Temnothorax

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Temnothorax
Temporal range: 37.2–0 Ma Eocene – Recent
Temnothorax adustus
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Crematogastrini
Alliance: Formicoxenus genus group
Genus: Temnothorax
Mayr, 1861
Type species
Myrmica recedens, now Temnothorax recedens
Diversity
504 species
7 fossil species
(Species Checklist, Species by Country)

Temnothorax adustus lateral view

Temnothorax adustus

Temnothorax adustus dorsal view

Specimen label

Synonyms
Evolutionary Relationships

Cardiocondyla

Ocymyrmex

Nesomyrmex

Xenomyrmex

Terataner

Atopomyrmex

Cataulacus

Carebara

Diplomorium

Melissotarsus

Rhopalomastix

Calyptomyrmex

Tetramorium

Cyphoidris

Dicroaspis

Aretidris

Vollenhovia

Dacetinops

Indomyrma

Crematogaster

Meranoplus

Lophomyrmex

Adlerzia

Recurvidris

Stereomyrmex

Trichomyrmex

Eutetramorium

Royidris

Malagidris

Vitsika

Huberia

Podomyrma

Liomyrmex

Metapone

Kartidris

Mayriella

Tetheamyrma

Dacatria

Proatta

Dilobocondyla

Secostruma

Acanthomyrmex

Myrmecina

Perissomyrmex

Pristomyrmex

some Lordomyrma

Propodilobus

Lasiomyrma

[some Lordomyrma

Ancyridris

some Lordomyrma

Paratopula

Poecilomyrma

Romblonella

Rotastruma

Gauromyrmex

Vombisidris

Temnothorax

Harpagoxenus

Formicoxenus

Leptothorax

Based on Ward et al. (2014) and Blaimer et al. (2018).
Evolutionary Relationships within Temnothorax
Temnothorax

Temnothorax andrei clade

Temnothorax sallei clade

Temnothorax rottenbergii clade

Temnothorax rugatulus clade

Temnothorax andersoni clade

Temnothorax obturator clade

Temnothorax salvini clade

Palearctic clade

Based on Prebus, 2021.

The workers of Temnothorax species are generally small. Colonies are typically monogynous, although facultative polygyny has been documented in several species (Alloway et al. 1983, Buschinger 1974, Buschinger and Linksvayer 2004, Rüppell et al. 1998). Colony populations are usually quite small, often with less than 100 workers (Beckers et al. 1989). However, several studies have found colonies of some species to be widely dispersed with several to many satellite nests (Alloway et al. 1983, Foitzik and Heinze 2001, Partridge et al. 1997). Many species are arboreal, living within hollow stems, old beetle or termite galleries, or in galls (Mackay 2000). Temnothorax species appear to be trophic generalists, feeding on a wide variety of scavenged items, including the elaiosomes of seeds (Espadaler 1997, Fokuhl et al. 2012). None have been documented to be active or aggressive predators. (Snelling et al. 2014)

The socially parasitic species formerly placed in the genus Myrmoxenus (synonymised with Temnothorax by Ward et al., 2014) vary strongly in fundamental life history traits, such as queen-worker ratio, the timing of sexual production, and mating behavior. Queens of these species generally take over nests of other Temnothorax species, kill the resident queen by throttling, and force the workers to take care of the social parasite’s brood. Young queens of Temnothorax ravouxi and other species produce large numbers of workers, which during “slaveraids” pillage host pupae from neighboring Temnothorax host colonies to increase the workforce in their own nests. Other species, such as Temnothorax corsicus, have lost caste polyphenism and rear only male and female sexual offspring. (Heinze et al., 2015.)


Photo Gallery

Identification

Antennal count is typically twelve segments but there are a small number of species with only eleven.

New World Temnothorax species groups

Eguchi, Bui and Yamane (2011) - The worker of Temnothorax is similar to that of Vombisidris and Cardiocondyla.

Keys including this Genus

Keys to Subgenera or Species Groups in this Genus

Keys to Species in this Genus

Distribution

Distribution and Richness based on AntMaps

Species by Region

Number of species within biogeographic regions, along with the total number of species for each region.

Afrotropical Region Australasian Region Indo-Australian Region Malagasy Region Nearctic Region Neotropical Region Oriental Region Palaearctic Region
Species 10 0 0 0 61 134 26 304
Total Species 2839 1735 3036 932 834 4378 1708 2836

Fossils

Fossils are known from: Baltic amber (Bartonian, Middle to Late Eocene), Bitterfeld amber (Bartonian, Middle to Late Eocene), Dominican amber, Dominican Republic (Burdigalian, Early Miocene), Rovno amber (Priabonian, Late Eocene).

Biology

Matthew Prebus is actively investigating the systematics and biology of the genus. The following is from a recent study (Prebus 2017, see this publication for citations):

General Biology

While the biology of most species of Temnothorax is poorly known, several taxa are frequently used as model organisms for social insect behavior and ecology. Studies on this genus have led investigators to discover new forms of social learning and inspired hypotheses about speciation and the origins of parasitism.

Temnothorax has a primarily Holarctic distribution, but notable exceptions include several species in Sub-Saharan Africa and many species in Mesoamerica, including the islands of the Caribbean . Generally, Temnothorax ants are encountered in the mesic forests of the Northern hemisphere at mid-to-high elevations, where they are typically found under rocks, in leaf litter, or as arboreal foragers. However, these ants have adapted to a broad variety of habitats, from arid deserts to tropical rain forests, and are found at elevations from sea level up to 4000 m. The workers of most species in the Holarctic region are diminutive, timid, slow moving, and cryptically colored. Because of their inconspicuous nature, these ants are often overlooked despite their broad geographical distribution. The islands of the Greater Antilles harbor many Temnothorax endemics, however, that provide an exception to this habitus: the islands Cuba and Hispaniola, for example, have more than thirty described endemic species, which are large, brightly colored, and capable of delivering powerful stings.

