Temnothorax

Temnothorax Temporal range: 37.2–0 Ma K Pa Eo Ol Mi Pl 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
536 species 1 fossil species (Species Checklist, Species by Country) Temnothorax adustus Specimen label
Synonyms
Antillaemyrmex Mann, 1920 Chalepoxenus Menozzi, 1923 Croesomyrmex Mann, 1920 Dichothorax Emery, 1895 Epimyrma Emery, 1915 Icothorax Hamann & Klemm, 1967 Leonomyrma Arnol'di, 1968 Macromischa Roger, 1863 Myrafant Smith, M.R., 1950 Myrmammophilus Menozzi, 1925 Myrmetaerus Soudek, 1925 Myrmoxenus Ruzsky, 1902 Protomognathus Wheeler, W.M., 1905

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

• 

  Temnothorax minutissimus alate gynes in an artificial nest with the host species, Temnothorax curvispinosus, workers and a queen.
• 

  Temnothorax rugatulus

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

• Key to Ant Genera of the Navajo Reservation
• Key to North American Genera of Myrmicinae
• Key to Vietnamese Myrmicinae Genera
• Temnothorax tebessae gapsensis
• Temnothorax tesquorum
• Temnothorax tuberum

Keys to Subgenera or Species Groups in this Genus

• Key to New World Temnothorax clades
• Temnothorax tebessae gapsensis
• Temnothorax tesquorum
• Temnothorax tuberum

Keys to Species in this Genus

• Key to Temnothorax of California
• Key to Afrotropical Temnothorax workers
• Key to Temnothorax of India
• Key to Temnothorax of mainland China
• Key to east Mediterranean Temnothorax muellerianus group
• Key to east Mediterranean Temnothorax recedens group
• Key to Temnothorax nylanderi species group workers
• Key to Temnothorax of Japan
• Key to Temnothorax andersoni species group workers
• Key to Temnothorax andrei species group workers
• Key to Temnothorax hispidus species group workers
• Key to Temnothorax longispinosus species group workers
• Key to Temnothorax nitens species group workers
• Key to Temnothorax schaumii species group workers
• Key to Temnothorax tricarinatus species group workers
• Key to the New World Temnothorax
• Key to Temnothorax of the Arabian Peninsula
• Key to Temnothorax interruptus group
• Key to Temnothorax of Pakistan
• Key to Greek species of Temnothorax anodonotoides species group
• Key to Temnothorax salvini clade workers
• Key to Temnothorax graecus species-group from Greece
• Key to Temnothorax of Nepal
• Key to the Cypriot Temnothorax species
• Key to the Temnothorax of Crete
• Key to Leptothorax and Temnothorax of China
• Key to Temnothorax corsicus species group
• Key to Temnothorax of eastern United States

Distribution

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	62	134	29	343
Total Species	2851	1736	3047	932	840	4391	1767	2925

Fossils

Fossils are known from: Baltic amber, Baltic Sea region, Europe (Priabonian, Late Eocene), Bitterfeld amber, Baltic Sea region, Europe (Priabonian, Late Eocene), Dominican amber, Dominican Republic (Burdigalian, Early Miocene), Rovno amber, Baltic Sea region, Europe (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

