Difference between revisions of "Myrmecophiles"
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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.
- 1 Blattodea
- 2 Coleoptera
- 2.1 Brentidae
- 2.2 Buprestidae
- 2.3 Carabidae
- 2.4 Chrysomelidae
- 2.5 Curculionidae
- 2.6 Dermestidae
- 2.7 Endomychidae
- 2.8 Histeridae
- 2.9 Hydrophilidae
- 2.10 Monotomidae
- 2.11 Ptinidae
- 2.12 Scarabaeidae
- 2.13 Salpingidae
- 2.14 Staphylinidae
- 2.15 Tenebrionidae
- 3 Diptera
- 4 Orthoptera
- 5 Lepidoptera / Butterflies
- 6 Mites
- 7 Collembola
- 8 Zygentoma
- 9 Pseudoscorpions
- 10 References
There are nine species of Attaphila, all are myrmecophiles in nests of leaf-cutting ants.
Paussinae have a predominantly pan-tropical distribution. They comprise about 800 species. Some members of which are known also as flanged bombardier beetles.
There are thirty-two species in this genus. All are assumed to be myrmecophiles.
The genera of the Paussidae of the Baltic amber. Zool Anz 68(1/2): 25-30. A single species, Eopaussus balticus from Baltic Amber.
There are seven species in this myrmecophilous genus of Carabidae.
There are 26 species in this myrmecophilous genus
A myrmecophilous genus of 12 species from South and Central America.
The genus Lebioderus Westwood, 1938, belongs to the subtribe Platyrhopalina Jeannel, 1946, of the tribe Paussini Latreille, 1807, and is represented by nine species from Southeast Asia, including Indonesia [Jawa (Java), Sumatera (Sumatra), and Kalimantan], Malaysia (Peninsular Malaysia), and the Philippines (Luzon) (Luna de Carvalho, 1987).
There are two extinct species in this genus from the Lower Oligocene, Florissant, Colorado.
A most species-rich genus with more than 368 described species, all are assumed to have an obligatory symbiosis with ants. In this symbiosis the beetles provide rewarding chemical secretions to their host ants and in return receive protection, a safe place for their vulnerable larvae to develop and a reliable source of protein-rich food: the ants, particularly the brood.
There are sixteen species in this genus.
- 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.
- Guyanemorpha *A newly discovered beetle species from the lowland rain forests of French Guiana coexists with ants. The colorful insect belongs to the Pseudomorphini tribe, a family of beetles known to live among ant species. Called Guyanemorpha spectabilis, the beetle’s large size and rich colors make it stand out in comparison with its dully colored relatives in the Western Hemisphere.
- Pseudomorpha * Pseudomorpha is a myrmecophilous genus of Carabid beetle restricted to the Western Hemisphere. Pseudomorpha and some related genera are obligatory myrmecophiles in their larval stages. There are 27 described species with more than 100 undescribed species. As far as is known, adults are found in and around ant nests and in the surrounding vicinity; females are ovoviviparous (Liebherr and Kavanaugh 1985); larvae are known to be ant nest inquilines (Lenko 1972; Erwin 1981), or perhaps living with termites (Ogueta 1967).
- Samiriamorpha *Samiriamorpha have been found in the arboreal nests of Azteca ants (Erwin and Geraci 2008).
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.
- 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
- 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 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 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).
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).
- Chlamydonia *The species of Chlamydonia all possess frontal tubercles, in two parallel, longitudinal series, median and lateral paired protuberances at the anterior pronotal margin, epipleural marginal stria doubled above the metathoracic leg, and humeral trichomes that are more or less elongate in orientation.
- Chlamydopsis * Species of Chlamydopsis are very diverse in morphology, and cannot be defined at present by any definite synapomorphies (Caterino 2003). The species currently contained in Chlamydopsis all share a visible scutellum and an upturned anterior pronotal margin, at least above the antennal cavities, if not along the entire margin. None are entirely flattened dorsally (like Ectatommiphila) and none have the strikingly elongate trichome setae seen in Eucurtia.
- Ectatommiphila *There are two Australian species in this myrmecophilous genus of Histeridae. Ectatommiphila is easily recognized by its large size (c. 4 mm, and flat dorsum. Its small trichomes open as transverse slits which are not raised above the level of the elytra.
- Eucurtia * The lone valid species Eucurtia comata (Blackburn) is recognisable principally by its large size and by the dense, elongate tufts of setae arising from the anterior and posterior elevations of the elytral trichomes
- Eucurtiopsis * Elytra broader than prothorax; trichome transversely incised; pronotum of most species with paired subapical dorsal processes; most species with branched setae on some part of the body.