Temnothorax ants are believed to be trophic generalists, and have been observed scavenging for a variety of food items, including dead insects, honeydew on leaves without tending aphids, honeydew from tended aphids and membracids, the axillary nectaries of bracken ferns and Baccharis (an Asteraceae genus), seeds, and elaiosomes. In competitions at baits, they appear to be opportunistic foragers, often absconding as more aggressive species arrive, and occasionally insinuating themselves into baits dominated by other species (Prebus, pers. obs.). Recruitment of workers to food and nest sites often appears to involve tandem running.

Nests of Temnothorax are generally small, often with fewer than 200 workers, which may be distributed among several satellite nests. With the worker caste of most species measuring less than four millimeters in length, these ants often take advantage of small cavities for their nesting sites, such as crevices in rocks, hollow nut shells, dead twigs, or directly in the soil. Temnothorax colonies tend to be monogynous, with only one queen per nest. This is hardly the rule, however: this genus displays a remarkable diversity of sociometry, with many species being either functionally monogynous (with several queens present, but only one actively producing eggs), or facultatively polygynous, both within and among species.

Many species appear to be arboreal specialists, nesting in dead hollow twigs on live trees, under bark, in galls, or under the roots of epiphytes.

Several taxa have intriguing life histories: it appears that Temnothorax and its sister group, the Leptothorax genus group (composed of Formicoxenus, Harpagoxenus, and Leptothorax) are unusually prone to developing a set of interrelated lifestyles collectively known as ‘social parasitism’.

Dulotic Temnothorax

Siefert et al. (2014) - Three slave-making species of the Temnothorax genus group (Bolton 2003) of the ant tribe Formicoxenini are known from North America. Each of the three represents an unmistakable combination of phenotypic characters. They all use species of the genus Temnothorax Forel, 1890 as hosts and cluster genetically with species of this genus (Beibl et al. 2005). The first species and outgroup to all the others (Beibl et al. 2005), Temnothorax americanus (Emery, 1895), is characterized by an elongated, semi-rectangular head capsule with extremely long antennal scobes that fully accommodate the short and flattened scape when it is folded back. These characters are a convergence to the Holarctic genus Harpagoxenus that belongs to the distantly related Leptothorax genus group. The second one, Temnothorax duloticus (Wesson, 1937), shows an acute, frontoventrally directed dent on the postpetiolar sternite, a high petiole with a bulky, truncate node that slopes down to the caudal cylinder with a distinct step. This particular petiolar shape and the reduction of antennal segments to 11 resemble the situation in the subgenus Mychothorax Ruzsky, 1904 of the genus Leptothorax Mayr, 1855. However, T. duloticus differs from the latter by the absence of a curved transverse carina on the stipes of the maxillae. On the third species….only little information exists to date (Herbers and Foitzik 2002, Beibl et al. 2005). Its phenotype is an unmistakable combination of an acute, frontoventrally directed dent on postpetiolar sternite, a stout, hump-backed mesosoma, small scape length, a high petiole that is in lateral aspect roughly triangular, a wide petiole and reduced mandibular dentition. Robin Stuart was the first who recognized the new species (Herbers and Foitzik 2002). We follow his proposal to name this slave-making species Temnothorax pilagens.

Phylogeny and Biogeography

Molecular analyses suggested the genus first arose 35Ma, with an initial succession of rapid diversification into distinctive clades. Early diversification occurred in the Neartic with large scale dispersal (Carribean, Neotropical mainland, Palearctic) following soon after the formation of the initial clades. The phylogeny clarifies and resolves numerous long standing problems with various past generic, subgeneric and species-group hypotheses. The data show support for Temnothorax originating as a terrestrial nesting species that later evolved arboreal species. Parasitism appears to have arisen numerous times, as it is not confined to a single node of the phylogeny.

Eguchi, Bui and Yamane (2011) - Temnothorax species are rare in Vietnam where they have been collected around or above 1000 m alt. in northern Vietnam.