Seifert 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 adlerzi	workerless inquiline	ant	Temnothorax exilis	host	Greece	Douwes et al., 1988; Báthori et al., 2024
Temnothorax affinis	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	rare host
Temnothorax affinis	host	ant	Temnothorax ravouxi	dulotic parasite	southern Europe	Czechowski & Czechowska, 2000; Báthori et al., 2024
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 albipennis	host	ant	Temnothorax ravouxi	dulotic parasite		Czechowski & Czechowska, 2000
Temnothorax albipennis	host	fungus	Myrmicinosporidium durum	pathogen	Italy	Buschinger et al., 2004; Espadaler & Santamaria, 2012
Temnothorax algerianus	dulotic parasite	ant	Temnothorax curtulus	host		Cagniant, 1969; Buschinger et al., 1990
Temnothorax algerianus	dulotic parasite	ant	Temnothorax gentilis	host		Buschinger et al., 1990
Temnothorax algerianus	dulotic parasite	ant	Temnothorax monjauzei	host		Buschinger et al., 1990
Temnothorax algerianus	dulotic parasite	ant	Temnothorax racovitzai	host	Spain	Báthori et al., 2024
Temnothorax algerianus	dulotic parasite	ant	Temnothorax spinosus	host	Algeria, Morocco	Buschinger et al., 1990; Báthori et al., 2024	most common slave
Temnothorax algerianus	dulotic parasite	ant	Temnothorax tebessae	host		Buschinger et al., 1990; Báthori et al., 2024
Temnothorax algerianus	dulotic parasite	ant	Temnothorax unifasciatus	host	Spain	Báthori et al., 2024
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	dulotic parasite
Temnothorax ambiguus	host	ant	Temnothorax duloticus	dulotic parasite
Temnothorax ambiguus	host	ant	Temnothorax pilagens	dulotic parasite
Temnothorax americanus	dulotic parasite	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	dulotic parasite	ant	Temnothorax curvispinosus	host		Beibl et al., 2005; see de la Mora et al., 2021
Temnothorax americanus	dulotic parasite	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	dulotic parasite		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	dulotic parasite	ant	Temnothorax gredosi	host		Báthori et al., 2024
Temnothorax bernardi	dulotic parasite	ant	Temnothorax recedens	host		Báthori et al., 2024
Temnothorax bikara	dulotic parasite	ant	Temnothorax makora	host	Japan	Japanese Ant Image Database
Temnothorax bikara	dulotic parasite	ant	Temnothorax spinosior	host	Japan	Japanese Ant Image Database
Temnothorax birgitae	workerless inquiline	ant	Temnothorax gracilicornis	host	Canary Islands	Schulz, 1994; Báthori et al., 2024
Temnothorax birgitae	workerless inquiline	ant	Temnothorax gracilicornis nivarianus	host	Canary Islands	Schulz, 1994; Báthori et al., 2024
Temnothorax brunneus	dulotic parasite	ant	Temnothorax marocanus	host	Morocco	Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax brunneus	dulotic parasite	ant	Temnothorax rottenbergii	host	Morocco	Buschinger & Douwes, 1990
Temnothorax corsicus	workerless inquiline	ant	Temnothorax exilis	host		Báthori et al., 2024
Temnothorax crasecundus	host	ant	Temnothorax ravouxi	dulotic parasite	Georgia	Csösz et al., 2015	as Temnothorax tamarae
Temnothorax curtisetosus	inquiline	ant	Temnothorax antigoni	host	Turkey	Salata & Borowiec, 2015
Temnothorax curtisetosus	inquiline	ant	Temnothorax oreades	host	Cyprus	Salata et al., 2024
Temnothorax curtulus	host	ant	Temnothorax algerianus	dulotic parasite		Cagniant, 1969; Buschinger et al., 1990
Temnothorax curvispinosus	host	ant	Temnothorax americanus	dulotic parasite
Temnothorax curvispinosus	host	ant	Temnothorax duloticus	dulotic parasite
Temnothorax curvispinosus	host	ant	Temnothorax minutissimus	workerless inquiline
Temnothorax duloticus	dulotic parasite	ant	Temnothorax ambiguus	host		Beibl et al., 2005; de la Mora et al., 2021
Temnothorax duloticus	dulotic parasite	ant	Temnothorax curvispinosus	host		Beibl et al., 2005; see de la Mora et al., 2021
Temnothorax duloticus	dulotic parasite	ant	Temnothorax longispinosus	host		Beibl et al., 2005; de la Mora et al., 2021
Temnothorax flavicornis	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	rare host
Temnothorax gentilis	host	ant	Temnothorax algerianus	dulotic parasite		Buschinger et al., 1990
Temnothorax gordiagini	dulotic parasite	ant	Temnothorax serviculus	host		Buschinger et al., 1984; Báthori et al., 2024
Temnothorax gracilicornis	host	ant	Temnothorax birgitae	workerless inquiline	Canary Islands	Schulz, 1994; Báthori et al., 2024
Temnothorax gracilicornis nivarianus	host	ant	Temnothorax birgitae	workerless inquiline	Canary Islands	Schulz, 1994; Báthori et al., 2024
Temnothorax gredosi	host	ant	Temnothorax bernardi	dulotic parasite		Báthori et al., 2024
Temnothorax interruptus	host	ant	Temnothorax muellerianus	dulotic parasite		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	dulotic parasite	ant	Temnothorax recedens	host		Buschinger et al., 1984; Seifert, 2018; Suefuji & Heinze, 2014; Suefuji & Heinze, 2015; de la Mora et al., 2021; Báthori et al., 2024