- Gomyopsis * The pronotal trichomes of Gomyopsis are unknown in the subfamily outside of Chlamydopsis. The sole species in this genus is similar in general body shape to Eucurtiopsis brendelli, from Sulawesi, with which it also shares a margined pronotum. It is not clear, however, if these features are indicative of a close relationships between the two.
- Kanakopsis * This distinctive chlamydopsine resembles members of the Orectoscelis lineage. However, it is distinguished by the combination of a distinct prementum, 3-segmented labial palpi, lack of an accessory epipleural stria, and lack of any frontal or anterior pronotal processes.
- Orectoscelis * Orectoscelis is herein restricted to those species with a completely hidden scutellum and circular, usually obliquely elevated humeral trichomes, fringed by a single continuous setal margin.
- Papuopsis * Papuopsis is distinguished from other genera by its laterally margined pronotum and elytra, lack of pronotal tubercles, short prothorax, and broadly rounded prosternal keel which extends nearly to the metasternum.
- Pheidoliphila *There are twenty-six species in this myrmecophilous genus of Histeridae. Lea originally characterised this genus as follows: ‘The species of this genus are all small. The prothorax is strongly raised in front, with the raised portions overhanging the head. The elytra are not depressed immediately begind the scutellar region, but there is a depression close to each shoulder; there is an oblique incision at each shoulder, but the shoulders themselves are not raised above the general level, although clothed internally. All the legs are fitted into grooves.’We have conceived of this group more broadly (Dégallier and Caterino 2005b), in particular including a few species that lack humeral trichomes (P. secqi, P. penatii, and P. oharai). All may still be characterised by the presence of some sort of bifid projection, usually prominent, extending dorsad, and often anterad of anterior pronotal margin. Where trichomes are present, they are small, oblique, and found in the extreme anterolateral corners of each elytron. Though hosts are now known for only a few species, all records are with ants in the genus Pheidole. Oke (1923) reported P. minuta being carried around by its hosts, held in the ants’ mandibles apparently by the humeral trichome. Pheidoliphila have been found throughout eastern Australia, as well as in Tasmania, and a single species placed here (P. micra) occurs in Papua New Guinea.
- Quasimodopsis * This taxon is very distinctive and phylogenetically isolated among the Chlamydopsinae. It is best defined and diagnosed based on the form of the prothorax which is very different from that of any other member of the subfamily. Although in gross terms it shares the hidden scutellum with Orectoscelis, Pheidoliphila, and others, the manner in which the scutellum is concealed, by the posteriorly prolonged pronotal margin, is very different.
- Teretriopsis * Because this genus is known from a single species, it is impossible to confidently determine characters by which it will consistently differ from other genera. It shares with Ceratohisterthe (apomorphic) lack of humeral trichomes, but differs from all known Ceratohister quite radically in body shape, mainly in its subcylindrical habitus. Its lack of pronotal processes also separates it from most Ceratohister (except for C. leai, which occurs in Sarawak, Bornean Malaysia).
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.
- Aritaerius *A myrmecophilous genus found in Arizona and Mexico. Host ants are unknown.
Haeterius brunneipennis in nest of Formica exsectoides
- Hippeutister *A myrmecophilous genus of six species found in North, Central, and South America. This genus is found in the nest of Solenopsis.
- Kaszabister *There are four species in this myrmecophilous genus that live in the nests of Solenopsis
- Pinaxister *A myrmecophilous genus found with Pheidole ants.
- Renclasea *This is a myrmecophilous genus of six species. Neivamyrmex is the probable host ant.
- Satrapes *This Palearctic genus of five species is characterized by its elongate and subcylindrical shape, short, very wide tibia, and triangular labrum. They live in ant nests of several genera including Lasius, Aphaenogaster, Formica, and Tetramorium.
- Sternocoelis *The genus Sternocoelis Lewis, 1888 is a small genus of myrmecophilous histerids with 27 described species distributed in the Mediterranean area with most species described from Morocco and Algeria.
- Terapus *There are fourteen species in this myrmecophilous genus of New World Histeridae. The host ant is the genus Pheidole.
- Ulkeus *There are six species in this myrmecophilous genus of Histeridae.
Cremastocheilina (Subtribe) eleven genera
- 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
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.
- Clinterocera *There are thirty-two species from China and adjacent regions; host ant Liometopum.
- Platysodes * Very little is known of the habitat and nothing of the life cycle of the enigmatic species in the genus Platysodes. However, they may be myrmecophilous since the mentum is extremely expanded and the maxillae are sclerotized. These are the same modifications that are seen in other species of Cremastocheilini that are myrmecophilous.