Association with Other Organisms

All Associate Records for Genus

Click here to show/hide associate data.
Taxon Relationship Associate Type Associate Taxon Associate Relationship Locality Source Notes
Temnothorax affinis host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 rare host
Temnothorax affinis host ant Temnothorax ravouxi slave maker southern Europe Czechowski & Czechowska, 2000
Temnothorax affinis host cestode Anomotaenia brevis parasite Germany Scharf et al., 2012; Laciny, 2021
Temnothorax affinis host cestode Dilepididae (type 1(?)) parasite Germany Buschinger, 1973; Laciny, 2021
Temnothorax affinis host fungus Myrmicinosporidium durum pathogen France, Croatia, Switzerland Buschinger & Winter, 1983; Espadaler & Santamaria, 2012
Temnothorax albipennis host ant Temnothorax ravouxi slave maker Czechowski & Czechowska, 2000
Temnothorax albipennis host fungus Myrmicinosporidium durum pathogen Italy Buschinger et al., 2004; Espadaler & Santamaria, 2012
Temnothorax algerianus slave maker ant Temnothorax curtulus host Buschinger et al., 1990
Temnothorax algerianus slave maker ant Temnothorax gentilis host Buschinger et al., 1990
Temnothorax algerianus slave maker ant Temnothorax monjauzei host Buschinger et al., 1990
Temnothorax algerianus slave maker ant Temnothorax spinosus host Algeria, Morocco Buschinger et al., 1990 the most common slave
Temnothorax allardycei host eucharitid wasp Orasema minuta parasite Heraty, 1990; Baker et al., 2019; Universal Chalcidoidea Database primary host
Temnothorax allardycei host eucharitid wasp Orasema minuta parasitoid Quevillon, 2018 encounter mode independent; direct transmission; transmission outside nest
Temnothorax ambiguus host ant Temnothorax americanus slave maker
Temnothorax ambiguus host ant Temnothorax duloticus slave maker
Temnothorax ambiguus host ant Temnothorax pilagens slave maker
Temnothorax americanus slave maker ant Temnothorax ambiguus host Beibl et al., 2005; Blatrix & Herbers, 2003; de la Mora et al., 2021; Herbers & Foitzik, 2002; Swan & Hare, 2012
Temnothorax americanus slave maker ant Temnothorax curvispinosus host Beibl et al., 2005; see de la Mora et al., 2021
Temnothorax americanus slave maker ant Temnothorax longispinosus host Beibl et al., 2005; see de la Mora et al., 2021
Temnothorax andrei xenobiont ant Camponotus yogi xenobiont United States Creighton & Snelling, 1966; Kanizsai et al., 2013 Chaparral. In living stems of ''Haplopappus pinifolius''.
Temnothorax andrei xenobiont ant Formica moki xenobiont Mann, 1911 nature of relationship unknown
Temnothorax andrei xenobiont ant Formica ravida xenobiont Mackay & Mackay, 1984 nature of relationship unknown
Temnothorax angustulus host fungus Myrmicinosporidium durum pathogen Espadaler & Santamaria, 2012
Temnothorax antigoni host ant Temnothorax curtisetosus inquiline Turkey Salata & Borowiec, 2015
Temnothorax aveli host ant Temnothorax kutteri slave maker Acosta Salmerón et al., 1983; Cagniant, 1973; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax aveli host cestode Dilepididae (type 1(?)) parasite Germany Buschinger, 1973; Laciny, 2021
Temnothorax bernardi host ant Temnothorax algerianus slave maker Buschinger et al., 1990
Temnothorax bikara slave maker ant Temnothorax makora host Japan Japanese Ant Image Database
Temnothorax bikara slave maker ant Temnothorax spinosior host Japan Japanese Ant Image Database
Temnothorax brunneus slave maker ant Temnothorax marocanus host Morocco Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax brunneus slave maker ant Temnothorax rottenbergii host Morocco Buschinger & Douwes, 1990
Temnothorax corsicus workerless inquiline ant Temnothorax exilis host
Temnothorax crasecundus host ant Temnothorax tamarae slave maker Georgia Csösz et al., 2015
Temnothorax curtisetosus inquiline ant Temnothorax antigoni host Turkey Salata & Borowiec, 2015
Temnothorax curtulus host ant Temnothorax algerianus slave maker Buschinger et al., 1990
Temnothorax curvispinosus host ant Temnothorax americanus slave maker
Temnothorax curvispinosus host ant Temnothorax duloticus slave maker
Temnothorax curvispinosus host ant Temnothorax minutissimus workerless inquiline
Temnothorax duloticus slave maker ant Temnothorax ambiguus host Beibl et al., 2005; de la Mora et al., 2021
Temnothorax duloticus slave maker ant Temnothorax curvispinosus host Beibl et al., 2005; see de la Mora et al., 2021
Temnothorax duloticus slave maker ant Temnothorax longispinosus host Beibl et al., 2005; de la Mora et al., 2021
Temnothorax exilis host ant Temnothorax corsicus workerless inquiline
Temnothorax exilis host ant Temnothorax kutteri slave maker Acosta Salmerón et al., 1983; Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 fairly common host
Temnothorax exilis host ant Temnothorax muellerianus slave maker Crete Salata, Borowiec & Trichas, 2018
Temnothorax exilis host fungus Myrmicinosporidium durum pathogen Espadaler & Santamaria, 2012
Temnothorax flavicornis host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 rare host
Temnothorax gentilis host ant Temnothorax algerianus slave maker Buschinger et al., 1990
Temnothorax gordiagini slave maker ant Temnothorax lichtensteini host Buschinger et al., 1984
Temnothorax gordiagini slave maker ant Temnothorax nylanderi host Buschinger et al., 1984
Temnothorax gordiagini slave maker ant Temnothorax serviculus host Buschinger et al., 1984
Temnothorax gordiagini slave maker ant Temnothorax unifasciatus host Buschinger et al., 1984
Temnothorax interruptus host ant Temnothorax muellerianus slave maker Beibl et al., 2007; Buschinger et al., 1988
Temnothorax interruptus host nematode Mermithidae (unspecified "Mermix") parasite Germany Gosswald, 1930; Laciny, 2021
Temnothorax kinomurai workerless inquiline ant Temnothorax makora host
Temnothorax kraussei slave maker ant Temnothorax recedens host Buschinger et al., 1984; Seifert, 2018; Suefuji & Heinze, 2014; Suefuji & Heinze, 2015; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax aveli host Acosta Salmerón et al., 1983; Beibl et al., 2007 as ''Temnothorax rabaudi'', rare host
Temnothorax kutteri slave maker ant Temnothorax aveli host Beibl et al., 2007; Cagniant, 1973 as ''Temnothorax berlandi'', rare host