Temnothorax kutteri	dulotic parasite	ant	Temnothorax aveli	host		Acosta Salmerón et al., 1983; Beibl et al., 2007	as ''Temnothorax rabaudi'', rare host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax aveli	host		Beibl et al., 2007; Cagniant, 1973	as ''Temnothorax berlandi'', rare host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax aveli	host		Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri	dulotic parasite	ant	Temnothorax exilis	host		Espadaler and Restrepo, 1983	as ''Temnothorax specularis'', fairly common host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax exilis	host		Acosta Salmerón et al., 1983	rare host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax exilis	host		Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri	dulotic parasite	ant	Temnothorax niger	host		Cagniant, 1973	rare host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax niger	host		Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri	dulotic parasite	ant	Temnothorax racovitzai	host		Buschinger et al., 1988	as ''Temnothorax massiliensis'', most common host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax racovitzai	host		Cagniant, 1973	rare host
Temnothorax kutteri	dulotic parasite	ant	Temnothorax racovitzai	host		Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri	dulotic parasite	ant	Temnothorax recedens	host		Espadaler & Restrepo, 1983; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax kutteri	dulotic parasite	ant	Temnothorax unifasciatus	host		Espadaler & Restrepo, 1983	rare host
Temnothorax lichtensteini	host	ant	Temnothorax gordiagini	dulotic parasite		Buschinger et al., 1984
Temnothorax lichtensteini	host	ant	Temnothorax menozzii	dulotic parasite		Báthori et al., 2024
Temnothorax lichtensteini	host	ant	Temnothorax ravouxi	dulotic parasite		Báthori et al., 2024
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	dulotic parasite
Temnothorax longispinosus	host	ant	Temnothorax duloticus	dulotic parasite
Temnothorax longispinosus	host	ant	Temnothorax pilagens	dulotic parasite
Temnothorax luteus	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	as ''Leptothorax tristis''
Temnothorax makora	host	ant	Temnothorax bikara	dulotic parasite	Japan	Japanese Ant Image Database
Temnothorax makora	host	ant	Temnothorax kinomurai	dulotic parasite	Japan	Japanese Ant Image Database
Temnothorax marae	host	ant	Temnothorax muellerianus	dulotic parasite	Sicily, Italy	Schifani et al., 2022
Temnothorax marocanus	host	ant	Temnothorax brunneus	dulotic parasite	Morocco	Buschinger & Douwes, 1990; Beibl et al., 2005; de la Mora et al., 2021
Temnothorax menozzii	dulotic parasite	ant	Temnothorax lichtensteini	host		Buschinger et al., 1984; Báthori et al., 2024
Temnothorax menozzii	dulotic parasite	ant	Temnothorax nylanderi	host		Buschinger et al., 1984; Báthori et al., 2024
Temnothorax menozzii	dulotic parasite	ant	Temnothorax serviculus	host		Buschinger et al., 1984; Báthori et al., 2024
Temnothorax menozzii	dulotic parasite	ant	Temnothorax unifasciatus	host		Buschinger et al., 1984; Báthori et al., 2024	rare
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 mongolicus	inquiline	ant	Temnothorax nassonovi	inquiline	Russian Federation	Antonov et al., 2024	co-occur in nests, relationship details unknown
Temnothorax monjauzei	host	ant	Temnothorax algerianus	dulotic parasite		Buschinger et al., 1990
Temnothorax muellerianus	dulotic parasite	ant	Temnothorax affinis	host		Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021	rare host
Temnothorax muellerianus	dulotic parasite	ant	Temnothorax exilis	host	Crete	Beibl et al., 2007; Salata, Borowiec & Trichas, 2018; de la Mora et al., 2021
Temnothorax muellerianus	dulotic parasite	ant	Temnothorax flavicornis	host		Beibl et al., 2007; Buschinger et al., 1988; de la Mora et al., 2021	rare host
Temnothorax muellerianus	dulotic parasite	ant	Temnothorax interruptus	host		Beibl et al., 2007; de la Mora et al., 2021
Temnothorax muellerianus	dulotic parasite	ant	Temnothorax luteus	host		Buschinger et al., 1988; de la Mora et al., 2021	as Leptothorax tristis
Temnothorax nadigi	host	ant	Temnothorax ravouxi	dulotic parasite	Poland	Czechowski & Czechowska, 2000
Temnothorax nassonovi	inquiline	ant	Temnothorax mongolicus	inquiline	Russian Federation	Antonov et al., 2024	co-occur in nests, relationship details unknown
Temnothorax niger	host	ant	Temnothorax kutteri	dulotic parasite		Cagniant, 1973; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021	rare host
Temnothorax nigriceps	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	common host
Temnothorax nigriceps	host	ant	Temnothorax ravouxi	dulotic parasite		Czechowski & Czechowska, 2000; Báthori et al., 2024
Temnothorax nigriceps	host	ant	Temnothorax stumperi	dulotic parasite		Báthori et al., 2024
Temnothorax nylanderi	host	ant	Temnothorax gordiagini	dulotic parasite		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	ant	Temnothorax menozzii	dulotic parasite		Báthori et al., 2024
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 oreades	host	ant	Temnothorax curtisetosus	inquiline	Cyprus	Salata et al., 2015
Temnothorax parvulus	host	cestode	Dilepididae (type 1(?))	parasite	Germany	Buschinger, 1973; Laciny, 2021