- Martineziana *Members of Martineziana are primarily associated with fire ants (Solenopsis spp.) (Stebnicka 2007). Other specimens have been collected with the Argentine ant, Iridomyrmex humilis (Mayr) (Stebnicka), and are occasionally collected at light.
Salpingidae or narrow-waisted bark beetles is a family of beetles, in the large suborder Polyphaga. The species are small, about 1.5 – 7 mm in length. This family is worldwide distributed and consists of about 45 genera and 300 species.
- Aphaenochara *This genus is closely allied to the genus Aleochara Gravenhorst, and is placed in the subtribe Aleocharina of the tribe Aleocharini.
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.
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.
- Archimicrodon * There are 45 species in this genus.
- Aristosyrphus * There are seven Neotropical species in this genus.
- Bardistopus * Form very slender; color black with the sides of front, pedicel and basoflagellomere, apex of scutellum, metanotum, broad lateral vitta on first abdominal segment and legs (except tarsi) reddish brown, tarsi yellowish white (from Mann, 1920).
- Carreramyia * There are five Neotropical species in this genus.
- Ceratophya * There are five Neotropical species in this genus.
- Pseudomicrodon * There are 14 known species. Larvae are found in ant nests. Native to the New World tropics.
- 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.
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.
- 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
There are a large number of mite species that live on and with ants.
A genus of mites that use ants for transportation (phoresy).
|Pheidole sp. alate||Caesarodispus pheidolei||Rahiminejad et al. 2015||described from mites found on alates in northern Iran|
|Tetramorium sp. alate||Caesarodispus khaustovi||Rahiminejad et al. 2015||described from mites found on alates in northern Iran|
|Tetramorium sp. alate||Caesarodispus nodijensis||Rahiminejad et al. 2015||described from mites found on alates in northern Iran|
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).
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).
- 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.
|Camponotus aethiops||Petalomium gottrauxi||Da Silva et al.. 2017||Known from Crimea and Iran|
|Myrmica ruginodis||Petalomium gottrauxi||Da Silva et al.. 2017||Known from Switzerland and Hungary|
|Neoponera verenae||Petalomium verenae||Da Silva et al., 2017||Brazil. Phoretic females attached to hairs between the first and second pairs of coxae|
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.
- Marachernes bellus (Harvey, 1992)
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- Da Silva, R. A., A. A. Khaustov, J. M. S. Lopes, J. H. C. Delabie, and A. R. Oliveira. 2017. A new species of Petalomium from Brazil with a redescription of Petalomium gottrauxi Mahunka 1977 (Acari: Heterostigmatina: Neopygmephoridae). Systematic and Applied Acarology. 22:1800-1812. doi:10.11158/saa.22.11.2
- Donisthorpe, H. 1927d. The guests of British ants, their habits and life-histories. London: G. Routledge and Sons, xxii + 244 pp.  124251
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- Harvey, M.S. 1992. A new genus of myrmecophilous Chernetidae from southern Australia (Pseudoscorpionida). Records of the Western Australian Museum 15: 763-775.
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- MARUYAMA, M., KOMATSU, T., KUDO, T., SHIMADA, T. & KINOMURA, K. 2013: The guests of Japanese ants. – Tokai University Press, Minamiyana, 208 pp.
- O'Keefe, S.T. 2000. Ant-like stone beetles, ants, and their associations(Coleoptera:Scydmaenidae;Hymenoptera:Formicidae;Isoptera). Journal of the New York Entomological Society. 108(3, 4): 273-303.
- Rahiminejad, V., H. Hajiqanbar, and A. A. Talebi. 2015. Three new species of the genus Caesarodispus (Acari: Microdispidae) associated with ants (Hymenoptera: Formicidae), with a key to species. Entomological Science. 18:461-469. doi:10.1111/ens.12149
- Parker, J. 2016. Myrmecophily in beetles (Coleoptera): evolutionary patterns and biological mechanisms. Myrmecological News(22): 65-108.
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- Sielezniew, M., D. Patricelli, R. Rutkowski, M. Witek, S. Bonelli, and M. M. Bus. 2015. Population genetics of the endangered obligatorily myrmecophilous butterfly Phengaris (=Maculinea) arion in two areas of its European range. Insect Conservation and Diversity. 8:505-516. doi:10.1111/icad.12129
- Smith, J.B. 1886. Ants' Nests and their inhabitants. The American Naturalist, vol. 20 (8):679-687.
- Wasmann, E. 1934. Die Ameisen, die Termiten und ihre Gäste. Regensburg: G. J. Manz, xviii + 148 pp.