Temnothorax kutteri slave maker ant Temnothorax aveli host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax exilis host Espadaler and Restrepo, 1983 as ''Temnothorax specularis'', fairly common host
Temnothorax kutteri slave maker ant Temnothorax exilis host Acosta Salmerón et al., 1983 rare host
Temnothorax kutteri slave maker ant Temnothorax exilis host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax niger host Cagniant, 1973 rare host
Temnothorax kutteri slave maker ant Temnothorax niger host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax racovitzai host Buschinger et al., 1988 as ''Temnothorax massiliensis'', most common host
Temnothorax kutteri slave maker ant Temnothorax racovitzai host Cagniant, 1973 rare host
Temnothorax kutteri slave maker ant Temnothorax racovitzai host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax recedens host Espadaler & Restrepo, 1983; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri slave maker ant Temnothorax unifasciatus host Espadaler & Restrepo, 1983 rare host
Temnothorax kutteri slave maker ant Temnothorax unifasciatus host Beibl et al., 2005; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax lichtensteini host ant Temnothorax gordiagini slave maker Buschinger et al., 1984
Temnothorax lichtensteini host cestode Choanotaenia crateriformis parasite Spain Espadaler & Riasol, 1983; Laciny, 2021
Temnothorax lichtensteini host fungus Myrmicinosporidium durum pathogen Espadaler & Santamaria, 2012
Temnothorax longispinosus host ant Temnothorax americanus slave maker
Temnothorax longispinosus host ant Temnothorax duloticus slave maker
Temnothorax longispinosus host ant Temnothorax pilagens slave maker
Temnothorax luteus host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 as ''Leptothorax tristis''
Temnothorax makora host ant Temnothorax bikara slave maker Japan Japanese Ant Image Database
Temnothorax makora host ant Temnothorax kinomurai slave maker Japan Japanese Ant Image Database
Temnothorax marae host ant Temnothorax muellerianus slave maker Sicily, Italy Schifani et al., 2022
Temnothorax marocanus host ant Temnothorax brunneus slave maker Morocco Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax minutissimus workerless inquiline ant Temnothorax curvispinosus host Beibl et al., 2005; Buschinger & Linksvayer, 2004; de la Mora et al., 2021; Johnson et al., 2008
Temnothorax monjauzei host ant Temnothorax algerianus slave maker Buschinger et al., 1990
Temnothorax muellerianus host fungus Myrmicinosporidium durum pathogen Italy Sanchez-Peña et al., 1993; Buschinger et al., 2004; Espadaler & Santamaria, 2012
Temnothorax muellerianus slave maker ant Temnothorax affinis host Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021 rare host
Temnothorax muellerianus slave maker ant Temnothorax exilis host Crete Beibl et al., 2007; Salata, Borowiec & Trichas, 2018; de la Mora et al., 2021
Temnothorax muellerianus slave maker ant Temnothorax flavicornis host Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021 rare host
Temnothorax muellerianus slave maker ant Temnothorax interruptus host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax muellerianus slave maker ant Temnothorax luteus host Buschinger et al., 1988; de la Mora et al., 2021 as Leptothorax tristis
Temnothorax muellerianus slave maker ant Temnothorax marae host Sicily, Italy Schifani et al., 2022; de la Mora et al., 2021
Temnothorax muellerianus slave maker ant Temnothorax nigriceps host Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021 common host
Temnothorax muellerianus slave maker ant Temnothorax racovitzai host Beibl et al., 2007; de la Mora et al., 2021
Temnothorax muellerianus slave maker ant Temnothorax recedens host Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021 common host
Temnothorax muellerianus slave maker ant Temnothorax rottenbergii host Beibl et al., 2007; de la Mora et al., 2021 as ''Temnothorax'' cf. ''rottenbergii''
Temnothorax muellerianus slave maker ant Temnothorax semiruber host Crete Beibl et al., 2007; Salata, Borowiec & Trichas, 2018; de la Mora et al., 2021
Temnothorax muellerianus slave maker ant Temnothorax tuberum host Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021 rare host
Temnothorax muellerianus slave maker ant Temnothorax unifasciatus host Beibl et al., 2007; Blatrix & Sermage, 2005; Buschinger et al., 1988; Seifert, 2018; de la Mora et al., 2021 most common host
Temnothorax nadigi host ant Temnothorax ravouxi slave maker Poland Czechowski & Czechowska, 2000
Temnothorax niger host ant Temnothorax kutteri slave maker Cagniant, 1973; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax nigriceps host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 common host
Temnothorax nigriceps host ant Temnothorax ravouxi slave maker Czechowski & Czechowska, 2000
Temnothorax nigriceps host nematode Mermithidae (unspecified "Mermix") parasite Germany Gosswald, 1930; Laciny, 2021
Temnothorax nylanderi host ant Temnothorax gordiagini slave maker Buschinger et al., 1983 ''T. nylanderi'' as the host of ''Temnothorax gordiagini'' had been suggested by Soudek, 1925, (cited in Buschinger et al., 1983) though this has never been confirmed and was probably a misidentification
Temnothorax nylanderi host cestode Anomotaenia brevis parasite France Gabrion et al., 1976; Laciny, 2021
Temnothorax nylanderi host cestode Anomotaenia brevis parasite Germany Scharf et al., 2012; Beros et al., 2015; Feldmeyer et al., 2016; Laciny, 2021
Temnothorax nylanderi host cestode Anomotaenia brevis parasite France (lab colony) Trabalon et al., 2000; Laciny, 2021
Temnothorax nylanderi host cestode Anomotaenia brevis parasitoid Quevillon, 2018 encounter mode secondary; indirect transmission; transmission outside nest
Temnothorax nylanderi host cestode Anomotaenia sp. parasite Spain Espadaler et al., 2011; Laciny, 2021
Temnothorax nylanderi host cestode Choanotaenia crateriformis parasite Spain Espadaler & Riasol, 1983; Laciny, 2021
Temnothorax nylanderi host cestode Dilepididae (type 1(?)) parasite Germany Buschinger, 1973; Laciny, 2021
Temnothorax nylanderi host fungus Myrmicinosporidium durum pathogen Espadaler & Santamaria, 2012
Temnothorax parvulus host cestode Dilepididae (type 1(?)) parasite Germany Buschinger, 1973; Laciny, 2021