Temnothorax pilagens	dulotic parasite	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	dulotic parasite	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 algerianus	dulotic parasite		Báthori et al., 2024
Temnothorax racovitzai	host	ant	Temnothorax kutteri	dulotic parasite		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	dulotic parasite		Beibl et al., 2007; Buschinger et al., 1988
Temnothorax racovitzai	host	fungus	Myrmicinosporidium durum	pathogen		Espadaler & Santamaria, 2012
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax affinis	host	southern Europe	Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018; Suefuji & Heinze, 2014; Báthori et al., 2024	relatively rare
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax albipennis	host		Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax crasecundus	host	Georgia	Csösz et al., 2015
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax lichtensteini	host	Poland	Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax nadigi	host	Poland	Czechowski & Czechowska, 2000; de la Mora et al., 2021; Seifert, 2018
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax nigriceps	host		Czechowski & Czechowska, 2000; de la Mora et al., 2021; Beibl et al., 2005; Seifert, 2018; Suefuji & Heinze, 2014; Báthori et al., 2024
Temnothorax ravouxi	dulotic parasite	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 ravouxi	dulotic parasite	ant	Temnothorax tuberum	host	eastern Europe	Czechowski & Czechowska, 2000; Beibl et al., 2007; Seifert, 2018
Temnothorax ravouxi	dulotic parasite	ant	Temnothorax unifasciatus	host	mainly western Europe	Beibl et al., 2007; Blatrix & Sermage, 2005; Czechowski & Czechowska, 2000; de la Mora et al., 2021; Delattre et al., 2013; Gratiashvili et al., 2014; Seifert, 2018; Suefuji & Heinze, 2014; Báthori et al., 2024
Temnothorax ravouxi	host	nematode	Mermithidae (unspecified "Mermix")	parasite	Germany	Gosswald, 1930; Laciny, 2021	as ''Epimyrma gösswaldi''
Temnothorax recedens	host	ant	Temnothorax bernardi	dulotic parasite		Báthori et al., 2024
Temnothorax recedens	host	ant	Temnothorax kraussei	dulotic parasite		Buschinger et al., 1988; de la Mora et al., 2021; Seifert, 2018; Suefuji & Heinze, 2014; Suefuji & Heinze, 2015; Báthori et al., 2024
Temnothorax recedens	host	ant	Temnothorax kutteri	dulotic parasite		Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021
Temnothorax recedens	host	ant	Temnothorax muellerianus	dulotic parasite		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	dulotic parasite	Morocco	Buschinger & Douwes, 1990
Temnothorax rottenbergii	host	ant	Temnothorax muellerianus	dulotic parasite		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	dulotic parasite	Crete	Salata, Borowiec & Trichas, 2018
Temnothorax serviculus	host	ant	Temnothorax gordiagini	dulotic parasite		Buschinger et al., 1984; Báthori et al., 2024
Temnothorax spinosior	host	ant	Temnothorax bikara	dulotic parasite	Japan	Japanese Ant Image Database
Temnothorax spinosus	host	ant	Temnothorax algerianus	dulotic parasite	Algeria, Morocco	Buschinger et al., 1990; Báthori et al., 2024	the most common host
Temnothorax spinosus	host	ant	Temnothorax tramieri	dulotic parasite	Algeria, Morocco	Cagniant, 1983
Temnothorax stumperi	dulotic parasite	ant	Temnothorax nigriceps	host		Báthori et al., 2024
Temnothorax stumperi	dulotic parasite	ant	Temnothorax tuberum	host		de la Mora et al., 2021; Seifert, 2018; Báthori et al., 2024
Temnothorax stumperi	dulotic parasite	ant	Temnothorax unifasciatus	host		de la Mora et al., 2021; Seifert, 2018
Temnothorax subditivus	xenobiont	ant	Crematogaster carinata	xenobiont
Temnothorax tramieri	dulotic parasite	ant	Temnothorax spinosus	host	Algeria, Morocco	Cagniant, 1983
Temnothorax tuberum	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	rare host
Temnothorax tuberum	host	ant	Temnothorax ravouxi	dulotic parasite	eastern Europe	Czechowski & Czechowska, 2000
Temnothorax tuberum	host	ant	Temnothorax stumperi	dulotic parasite		Seifert, 2018; Báthori et al., 2024
Temnothorax tuberum	host	nematode	Mermithidae (unspecified "Mermix")	parasite	Germany	Gosswald, 1930; Laciny, 2021
Temnothorax unifasciatus	host	ant	Temnothorax algerianus	dulotic parasite		Báthori et al., 2024
Temnothorax unifasciatus	host	ant	Temnothorax gordiagini	dulotic parasite		Buschinger et al., 1984
Temnothorax unifasciatus	host	ant	Temnothorax kutteri	dulotic parasite		Espadaler & Restrepo, 1983; Buschinger et al., 1988; Beibl et al., 2007; de la Mora et al., 2021	rare host
Temnothorax unifasciatus	host	ant	Temnothorax menozzii	dulotic parasite		Báthori et al., 2024
Temnothorax unifasciatus	host	ant	Temnothorax muellerianus	dulotic parasite		Buschinger et al., 1988	common host
Temnothorax unifasciatus	host	ant	Temnothorax ravouxi	dulotic parasite	mainly western Europe	Czechowski & Czechowska, 2000; Báthori et al., 2024
Temnothorax unifasciatus	host	ant	Temnothorax stumperi	dulotic parasite		Seifert, 2018
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

