrmecophiles may occupy a variety of ecological niches within their host ant colony. Some consume waste materials in the nests, such as dead ants, dead larvae, or fungi growing in the nest. A few feed on external secretions of ants and some are fed directly by their host ants. Some myrmecophiles feed on the stored food supplies of ants, and a few are predatory on ant eggs, larvae, pupae or even adults. Others benefit the ants by providing a food source for them. Many myrmecophilous relationships are obligate, meaning one or the other participant requires the relationship for survival. Some associations are facultative, benefiting one or both participants but not being necessary to their survival. Many myrmecophiles await discovery and for many the nature of the relationship with their host is unknown.



Ectobiidae
There are nine species of Attaphila, all are myrmecophiles in nests of leaf-cutting ants.


 * Attaphila

Brentidae

 * Systellus

Buprestidae

 * Habroloma myrmecophila

Harpalinae

 * Helluomorphoides

Paussinae
Paussinae have a predominantly pan-tropical distribution. They comprise about 800 species. Some members of which are known also as flanged bombardier beetles.


 * Arthropterus

Pseudomorphinae

 * Cryptocephalomorpha *Species of the Cryptocephalomorpha genus are exceptional because rather than lay eggs, these beetles give birth to larvae, which are more mobile in case of an attack from the ants. This appears to be one of their adaptations to life among the ants. The morphology of the larvae is also more complex than that of an egg. Larvae may have glands that can secrete scents to hide their presence or stave off the ants.

Cryptocephalinae
Myrmecophilous cryptocephalines (34 species in 14 genera) primarily live among formicine and myrmecines ants as hosts. These two ant lineages are putative sister-groups, with their root-node dated to between 77–90 mya. In the New World tropics, the relatively recent radiation of ants from moist forests to more xeric ecosystems might have propelled the association of cryptocephalines and ant nests.


 * Agrain FA, Buffington ML, Chaboo CS, Chamorro ML, Schöller M. Leaf beetles are ant-nest beetles: the curious life of the juvenile stages of case-bearers (Coleoptera, Chrysomelidae, Cryptocephalinae). Zookeys. 2015 Dec 17;(547):133-64.


 * Anomoea * larvae are myrmecophiles having an association with ants of the genus Formica.


 * Cadmus
 * Clytra
 * Clytrasoma
 * Coenobius
 * Coscinoptera
 * Cryptocephalus
 * Griburius
 * Hockingia
 * Isnus
 * Labidostomis
 * Lachnaia
 * Megalostomis
 * Pachybrachis
 * Smaragdina
 * Tituboea

Curculionidae

 * Liometophilus *Liometophilus manni Fall, this weevil is a mymecophile living in the nest of the ant Liometopum apiculatum Mayr. Rare in collections, with the type series of 2 specimens found in the runways of different colonies in Ramsey Canyon. Also found on the undersides of stones covering the entrance to the ant's nest. I have seen specimens from the Huachuca Mts., Chiricahua Mts., and Dragoon Mts. in AZ; Grant County NM; and Hidalgo Mexico, always in small numbers. "Charles W. O'Brien" 2009

Dermestidae

 * Thorictus *All species in the genus Thorictus are myrmecophilous and demonstrate two types of life history patterns with regards to their hosts. Some are Cataglyphis specialists, such as Thorictus foreli, a system that was described early on by authors like Wasmann and Forel (see Wheeler (1910) and again later by other researchers (Reichensperger, 1925, Banck, 1927). These Thorictus species are phoretic, i.e., they remain attached to ant antennae. Phoresy is the name of the association in which an organism attaches itself to a host body part and is subsequently transported by its host. Others, like Thorictus grandicollis, roam freely in the nest and are hosted by various ant species (Messor, Pheidole, etc.); they are considered to be generalists (Cammaerts and Cammaerts, 1994). All the Thorictus species, specialists and generalists, are deemed to be mainly detritivorous (Sanchez-Pinero and Gomez, 1995).