Temnothorax pilagens slave maker ant Temnothorax ambiguus host Michigan, New York, & Vermont, United States de la Mora et al., 2021; Fleeberg & Foitzik, 2016; Seifert et al., 2014
Temnothorax pilagens slave maker ant Temnothorax longispinosus host Michigan, New York, & Vermont, United States de la Mora et al., 2021; Fleeberg & Foitzik, 2016; Seifert et al., 2014
Temnothorax purpuratus host eupelmid wasp Anastatus myrmecobius parasite Universal Chalcidoidea Database associate
Temnothorax purpuratus host eupelmid wasp Anastatus myrmecobius parasitoid Quevillon, 2018 multiple encounter modes; direct transmission; transmission outside nest
Temnothorax racovitzai host ant Temnothorax kutteri slave maker Buschinger et al., 1988; Cagniant, 1973; Beibl et al., 2007; de la Mora et al., 2021 most common host
Temnothorax racovitzai host ant Temnothorax muellerianus slave maker Beibl et al., 2007; Buschinger et al., 1988
Temnothorax racovitzai host fungus Myrmicinosporidium durum pathogen Espadaler & Santamaria, 2012
Temnothorax ravouxi host nematode Mermithidae (unspecified "Mermix") parasite Germany Gosswald, 1930; Laciny, 2021 as ''Epimyrma gösswaldi''
Temnothorax ravouxi slave maker ant Temnothorax affinis host southern Europe Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018; Suefuji & Heinze, 2014 relatively rare
Temnothorax ravouxi slave maker ant Temnothorax albipennis host Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018
Temnothorax ravouxi slave maker ant Temnothorax nadigi host Poland Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018
Temnothorax ravouxi slave maker ant Temnothorax nigriceps host Czechowski & Czechowska, 2000; de la Mora et al., 2021; Beibl et al., 2005; Seifert, 2018; Suefuji & Heinze, 2014
Temnothorax ravouxi slave maker ant Temnothorax tuberum host eastern Europe Czechowski & Czechowska, 2000; Beibl et al., 2007; Seifert, 2018
Temnothorax ravouxi slave maker ant Temnothorax unifasciatus host mainly western Europe Beibl et al., 2007; Blatrix & Sermage, 2005; Czechowski & Czechowska, 2000; Delattre et al., 2013; Seifert, 2018; Suefuji & Heinze, 2014
Temnothorax recedens host ant Temnothorax kraussei slave maker Buschinger et al., 1988; de la Mora et al., 2021; Seifert, 2018; Suefuji & Heinze, 2014; Suefuji & Heinze, 2015
Temnothorax recedens host ant Temnothorax kutteri slave maker Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax recedens host ant Temnothorax muellerianus slave maker Buschinger et al., 1988; de la Mora et al., 2021 common host
Temnothorax recedens host fungus Myrmicinosporidium durum pathogen France, Croatia, Switzerland Buschinger & Winter, 1983; Espadaler & Santamaria, 2012
Temnothorax recedens host gregarine Mattesia geminata parasite Germany, Italy Buschinger & Kleespies, 1999
Temnothorax rottenbergii host ant Temnothorax brunneus slave maker Morocco Buschinger & Douwes, 1990
Temnothorax rottenbergii host ant Temnothorax muellerianus slave maker Beibl et al., 2007; Buschinger et al., 1988
Temnothorax rugatulus host cestode Dilepididae unspecified (cf. Anomotaenia) parasite Arizona/New Mexico, United States Heinze et al., 1998; Laciny, 2021
Temnothorax semiruber host ant Temnothorax muellerianus slave maker Crete Salata, Borowiec & Trichas, 2018
Temnothorax serviculus host ant Temnothorax gordiagini slave maker Buschinger et al., 1984
Temnothorax spinosior host ant Temnothorax bikara slave maker Japan Japanese Ant Image Database
Temnothorax spinosus host ant Temnothorax algerianus slave maker Algeria, Morocco Buschinger et al., 1990 the most common host
Temnothorax spinosus host ant Temnothorax tramieri slave maker Algeria, Morocco Cagniant, 1983
Temnothorax stumperi slave maker ant Temnothorax tuberum host de la Mora et al., 2021; Seifert, 2018
Temnothorax stumperi slave maker ant Temnothorax unifasciatus host de la Mora et al., 2021; Seifert, 2018
Temnothorax subditivus xenobiont ant Crematogaster carinata xenobiont
Temnothorax tamarae slave maker ant Temnothorax crasecundus host Georgia Csösz et al., 2015
Temnothorax tamarae slave maker ant Temnothorax sp. NYL2 host Georgia de la Mora et al., 2021; Gratiashvili et al., 2014 an undescribed species closely related to ''Temnothorax nylanderi''
Temnothorax tamarae slave maker ant Temnothorax unifasciatus host Georgia Gratiashvili et al., 2014 needs confirmation
Temnothorax tramieri slave maker ant Temnothorax spinosus host Algeria, Morocco Cagniant, 1983
Temnothorax tuberum host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 rare host
Temnothorax tuberum host ant Temnothorax ravouxi slave maker eastern Europe Czechowski & Czechowska, 2000
Temnothorax tuberum host ant Temnothorax stumperi slave maker Seifert, 2018
Temnothorax tuberum host fungus Myrmicinosporidium durum pathogen Germany Holldobler, 1933; Espadaler & Santamaria, 2012
Temnothorax tuberum host nematode Mermithidae (unspecified "Mermix") parasite Germany Gosswald, 1930; Laciny, 2021
Temnothorax unifasciatus host ant Temnothorax gordiagini slave maker Buschinger et al., 1984
Temnothorax unifasciatus host ant Temnothorax kutteri slave maker Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021 rare host
Temnothorax unifasciatus host ant Temnothorax muellerianus slave maker Buschinger et al., 1988 common host
Temnothorax unifasciatus host ant Temnothorax ravouxi slave maker mainly western Europe Czechowski & Czechowska, 2000
Temnothorax unifasciatus host ant Temnothorax stumperi slave maker Seifert, 2018
Temnothorax unifasciatus host ant Temnothorax tamarae slave maker Georgia Gratiashvili et al., 2014 needs confirmation
Temnothorax unifasciatus host cestode Anomotaenia sp. parasite Spain Espadaler et al., 2011; Laciny, 2021
Temnothorax unifasciatus host cestode Choanotaenia crateriformis parasite Spain Espadaler & Riasol, 1983; Laciny, 2021
Temnothorax unifasciatus host cestode Dilepididae (type 1(?)) parasite Germany Buschinger, 1973; Laciny, 2021
Temnothorax unifasciatus host fungus Myrmicinosporidium durum pathogen Italy, France, Croatia, Switzerland Buschinger & Winter, 1983; Sanchez-Peña et al., 1993; Buschinger et al., 2004; Espadaler & Santamaria, 2012
Temnothorax unifasciatus host gregarine Mattesia geminata parasite Germany, Italy Buschinger & Kleespies, 1999
Temnothorax unifasciatus host nematode Mermithidae (unspecified "Mermix") parasite Germany Gosswald, 1930; Laciny, 2021