 Explore: Show all Flight Month data or Search these data. See also a list of all data tables or learn how data is managed.

Click here to show/hide associate data.

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)

Morphology

Worker Morphology

 Explore: Show all Worker Morphology data or Search these data. See also a list of all data tables or learn how data is managed.

 • 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

• See additional details at the Ant Chromosome Database.

 Explore: Show all Karyotype data or Search these data. See also a list of all data tables or learn how data is managed.

Click here to show/hide karyotype data.

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 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 flavicornis	11			Italy	Fischer, 1987	as ''Leptothorax flavicornis''
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 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 menozzii	10			Croatia	Buschinger et al., 1983; Fischer, 1987	as ''Myrmoxenus gordiagini''
Temnothorax niger	18			France; Italy	Fischer, 1987	as ''Leptothorax niger''
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 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''

Phylogeny

Placement within the subfamily Myrmicinae

Myrmicinae

Myrmicini ⊞(2 genera) ⊟ Manica  (6 species, 2 fossil species) Myrmica  (214 species, 18 fossil species) Pogonomyrmecini ⊞(3 genera) ⊟ Patagonomyrmex  (3 species, 0 fossil species) Hylomyrma  (30 species, 0 fossil species) Pogonomyrmex  (95 species, 1 fossil species) Stenammini ⊞(7 genera) ⊟ Stenamma  (85 species, 1 fossil species) Novomessor  (3 species, 0 fossil species) Veromessor  (10 species, 0 fossil species) some Aphaenogaster  (219 species, 14 fossil species) some Aphaenogaster Messor  (169 species, 1 fossil species) Goniomma  (10 species, 1 fossil species) Oxyopomyrmex  (12 species, 0 fossil species) Solenopsidini ⊞(23 genera) ⊟ Stegomyrmex  (5 species, 0 fossil species) Dolopomyrmex  (1 species, 0 fossil species) some Rogeria  (40 species, 0 fossil species) Bariamyrma  (1 species, 0 fossil species) some Rogeria Solenopsis  (215 species, 9 fossil species) Kempfidris  (1 species, 0 fossil species) Tropidomyrmex  (1 species, 0 fossil species) Austromorium  (2 species, 0 fossil species) Chelaner  (53 species, 0 fossil species) some Monomorium Oxyepoecus  (21 species, 0 fossil species) Megalomyrmex  (45 species, 0 fossil species) Epelysidris  (1 species, 0 fossil species) Erromyrma  (1 species, 0 fossil species) Tyrannomyrmex  (4 species, 0 fossil species) Unicumyrmex  (1 species, 0 fossil species) Anillomyrma  (2 species, 0 fossil species) Myrmicaria  (71 species, 0 fossil species) some Monomorium Syllophopsis  (23 species, 0 fossil species) some Adelomyrmex  (31 species, 0 fossil species) Cryptomyrmex  (2 species, 0 fossil species) some Adelomyrmex Baracidris  (3 species, 0 fossil species) Attini ⊞(49 genera) ⊟ Ochetomyrmex  (2 species, 0 fossil species) Tranopelta  (2 species, 0 fossil species) Diaphoromyrma  (1 species, 0 fossil species) Lachnomyrmex  (16 species, 0 fossil species) Blepharidatta  (4 species, 0 fossil species) Allomerus  (8 species, 0 fossil species) Wasmannia  (11 species, 0 fossil species) Pheidole  (1,297 species, 8 fossil species) Cephalotes  (123 species, 16 fossil species) Procryptocerus  (44 species, 0 fossil species) Strumigenys  (882 species, 4 fossil species) Phalacromyrmex  (1 species, 0 fossil species) Pilotrochus  (1 species, 0 fossil species) Protalaridris  (7 species, 0 fossil species) Rhopalothrix  (19 species, 0 fossil species) Basiceros  (9 species, 0 fossil species) Octostruma  (35 species, 0 fossil species) Eurhopalothrix  (55 species, 0 fossil species) Talaridris  (1 species, 0 fossil species) Acanthognathus  (7 species, 1 fossil species) Daceton  (2 species, 0 fossil species) Lenomyrmex  (7 species, 0 fossil species) Microdaceton  (4 species, 0 fossil species) Orectognathus  (29 species, 0 fossil species) Colobostruma  (16 species, 0 fossil species) Epopostruma  (20 species, 0 fossil species) Mesostruma  (9 species, 0 fossil species) Paleoattina Apterostigma  (44 species, 2 fossil species) Mycocepurus  (6 species, 0 fossil species) Myrmicocrypta  (31 species, 0 fossil species) Neoattina Cyatta  (1 species, 0 fossil species) Kalathomyrmex  (1 species, 0 fossil species) Mycetarotes  (4 species, 0 fossil species) Mycetosoritis  (2 species, 0 fossil species) some Cyphomyrmex  (23 species, 2 fossil species) some Cyphomyrmex Paramycetophylax  (1 species, 0 fossil species) Mycetophylax  (21 species, 0 fossil species) Mycetagroicus  (4 species, 0 fossil species) Mycetomoellerius  (33 species, 1 fossil species) Sericomyrmex  (11 species, 0 fossil species) Xerolitor  (1 species, 0 fossil species) Paratrachymyrmex  (9 species, 0 fossil species) Trachymyrmex  (9 species, 0 fossil species) Amoimyrmex  (3 species, 0 fossil species) Atta  (20 species, 1 fossil species) some Acromyrmex  (51 species, 0 fossil species) some Acromyrmex Pseudoatta  (2 species, 0 fossil species) Crematogastrini ⊞(65 genera) ⊟ Rostromyrmex  (1 species, 6 fossil species) Cardiocondyla  (90 species, 0 fossil species) Ocymyrmex  (34 species, 0 fossil species) Nesomyrmex  (84 species, 2 fossil species) Xenomyrmex  (5 species, 0 fossil species) Terataner  (14 species, 0 fossil species) Atopomyrmex  (3 species, 0 fossil species) Cataulacus  (65 species, 3 fossil species) Carebara  (251 species, 9 fossil species) Diplomorium  (1 species, 0 fossil species) Melissotarsus  (4 species, 1 fossil species) Rhopalomastix  (14 species, 0 fossil species) Calyptomyrmex  (38 species, 0 fossil species) Strongylognathus  (27 species, 0 fossil species), Tetramorium  (603 species, 2 fossil species) Cyphoidris  (4 species, 0 fossil species) Dicroaspis  (2 species, 0 fossil species) Aretidris  (2 species, 0 fossil species) Vollenhovia  (83 species, 3 fossil species) Dacetinops  (7 species, 0 fossil species) Indomyrma  (2 species, 0 fossil species) Crematogaster  (785 species, 0 fossil species) Meranoplus  (93 species, 0 fossil species) Lophomyrmex  (13 species, 0 fossil species) Adlerzia  (1 species, 0 fossil species) Recurvidris  (12 species, 0 fossil species) Stereomyrmex  (3 species, 0 fossil species) Trichomyrmex  (29 species, 0 fossil species) Eutetramorium  (3 species, 0 fossil species) Royidris  (15 species, 0 fossil species) Malagidris  (6 species, 0 fossil species) Vitsika  (16 species, 0 fossil species) Huberia  (2 species, 0 fossil species) Podomyrma  (62 species, 1 fossil species) Liomyrmex  (1 species, 0 fossil species) Metapone  (33 species, 0 fossil species) Kartidris  (6 species, 0 fossil species) Mayriella  (9 species, 0 fossil species) Tetheamyrma  (2 species, 0 fossil species) Dacatria  (1 species, 0 fossil species) Proatta  (1 species, 0 fossil species) Dilobocondyla  (22 species, 0 fossil species) Secostruma  (1 species, 0 fossil species) Acanthomyrmex  (19 species, 0 fossil species) Myrmecina  (106 species, 0 fossil species) Perissomyrmex  (6 species, 0 fossil species) Pristomyrmex  (61 species, 3 fossil species) some Lordomyrma  (36 species, 0 fossil species) Propodilobus  (1 species, 0 fossil species) Lasiomyrma  (4 species, 0 fossil species) some Lordomyrma Ancyridris  (2 species, 0 fossil species) some Lordomyrma Paratopula  (12 species, 0 fossil species) Poecilomyrma  (2 species, 0 fossil species) Romblonella  (10 species, 0 fossil species) Rotastruma  (3 species, 0 fossil species) Gauromyrmex  (3 species, 0 fossil species) Vombisidris  (19 species, 0 fossil species) Temnothorax  (536 species, 1 fossil species) Harpagoxenus  (4 species, 0 fossil species) Formicoxenus  (8 species, 0 fossil species) Leptothorax  (20 species, 0 fossil species)