Endomychidae
Endomychidae includes a number of species closely associated with social insects, particularly ants and termites. Wasmann (1894) listed 11 endomychids associated with ants. Price and Young (2007) noted the close proximity of adults of Rhanidea unicolor to a colony of Lasius ants, although no direct association was inferred. Myrmecophily is the most common form of social insect inquilinism among endomychids; however, termitophilous and melittophilous species are also known. Endomychids have also been recovered from birds' nests.

Histeridae
Histeridae is worldwide in distribution with just under 4,300 known species, grouped into about 350 genera. It reaches its highest diversity in the tropics. Both subfamilies Chlamydopsinae, mainly distributed in southern Asia, Pacific, and Australia, and Haeteriinae contain myrmeco- or termitophilous species. It is accepted that myrmecophiles feed on the larvae of ants or other insects or even regurgitated food from the host ants (Lapeva-Gjonova, 2013).

Chlamydopsinae
With 177 species, chlamydopsines form the second significant radiation of histerid myrmecophiles, with a single species recorded in association with termites (CATERINO & DÉGALLIER 2007). The group is confined principally to the tropics of the Australasian, Indomalayan and Oceanian regions, but extends northwards into subtropical Japan. Most species appear to be morphologically specialised obligate colony guests, possessing trichomes that envelop deep clefts or depressions on the elytral humeri, or more rarely on the pronotum. Unfortunately, the biology of chlamydopsines is exceptionally poorly known, although the beetles have been observed feeding on ant larvae and being carried by their hosts, which may use the elytral clefts as grasping notches (OKE 1923). Although the majority of species have been collected using flight intercept traps and thus lack host association data (TISHECHKIN 2009), collections made directly from colonies reveal that as a group, chlamydopsines utilize diverse host ants, although ponerines appear to be the most commonly used ant subfamily (CATERINO & DÉGALLIER 2007). Chlamydopsine larvae await discovery.


 * Ceratohister * Ceratohister is here defined to include Orectoscelis-like species which have lost any indication of elytral trichomes, and which lack the elevated elytral carinae of the undescribed Fijian species and some Australian species of Pheidoliphila (e.g. P. storeyi Dégallier & Caterino, P. dahlgreni Dégallier & Caterino).

Dendrophilinae

 * Dendrophilus

Haeteriinae
Haeteriines (Fig. 2D) number 335 described species in more than 100 genera, although the true diversity is far larger (M. Caterino, pers. comm.). Monophyly of the group is highly likely, aside from a few aberrant genera of questionable placement (KOVARIK & CATERINO 2005), and the subfamily may rank alongside Paussini (Carabidae, discussed above) and Clavigeritae (Staphylinidae: Pselaphinae, discussed below) as one of the most speciose clades of beetle myrmecophiles. The group also includes scattered termitophilous species (TISHECHKIN 2005). As with paussines and clavigerites, many genera include one or just a few species – a symptom of the extreme morphological diversity of the subfamily that can obscure even close phylogenetic relationships. Haeteriines range from small species with relatively generalised morphologies, to larger-bodied and heavily armoured species with dramatic cuticular protruberances, particularly on the pronotum, most likely for withstanding grasping ant mandibles. Trichomes are present in various taxa, and some have very elongate legs to grasp or groom hosts (AKRE 1968, KISTNER 1982). Haeteriines are predominantly Neotropical, but extend into North America and three genera also occur in the Palaearctic. Individual haeteriine genera are adapted to their ant hosts in diverse ways: some are attacked or largely ignored by their hosts, while others are highly integrated guests, which may be fed trophallactically by workers (WHEELER 1908, HENDERSON & JEANNE 1990). A recent biochemical study of one of the European genera, Sternocoelis, provided evidence that the beetles can actively synthesise cuticular hydrocarbon blends to match the odour profiles of their Aphaenogaster host colonies (LENOIR & al. 2012). The greatest number of haeteriine species are army ant inquilines (HELAVA & al. 1985, TISHECHKIN 2005), with almost 30 genera associated with Eciton alone. Adults appear to be highly host-specific (TISHECHKIN 2005), and are attracted to trails of their hosts and repelled by trails of non-host species (AKRE & RETTENMEYER 1968). The beetles have been observed feeding on the ants' brood and harvested food, and to run with or phoretically attach to workers during emigrations (AKRE 1968). DNA sequencing of histerid larvae sifted from under Eciton bivouacs and refuse dumps successfully identified haeteriines (CATERINO & TISHECHKIN 2006), implying that the entire lifecycles of at least some of these army ant guests take place in or around, host colonies.