Flight Period

All Flight Records for Genus

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Taxon Month Source Notes
Temnothorax affinis Jul Aug antkeeping.info
Temnothorax albipennis Jul antkeeping.info
Temnothorax andrei Jun Jul Cole, 1958; antkeeping.info
Temnothorax caguatan Jun Jul antkeeping.info
Temnothorax congruus Jul Japan
Temnothorax corticalis Jul Aug Sep antkeeping.info
Temnothorax crassispinus Jul Aug Sep antkeeping.info
Temnothorax curvispinosus Jun Jul Aug antkeeping.info
Temnothorax interruptus Jun Jul Aug Sep antkeeping.info
Temnothorax nigriceps Jun Jul Aug Sep antkeeping.info
Temnothorax nylanderi Jul Aug Sep antkeeping.info
Temnothorax parvulus Aug Sep antkeeping.info
Temnothorax ravouxi Jun Jul Aug antkeeping.info
Temnothorax saxonicus Jul antkeeping.info
Temnothorax sordidulus Jul antkeeping.info
Temnothorax spinosior Jul Japan
Temnothorax tuberum Jun Jul Aug antkeeping.info
Temnothorax unifasciatus Jun Jul Aug antkeeping.info
Temnothorax vivianoi Aug Schifani et al., 2022 Sicily, Italy

Life History Traits

  • Mean colony size: 47-325 (Greer et al., 2021)
  • Compound colony type: dulosis; inquilinism (Greer et al., 2021)
  • Nest site: hypogaeic; arboreal (Greer et al., 2021)
  • Diet class: omnivore (Greer et al., 2021)
  • Foraging stratum: subterranean/leaf litter; arboreal (Greer et al., 2021)
  • Foraging behaviour: cooperative (Greer et al., 2021)

Castes

Morphology

Worker Morphology

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 • Eyes: 11-100 ommatidia • Pronotal Spines: absent • Mesonotal Spines: absent • Propodeal Spines: dentiform; present • Petiolar Spines: absent • Caste: none or weak • Sting: present • Metaplural Gland: absent; present • Cocoon: absent

Karyotype

Species Uncertain

  • Temnothorax cf. interrupus: n = 12 (Greece) (Fischer, 1987) (as Leptothorax cf. interruptus).
  • Temnothorax cf. lichtensteini: n = 14 (France; Spain) (Fischer, 1987) (as Leptothorax cf. lichtensteini).
  • Temnothorax: n = 9 (Croatia, France, Greece) (Fischer, 1987) (as Leptothorax spp., tuberum group).