See Phylogeny of Myrmicinae for details.

Relationships among major species clades within the genus

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.

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.

As evidenced by the extensive taxonomic changes shown above, this is a diverse genera (one of the seven ant genera with > 500 species) and taxonomically-complex collection of species. The genus and its members have been studied and revised by many ant taxomomists. This has resulted in a variety of species being assigned to various genera and sub-genera closely aligned with Temnothorax and in turn many species being subsumed back into Temnothorax from genera and sub-genera deemed to be synonoms.

Matthew Prebus (2015, 2017, 2020, 2021) is currently working to comprehensively resolve the taxonomy and systematics of the genus on a worldwide basis. The foundation of this work is explained in Prebus (2017):

Temnothorax is a highly morphologically and behaviorally diverse genus. Despite recent taxonomically focused molecular phylogenies [Beibl, Stuart, Heinze 2005, Beibl, Buschinger, Foitzik 2007, Heinze, et al. 2015] and extensive taxonomic works [Mackay 2000, Baroni Urbani, C. 1978], Temnothorax remains widely renowned as one of the most taxonomically difficult ant genera [Buschinger, 1981]: its status as a phylogenetically natural group has been repeatedly revised [Bolton 2003] and called into question [Ward, 2007]. This is a product of multiple factors: subtle morphological variation on a global scale among groups of species, striking departures from the morphological norm in others, morphological convergence with distantly related genera [Ward, et al. 2015], weak diagnostic characters for the genus as a whole, and regional treatments of species in the absence of global understanding of the clade structure (Prebus 2017, Additional file 1). The current absence of an infrageneric classification system for Temnothorax is not for a lack of effort, though: past taxonomists erected five subgenera to compartmentalize the species diversity of this genus, none of which are currently recognized due to a lack of clear diagnostic characters [Bolton 2003]. My studies of Temnothorax are designed to elucidate the phylogeny of Temnothorax on a global scale in order to gain insight into the structure of relationships within the genus, verify that the genus is monophyletic as currently diagnosed, understand the biogeographical history and taxonomic evolution, and revise the taxonomy of Temnothorax.