 * Aemulister

Hydrophilidae

 * Chimaerocyon

Monotomidae

 * Monotoma

Ptinidae

 * Diplocotes

Coenochilina (Subtribe)

 * Coenochilus

Cremastocheilina (Subtribe) eleven genera
hind angles, lacking trichomes. For the time being I consider Psilocnemis the closest known relative of Centrochilus. Distribution. — Nearctic: Mexico; one species known. Bionomics. — Unknown, but association with social insects suspected.
 * Centrochilus *At first sight Centrochilus seems intermediate between Cremastocheilus and Psilocnemis, the former having a sophisticated set of prothoracic trichomes, the latter having a simple prothorax with obsolete

Aphodiinae

 * Euparia

Salpingidae
Revision of Western Hemisphere Dacoderinae with a phylogenetic analysis of world taxa (Coleoptera: Salpingidae)
 * Dacoderus

Aleocharinae

 * Amidobia
 * Aphaenochara
 * Crematoxenus
 * Doryloxenus
 * Ecitocryptus


 * Dinarda

Lomechusini

 * Drusilla sparsa / Crematogaster osakensis
 * Ecitonidia


 * Leptogenopapus


 * Lomechusa


 * Zyras

Oxypodini

 * Labidopullus
 * Oxypoda
 * Thiasophila

Pselaphinae

 * Adranes

Scydmaeninae
Includes 117 myrmecophilous species in 20 genera.
 * O'Keefe, Sean (2000). "Ant-Like Stone Beetles, Ants, and Their Associations (Coleoptera: Scydmaenidae; Hymenoptera: Formicidae; Isoptera)". ‘’J. N. Y. Entomol. Soc.’’ (NYC: N. Y. Entomol. Soc.) 108 (3/4): 273–303.


 * Cephennium
 * Chevrolatia
 * Euconnus
 * Papusus
 * Plaumanniola
 * Scydmaenus

Tachyporinae

 * Vatesus

Adelostomini

 * Cimiciopsis


 * Alaudes

Tenebrioninae

 * Bycrea

Microdontinae
There are 43 genera and more than 400 species in this subfamily of Syrphidae. The Microdontinae are Myrmecophiles, meaning they live in the nests of ants. Larval Microdontinae are scavengers or predators in ant nests, and, in contrast to other syrphid larvae, have no readily apparent body segmentation. Some species also do not exhibit the typical adult flower-visiting behaviour of other hoverflies, but instead remain near their larval host colonies (some of these species have no functional mouthparts and cannot feed as adults)


 * Afromicrodon *  All five species are restricted to Madagascar and the Comoros Islands.


 * Mixogaster
 * Oligeriops
 * Omegasyrphus
 * Paragodon
 * Paramicrodon
 * Paramixogaster
 * Parocyptamus
 * Peradon
 * Piruwa
 * PseudomicrodonThere are 14 known species. Larvae are found in ant nests. Native to the New World tropics.
 * Ptilobactrum
 * Rhoga
 * Rhopalosyrphus The larvae feed as scavengers in the nests of ants, Pseudomyrmecinae. There are nine native species from the southern United States to northern Argentina.
 * Schizoceratomyia
 * Serichlamys
 * Spheginobaccha
 * Stipomorpha
 * Sulcodon
 * Surimyia
 * Thompsodon
 * Ubristes

Myrmecophilidae
There are five genera of ant-loving crickets in this family and around 100 species. They are obligate inquilines within ant nests. World-wide in distribution, many species are found with different species and genera of ant hosts. Many ant hosts are still unknown. All species are very small, yellow, brown, or nearly black in color, wingless, and flattened, and resemble small cockroach nymphs. They do not produce sound, and lack both wings and tympanal organs ("ears") on the front tibia. There are sixty-three species in this genus.