All Karyotype Records for Genus

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Taxon Haploid Diploid Karyotype Locality Source Notes
Temnothorax 12 Greece Fischer, 1987 as ''Leptothorax'' cf. ''interruptus''
Temnothorax 14 France; Spain Fischer, 1987 as ''Leptothorax'' cf. ''lichtensteini''
Temnothorax 9 Croatia, France, Greece Fischer, 1987 as ''Leptothorax'' spp., ''tuberum'' group
Temnothorax adlerzi 10 Greece Douwes et al., 1988 as ''Epimyrma adlerzi''
Temnothorax affinis 9 France, Germany, Switzerland Fischer, 1987 as ''Leptothorax affinis''
Temnothorax albipennis 8 Germany Fischer, 1987 as ''Leptothorax albipennis'', ''L. tuberointerruptus'', ''Stenamma albipennis''
Temnothorax albipennis 8 Great Britain Orledge, 1998 as ''Leptothorax albipennis'', ''L. tuberointerruptus'', ''Stenamma albipennis''
Temnothorax algerianus 10 Douwes et al., 1988 as ''Epimyrma algeriana''
Temnothorax ambiguus 22 Canada Fischer, 1987 as ''Leptothorax ambiguus''
Temnothorax americanus 11 Canada; USA Fischer, 1987 as ''Harpagoxenus americanus''
Temnothorax andrei 10 USA Fischer, 1987 as ''Leptothorax andrei''
Temnothorax angustulus 23 France Fischer, 1987 as ''Leptothorax angustulus''
Temnothorax aveli 9 France Fischer, 1987 as ''Leptothorax rabaudi''
Temnothorax bernardi 10 Spain Buschinger et al., 1986; Fischer, 1987 as ''Epimyrma bernardi''
Temnothorax congruus 9 18 Japan Imai & Kubota, 1972 as ''Leptothorax congruus''
Temnothorax corsicus 10 France Buschinger et al., 1986; Fischer, 1987 as ''Epimyrma corsica''
Temnothorax corticalis 12 Germany Fischer, 1987 as ''Leptothorax corticalis''
Temnothorax corticalis 9 Crozier, 1975 as ''Leptothorax corticalis''
Temnothorax curvispinosus 23 USA Fischer, 1987 as ''Leptothorax curvispinosus''
Temnothorax duloticus 23 Canada Fischer, 1987 as ''Leptothorax duloticus''
Temnothorax duloticus 24 Canada Fischer, 1987 as ''Leptothorax duloticus''
Temnothorax duloticus 25 Canada Fischer, 1987 as ''Leptothorax duloticus''
Temnothorax exilis 11 Croatia; France; Italy; Turkey Fischer, 1987 as ''Leptothorax exilis''
Temnothorax exilis 17 Italy Fischer, 1987 as ''Leptothorax specularis''
Temnothorax flavicornis 11 Italy Fischer, 1987 as ''Leptothorax flavicornis''
Temnothorax gordiagini 10 Croatia Buschinger et al., 1983; Fischer, 1987 as ''Myrmoxenus gordiagini''
Temnothorax gredosi 17 Spain Fischer, 1987 as ''Leptothorax gredosi''
Temnothorax interruptus 24 Germany Hauschteck-Jungen & Jungen, 1983 as ''Leptothorax interruptus''
Temnothorax interruptus 12 Croatia; Germany Fischer, 1987 as ''Leptothorax interruptus''
Temnothorax kraussei 10 France, Italy, Spain Fischer, 1987 as ''Epimyrma kraussei''
Temnothorax kutteri 12 France Fischer, 1987 as ''Chalepoxenus kutteri''
Temnothorax lichtensteini 14 Croatia, France, Italy Fischer, 1987 as ''Leptothorax lichtensteini''
Temnothorax longispinosus 12 Canada Fischer, 1987 as ''Leptothorax longispinosus''
Temnothorax longispinosus 12 24 USA Crozier, 1970b as ''Leptothorax longispinosus''
Temnothorax luteus 21 France; Switzerland Fischer, 1987 as ''Leptothorax tristis''
Temnothorax mediterraneus 14 28 France, Italy, Spain Fischer, 1987 as ''Leptothorax kraussei''
Temnothorax melas 13 France Fischer, 1987 as ''Leptothorax melas''
Temnothorax muellerianus 12 Croatia; France; Greece; Italy Fischer, 1987 as ''Chalepoxenus siciliensis''
Temnothorax niger 18 France; Italy Fischer, 1987 as ''Leptothorax niger''
Temnothorax nigriceps 18 Switzerland Hauschteck, 1962 as ''Leptothorax nigriceps''
Temnothorax nigriceps 9 Croatia; France; Germany; Italy Fischer, 1987 as ''Leptothorax nigriceps''
Temnothorax nylanderi 11 Croatia, France, Germany Fischer, 1987 as ''Leptothorax nylanderi''
Temnothorax nylanderi 11 22 Switzerland Hauschteck-Jungen & Jungen, 1983 as ''Leptothorax nylanderi''
Temnothorax parvulus 14 France, Italy, Switzerland Fischer, 1987 as ''Leptothorax parvulus''
Temnothorax racovitzai 21 France Fischer, 1987 as ''Leptothorax racovitzai''
Temnothorax ravouxi 10 France; Germany; Italy; Switzerland Buschinger et al., 1986; Fischer, 1987 as ''Epimyrma ravouxi''
Temnothorax recedens 12 Italy Fischer, Karl, 1987 as ''Leptothorax recedens''
Temnothorax rottenbergii 11 Italy Fischer, 1987 as ''Leptothorax rottenbergii''
Temnothorax rougeti 8 Fischer, 1987 as ''Leptothorax rougeti'' (subspecies of ''Temnothorax unifasciatus'')
Temnothorax rugatulus 26 USA Taber & Cokendolpher, 1988 as ''Leptothorax rugatulus''
Temnothorax rugatulus 27 USA Taber & Cokendolpher, 1988 as ''Leptothorax rugatulus''
Temnothorax rugatulus 14 USA Fischer, 1987 as ''Leptothorax rugatulus''
Temnothorax schaumii 9 USA Hauschteck-Jungen & Jungen, 1983 as ''Leptothorax schaumii''
Temnothorax sordidulus 11 Croatia, Italy, Switzerland Fischer, 1987 as ''Leptothorax sordidulus''
Temnothorax spinosior 12 24 Japan Imai, 1966; Imai, 1969; Imai, 1974; Imai & Yosida, 1964; Fischer, 1987 as ''Leptothorax spinosior''
Temnothorax spinosus 16 Algeria Fischer, 1987 as ''Leptothorax spinosus''
Temnothorax stumperi 10 France; Switzerland Buschinger et al., 1986; Fischer, 1987 as ''Epimyrma stumperi''
Temnothorax tuberum 9 France, Sweden, Switzerland Fischer, 1987 as ''Leptothorax tuberum''
Temnothorax tuberum 9 18 Switzerland Hauschteck, 1962; Hauschteck-Jungen & Jungen, 1983 as ''Leptothorax tuberum''
Temnothorax unifasciatus 9 Croatia; France; Germany; Greece; Italy Fischer, 1987 as ''Leptothorax unifasciatus''
Temnothorax unifasciatus 9 18 Germany; Switzerland Hauschteck-Jungen & Jungen, 1983 as ''Leptothorax unifasciatus''