In addition, Sándor Csősz, Sebastian Salata and Lech Borowiec, and numerous co-authors, are actively revising the European Temnothorax species, e.g., Csösz, Heinze & Mikó 2015; Csősz, Salata & Borowiec 2018; Salata, Srodon & Boroweic 2023; Salata & Borowiec 2015, 2019, 2022; Salata, Borowiec & Trichas 2018; Seifert & Csősz 2015.

References

• Arnol'di, K. V.; Dlussky, G. M. 1978. Superfam. Formicoidea. 1. Fam. Formicidae - ants. Pp. 519-556 in: Medvedev, G. S. (ed.) Keys to the insects of the European part of the USSR. Vol. 3. Hymenoptera. Part 1. Opredeliteli Faune SSSR 119:3-584. (page 543, Temnothorax as genus)
• Atanassov, N.; Dlussky, G. M. 1992. Fauna of Bulgaria. Hymenoptera, Formicidae. Fauna Bûlg. 22: 1-310 (page 125, Temnothorax as genus)
• Barden, P. 2017. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages. Myrmecological News 24: 1-30.
• Baroni Urbani, C. 1971c. Catalogo delle specie di Formicidae d'Italia (Studi sulla mirmecofauna d'Italia X). Memorie della Societa Entomologica Italiana 50: 5-287 (page 96, Temnothorax as junior synonym of Leptothorax)
• Bernard, F. 1967a [1968]. Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale. Paris: Masson, 411 pp. (page 185, Temnothorax as genus)
• Blaimer, B.B., Ward, P.S., Schultz, T.R., Fisher, B.L., Brady, S.G. 2018. Paleotropical diversification dominates the evolution of the hyperdiverse ant tribe Crematogastrini (Hymenoptera: Formicidae). Insect Systematics and Diversity 2(5): 3; 1-14 (doi:10.1093/isd/ixy013).
• Bolton, B. 1982. Afrotropical species of the myrmecine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor and Cataulacus (Formicidae). Bulletin of the British Museum (Natural History). Entomology, 46: 307-370 (page 319, Temnothorax as junior synonym of Leptothorax)
• Bolton, B. 1994. Identification guide to the ant genera of the world. Cambridge, Mass.: Harvard University Press, 222 pp. (page 105, Temnothorax as junior synonym of Leptothorax)
• Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 252, 270, Temnothorax as genus; Temnothorax senior synonym of Antillaemyrmex, Croesomyrmex, Dichothorax, Icothorax, Macromischa, Myrmammophilus, Myrafant)
• Bondroit, J. 1918. Les fourmis de France et de Belgique. Ann. Soc. Entomol. Fr. 87: 1-174 (page 117, Temnothorax as subgenus of Leptothorax)
• Boudinot, B.E. 2019. Hormigas de Colombia. Cap. 15. Clave para las subfamilias y generos basada en machos. Pp. 487-499 in: Fernández, F., Guerrero, R.J., Delsinne, T. (eds.) 2019d. Hormigas de Colombia. Bogotá: Universidad Nacional de Colombia, 1198 pp.
• Brown, W. L., Jr. 1973b. A comparison of the Hylean and Congo-West African rain forest ant faunas. Pp. 161-185 in: Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Wash (page 185, Temnothorax as junior synonym of Leptothorax [provisional])
• Burchill, A.T., Moreau, C.S. 2016. Colony size evolution in ants: macroevolutionary trends. Insectes Sociaux 63, 291–298 (doi:10.1007/s00040-016-0465-3).
• Buschinger, A. 2001. Multiple Hybridisierung von Arten der Ameisen-Gattung Epimyrma (Hymenoptera: Formicidae), und Beobachtungen zur Ausbeutung der Wirtsarten durch die Parasiten. Myrmecol. Nachr. 4, 25–35.
• Buschinger, A.; Linksvayer, T. A. 2004. Novel blend of life history traits in an inquiline ant, Temnothorax minutissimus, with description of the male (Hymenoptera: Formicidae). Myrmecol. Nachr. 6: 67-76.
• Cantone S. 2018. Winged Ants, The queen. Dichotomous key to genera of winged female ants in the World. The Wings of Ants: morphological and systematic relationships (self-published).
• Dalla Torre, K. W. von. 1893. Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann, 289 pp. (page 122, Temnothorax in Myrmicinae; Temnothorax as subgenus of Leptothorax)
• Dlussky, G. M.; Fedoseeva, E. B. 1988. Origin and early stages of evolution in ants. Pp. 70-144 in: Ponomarenko, A. G. (ed.) Cretaceous biocenotic crisis and insect evolution. Moskva: Nauka, 232 pp. (page 79, Temnothorax as genus)
• Donisthorpe, H. 1943h. A list of the type-species of the genera and subgenera of the Formicidae. [concl.]. Ann. Mag. Nat. Hist. 11(10): 721-737 (page 731, Temnothorax in Myrmicinae, Leptothoracini; Temnothorax as subgenus of Leptothorax)