 * Myrmecophilus


 * Myrmophilellus
 * Camponophilus
 * Eremogryllodes
 * Microbothriophylax

Lycaenidae

 * Phengaris (=Maculinea) (Lycaenidae) - Sielezniew et al. (2015) - Caterpillars develop on specific host plants (depending on species: Thymus or Origanum, Gentiana and Sanguisorba) and complete their development inside the nests of specific red ants (Myrmica sp.) as social parasites feeding on the hosts’ brood, or being fed by trophallaxis (Thomas, 1995).

Hovestadt et al. (2019) used a population-genetic model to show how individual Maculinea species could theoretically switch ant hosts.


 * ant hosts
 * Myrmica schencki host of Mountain Alcon Blue (Phengaris rebeli) - fairly well studied association, see M. schencki species page
 * Myrmica lonae host of Large Blue (Phengaris arion)

Mites
There are a large number of mite species that live on and with ants.

A genus of mites that use ants for transportation (phoresy).
 * Caesarodispus

Dos Santos Lopes et al. (2015) - Most Oplitidae are myrmecophilous or termitophilous (Hunter and Farrier 1975, 1976), with adults as the most commonly encountered instar on the ants. Available data suggest that host specificity in Oplitidae is quite high, with individual oplitid species usually associated with only one, or a few closely related, host species (Hunter and Farrier 1975, 1976; Campbell et al. 2013). Oplitis is found worldwide, but current understanding of local faunas varies widely among regions. For example, while the European fauna is fairly well studied, the Neotropical fauna is still relatively poorly known (Sellnick 1926, 1954, 1963; Zirngiebl-Nicol and Hirschmann 1973a, 1973b; Hirschmann 1975, 1991; Hiramatsu and Hirschmann 1983; Kontschán and Starý 2012).
 * Oplitis

A survey of mites on ants in the genus Neoponera from the state of Bahia, northeastern Brazil, revealed a new Oplitis species of the paradoxa-Gruppe (Hirschmann 1991), interpreted here as Oplitis s.s., which will be described for the adult instars. This species is associated with three closely related Neoponera species in the Apicalis complex (Wild 2005).

Dos Santos Lopes et al. (2015) - Associated with: Neoponera verenae, Neoponera inversa and Neoponera apicalis.
 * Oplitis apicalis

A total of 291 Oplitis specimens were recovered attached to ants. Five of these specimens, recovered from three different nests of N. verenae, were identified as O. camponoti (Hirschmann 1991). Most likely this is accidental, as O. camponoti is normally associated with ants in the genus Camponotus (Hirschmann 1991).

Da Silva et al. (2017) - The mite genus Petalomium Cross 1965 (Acari: Heterostigmatina: Neopygmephoridae) includes about 40 described species, most of them associated with ants (Hymenoptera: Formicidae) (Khaustov 2015). The papers referring to Petalomium in Neotropical region are Mahunka (1981), describing P. affinitum from Santa Lucia, West Indies, and Berghoff et al. (2009), reporting 2 unidentified species from Republic of Panama. The aim of this study is to describe a new species associated with workers of the poneromorph ant Neoponera verenae from southern Bahia, Northeastern Brazil. For purposes of comparison, a closely related species, Petalomium gottrauxi Mahunka 1977 is redescribed based on the paratype and specimens from the Crimean Peninsula.
 * Petalomium

Atelurinae
Atelurinae is a subfamily of primitive insects belonging to the order Zygentoma. Once considered an independent family, it is now treated as a subfamily within the Nicoletiidae. They are generally found in association with ants or termites, living as inquilines in the hosts' nests. They are typically small, tear-drop or sub-ovoid in body shape, light yellow in color and lacking eyes. The subfamily is quite diverse, with more than 140 described species in about 70 genera; many of the genera are monotypic.

Pseudoscorpions

 * Marachernes bellus (Harvey, 1992)