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • TEMNOTHORAX [Myrmicinae: Formicoxenini]
    • Temnothorax Mayr, 1861: 68. Type-species: Myrmica recedens, by monotypy.
    • Temnothorax junior synonym of Leptothorax: Forel, 1890a: lxxii.
    • Temnothorax revived from synonymy as subgenus of Leptothorax: Forel, 1892i: 315; maintained as subgenus: Emery, 1915g: 24; Forel, 1915d: 27; Bondroit, 1918: 117; Wheeler, W.M. 1922a: 679; Emery, 1924d: 259.
    • Temnothorax junior synonym of Leptothorax: Baroni Urbani, 1971c: 96; Bolton, 1982: 319; Bolton, 1994: 105.
    • Temnothorax revived status as genus: Bernard, 1967: 185; Arnol'di & Dlussky, 1978: 543; Dlussky & Fedoseeva, 1988: 79; Atanasov & Dlussky, 1992: 125; Bolton, 2003: 252, 270.
    • Temnothorax senior synonym of Antillaemyrmex, Croesomyrmex, Dichothorax, Icothorax, Macromischa, Myrmammophilus, Myrafant: Bolton, 2003: 252, 270.
    • Current subgenera of Temnothorax: nominal plus Gonepimyrma.
  • ANTILLAEMYRMEX [junior synonym of Temnothorax]
    • Antillaemyrmex Mann, 1920: 408 [as subgenus of Macromischa]. Type-species: Macromischa (Antillaemyrmex) terricola, by original designation.
    • Antillaemyrmex raised to genus: Wheeler, W.M. 1931b: 4 (in text).
    • Antillaemyrmex junior synonym of Macromischa: Smith, M.R. 1937: 849; Smith, M.R. 1939e: 502.
    • Antillaemyrmex junior synonym of Leptothorax: Snelling, R.R. 1986: 154.
    • Antillaemyrmex junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • CHALEPOXENUS [junior synonym of Temnothorax]
    • Chalepoxenus Menozzi, 1923a: 257. Type-species: Chalepoxenus gribodoi (junior synonym of Leptothorax muellerianus), by monotypy.
    • [Name of type-species initially misspelled gridoboi, p. 257; correct spelling as gribodoi, p. 258.]
    • Chalepoxenus senior synonym of Leonomyrma: Buschinger, 1987: 117.
    • Chalepoxenus junior synonym of Temnothorax: Ward et al., 2014: 15.
  • CROESOMYRMEX [junior synonym of Temnothorax]
    • Croesomyrmex Mann, 1920: 408 [as subgenus of Macromischa]. Type-species: Macromischa (Croesomyrmex) wheeleri, by original designation.
    • Croesomyrmex raised to genus: Wheeler, W.M. 1931b: 4 (in text).
    • Croesomyrmex junior synonym of Macromischa: Smith, M.R. 1937: 849; Smith, M.R. 1939e: 502.
    • Croesomyrmex junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • DICHOTHORAX [junior synonym of Temnothorax]
    • Dichothorax Emery, 1895c: 323 [as subgenus of Leptothorax]. Type-species: Leptothorax (Dichothorax) pergandei, by subsequent designation of Wheeler, W.M. 1911f: 161.
    • Dichothorax raised to genus: Ashmead, 1905b: 383.
    • Dichothorax subgenus of Leptothorax: Wheeler, W.M. 1910g: 139.
    • Dichothorax junior synonym of Leptothorax: Bolton, 1982: 319.
    • Dichothorax junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • EPIMYRMA [junior synonym of Myrmoxenus]
    • Epimyrma Emery, 1915a: 262. Type-species: Epimyrma kraussei, by original designation.
    • Epimyrma senior synonym of Myrmoxenus (and its junior synonym Myrmetaerus): Buschinger, 1990: 245 (provisional synoynym); Bolton, 1994: 105 [incorrect seniority].
    • Epimyrma junior synonym of Myrmoxenus: Schulz & Sanetra, 2002: 162.
  • ICOTHORAX [junior synonym of Temnothorax]
    • Icothorax Hamann & Klemm, 1967: 415 [as subgenus of Leptothorax]. Type-species: Leptothorax (Icothorax) megalops, by monotypy.
    • Icothorax junior synonym of Myrafant: Baroni Urbani, 1978b: 556; Bolton, 1995b: 32.
    • Icothorax junior synonym of Chalepoxenus: Hölldobler & Wilson, 1990: 13.
    • Icothorax junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • LEONOMYRMA [junior synonym of Chalepoxenus]
    • Leonomyrma Arnol'di, 1968: 1809. Type-species: Leonomyrma spinosa, by original designation.
    • Leonomyrma junior synonym of Chalepoxenus: Buschinger, 1987: 117.
  • MACROMISCHA [junior synonym of Temnothorax]
    • Macromischa Roger, 1863a: 184. Type-species: Macromischa purpurata, by subsequent designation of Wheeler, W.M. 1911f: 166.
    • Macromischa subgenus of Leptothorax: Baroni Urbani, 1978b: 398.
    • Macromischa revived status as genus: Smith, D.R. 1979: 1390.
    • Macromischa senior synonym of Antillaemyrmex, Croesomyrmex: Smith, M.R. 1937: 849; Smith, M.R. 1939e: 502; Baroni Urbani, 1978b: 398.
    • Macromischa junior synonym of Leptothorax: Snelling, R.R. 1986: 154; Bolton, 1994: 105; Bolton, 1995b: 34.
    • Macromischa junior synonym of Temnothorax: Bolton, 2003: 252, 271.
  • MYRAFANT [junior synonym of Temnothorax]
    • Myrafant Smith, M.R. 1950: 30 [as subgenus of Leptothorax]. Type-species: Leptothorax curvispinosus, by original designation.
    • Myrafant senior synonym of Icothorax: Baroni Urbani, 1978b: 556.
    • Myrafant junior synonym of Leptothorax: Bolton, 1982: 319.
    • Myrafant junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • MYRMAMMOPHILUS [junior synonym of Temnothorax]
    • Myrmammophilus Menozzi, 1925d: 29 [as subgenus of Leptothorax]. Type-species: Leptothorax (Myrmammophilus) finzii, by monotypy.
    • Myrmammophilus junior synonym of Leptothorax: Bolton, 1982: 319.
    • Myrmammophilus junior synonym of Temnothorax: Bolton, 2003: 253, 271.
  • MYRMETAERUS [junior synonym of Myrmoxenus]
    • Myrmetaerus Soudek, 1925a: 33. Type-species: Myrmetaerus microcellatus (junior synonym of Myrmoxenus gordiagini), by monotypy.
    • [Myrmetaerus also described as new by Soudek, 1925b: 14.]
    • Myrmetaerus junior synonym of Camponotus: Snelling, R.R. 1981: 404 (error).
    • Myrmoxenus junior synonym of Epimyrma: Bolton, 1994: 105.
    • Myrmetaerus junior synonym of Myrmoxenus: Buschinger, Winter & Faber, 1984: 336; Bolton, 2003: 248.
  • MYRMOXENUS [junior synonym of Temnothorax]
    • Myrmoxenus Ruzsky, 1902b: 474. Type-species: Myrmoxenus gordiagini, by monotypy.
    • Myrmoxenus senior synonym of Myrmetaerus: Buschinger, Winter & Faber, 1984: 336.
    • Myrmoxenus junior synonym of Epimyrma: Bolton, 1994: 105 [incorrect seniority].
    • Myrmoxenus senior synonym of Epimyrma: Schulz & Sanetra, 2002: 162.
    • Myrmoxenus junior synonym of Temnothorax: Ward et al., 2014: 15.
  • PROTOMOGNATHUS [junior synonym of Temnothorax]
    • Protomognathus Wheeler, W.M. 1905a: 3 [as subgenus of Tomognathus]. Type-species: Tomognathus americanus, by monotypy.
    • Protomognathus junior synonym of Harpagoxenus: Emery, 1924d: 265.
    • Protomognathus revived from synonymy and raised to genus: Cover, in Hölldobler & Wilson, 1990: 65.
    • Protomognathus junior synonym of Temnothorax: Ward et al., 2014: 15.

References