• Emery, C. 1877b. Saggio di un ordinamento naturale dei Mirmicidei, e considerazioni sulla filogenesi delle formiche. Bull. Soc. Entomol. Ital. 9: 67-83 (page 81, Temnothorax in Myrmicidae, Myrmicidae)
• Emery, C. 1915e. Formiche raccolte nell'Eritrea dal Prof. F. Silvestri. Boll. Lab. Zool. Gen. Agrar. R. Sc. Super. Agric. 10: 3-26 (page 24, Temnothorax as subgenus of Leptothorax)
• Emery, C. 1916a [1915]. Fauna entomologica italiana. I. Hymenoptera.-Formicidae. Bull. Soc. Entomol. Ital. 47: 79-275 (page 176, Temnothorax as subgenus of Leptothorax)
• Emery, C. 1924f [1922]. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [concl.]. Genera Insectorum 174C: 207-397 (page 259, Temnothorax in Myrmicinae, Leptothoracini; Temnothorax as subgenus of Leptothorax)
• Emery, C.; Forel, A. 1879. Catalogue des Formicides d'Europe. Mitt. Schweiz. Entomol. Ges. 5: 441-481 (page 459, Temnothorax in Myrmicinae [Myrmicidae])
• Fernandez, F., Guerrero, R.J., Sánchez-Restrepo, A.F. 2021. Sistemática y diversidad de las hormigas neotropicales. Revista Colombiana de Entomología 47, 1–20 (doi:10.25100/socolen.v47i1.11082).
• Forel, A. 1890b. Fourmis de Tunisie et de l'Algérie orientale. Ann. Soc. Entomol. Belg. 34:lxi-lxxvi. (page lxxxii, Temnothorax as junior synonym of Leptothorax)
• Forel, A. 1892j. Die Ameisenfauna Bulgariens. (Nebst biologischen Beobachtungen.). Verh. K-K. Zool.-Bot. Ges. Wien 42: 305-318 (page 315, Temnothorax as subgenus of Leptothorax)
• Forel, A. 1915d. Fauna insectorum helvetiae. Hymenoptera. Formicidae. Die Ameisen der Schweiz. Mitt. Schweiz. Entomol. Ges. 12(B Beilage: 1-77 (page 27, Temnothorax as subgenus of Leptothorax)
• Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 245, Temnothorax in Myrmicinae, Leptothoracini; Temnothorax as subgenus of Leptothorax)
• Heinze, J., Buschinger, A., Poettinger, T. & Suefuji, M. 2015. Multiple convergent origins of workerlessness and inbreeding in the socially parasitic ant genus Myrmoxenus. PLoS ONE 10(7): e0131023. doi:10.1371/journal.pone.0131023.
• Jessen, K., Klinkicht, M. 1990. Hybridization in the social parasitic ant genus Epimyrma (Hymenoptera, Formicidae). Insectes Sociaux 37, 273–293.
• Jansen, G., Savolainen, R. 2010. Molecular phylogeny of the ant tribe Myrmicini (Hymenoptera: Formicidae). Zoological Journal of the Linnean Society 160(3), 482–495 (doi:10.1111/j.1096-3642.2009.00604.x).
• MacKay, W. P. 2000. A review of the New World ants of the subgenus Myrafant, (Genus Leptothorax) (Hymenoptera: Formicidae). Sociobiology. 36:265-444.
• Mayr, G. 1861. Die europäischen Formiciden. Nach der analytischen Methode bearbeitet. Wien: C. Gerolds Sohn, 80 pp. (page 68, Temnothorax in Myrmicinae [Myrmicidae])
• Mayr, G. 1865. Formicidae. In: Reise der Österreichischen Fregatte "Novara" um die Erde in den Jahren 1857, 1858, 1859. Zoologischer Theil. Bd. II. Abt. 1. Wien: K. Gerold's Sohn, 119 pp. (page 21, Temnothorax in Myrmicinae [Myrmicidae])
• Prebus, M. 2017. Insights into the evolution, biogeography and natural history of the acorn ants, genus Temnothorax Mayr (hymenoptera: Formicidae). Bmc Evolutionary Biology. 17:250. doi:10.1186/s12862-017-1095-8
• Prebus, M.M. 2020. Phylogenomic species delimitation in the ants of the Temnothorax salvini group (Hymenoptera: Formicidae): an integrative approach. Systematic Entomology. 46:307-326. doi:10.1111/syen.12463
• Prebus, M.M. 2021. Taxonomic revision of the Temnothorax salvini clade (Hymenoptera: Formicidae), with a key to the clades of New World Temnothorax. PeerJ 9, e11514 doi:10.7717/peerj.11514.
• Ruzsky, M. 1905b. The ants of Russia. (Formicariae Imperii Rossici). Systematics, geography and data on the biology of Russian ants. Part I. Tr. Obshch. Estestvoispyt. Imp. Kazan. Univ. 38(4-6 6: 1-800 (page 607, Temnothorax as subgenus of Leptothorax)
• Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 139, Temnothorax as subgenus of Leptothorax)
• Wheeler, W. M. 1922i. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bull. Am. Mus. Nat. Hist. 45: 631-710 (page 679, Temnothorax as subgenus of Leptothorax)