Temporal range: 48.6–0 Ma Middle Eocene – Recent
12 fossil species
(Species Checklist, Species by Country)
|Based on Ward et al., 2014|
This is a large and diverse genus which is found throughout much of the world (except southern South America and southern Africa).
- 1 Identification
- 2 Distribution
- 3 Biology
- 4 Castes
- 5 Morphology
- 6 Nomenclature
- 7 References
Antennae are 12 segmented (including the scape) with a 4 segmented club. In side view the propodeum is depressed below the level of the pronotum and forward (anterior) section of the mesonotum, these two regions being connected by the steeply sloping rear (posterior) section of the mesonotum. All workers from a nest are approximately the same size (monomorphic).
Aphaenogaster is most often confused with Pheidole or possibly Carebara. They can be separated from Pheidole by the 4 segmented rather than 3 segmented club and larger body size (over 3.4mm long), and from Carebara by the 12 segmented antennae (they are 11 segmented in Carebara). Additionally, both Pheidole and Carebara have major and minor workers (Pheidole is dimorphic, Carebara polymorphic) while Aphaenogaster has only a single worker caste (is monomorphic).
The Australian species of Aphaenogaster show differences which are little more than variation on a theme. This is in contrast to the nearby Papua New Guinea fauna where morphological variation is considerable. This difference suggests that the Australian fauna is composed of closely related species while that of PNG consists of several more distantly related lineages.
North American species
These ants can usually be easily distinguished by their elongate, slender habitus (general appearance). Their head is usually longer than broad, eye large, convex and placed at the middle of the head. The mesonotum of the worker is elongate and depressed, the propodeum usually has a pair of spines or small teeth. The workers could be confused with the minor workers of Pheidole, but differ in usually being much larger (over 3 mm total length, usually less than 3 mm in Pheidole), and that the antennal club is poorly defined and consists of four segments (well defined in Pheidole and usually consisting of three segments).
Eguchi, Bui and Yamane (2011) - Worker monomorphic; head in full-face view oval or elliptical, often with extremely elongate neck; frontal carina, if distinct, not extending beyond the level of eye in full-face view; antennal scrobe absent; parafrontal ridge or carina(e) often present; median portion of clypeus convex anteriad, sometimes with a shallow emargination at midpoint; posteromedian portion of clypeus moderately or relatively broadly inserted between frontal lobes; masticatory margin with apical and 2 distinct preapical teeth followed by several smaller teeth or denticles; palp formula 5,3 or 4,3; antenna 12-segmented, gradually incrassate toward apex or with an indistinct 4-segmented club; eye medium sized; mesosoma elongate; promesonotum forming a dome; promesonotal suture weakly present or absent dorsally; metanotal groove moderately or strongly impressed dorsally; propodeal spines varying in size and shape (rarely reduced to tiny denticles or rounded angles); propodeal lobe round or subtriangular with blunt angles; petiole consisting of an anterior peduncle and a node (separation between peduncle and node sometimes indistinct); gastral shoulder absent.
The worker of Aphaenogaster is similar to the minor worker of Pheidole (larger species) and the worker of Kartidris and Myrmica. In the minor worker of Pheidole the masticatory margin of the mandible bears 1 or 2 small teeth between the preapical tooth and the 3rd large tooth. In the worker of Kartidris the vertex has a broad depressed area between eyes, the masticatory margin of the mandible has 5 distinct teeth, the antennal club is distinctly 3-segmented, and the propodeum is unarmed. In the worker of Myrmica the promesonotum is only slightly raised and the propodeal lobe is well-developed as a triangular or sharp lamella. In addition, the palp formula is always 6,4 in Myrmica as opposed to 5,3 or 4,3 in Aphaenogaster.
There are a number of Aphaenogaster species groups. These have mainly served as a convenience for organizing revisionary studies that focus on a morphologically similar set of species from a particular region: Aphaenogaster species groups
|See images of species within this genus|
Keys including this Genus
- Key to Ant Genera of the Navajo Reservation
- Key to Australian Genera of Myrmicinae
- Key to North American Genera of Myrmicinae
- Key to Vietnamese Myrmicinae Genera
- Key to the Ant Genera of New Mexico
Keys to Species in this Genus
- Key to Australian Aphaenogaster Species
- Key to US Aphaenogaster species
- Key to Aphaenogaster European testaceopilosa-group workers
- Key to Aphaenogaster workers of Europe
- Key to Aphaenogaster European testaceopilosa-group queens
- Key to Aphaenogaster European testaceopilosa-group males
- Key to Aphaenogaster of Iran
- Key to Mediterranean Aphaenogaster cecconii group species
- Key to Iberian Peninsula Aphaenogaster species
- Key to Aphaenogaster phalangium complex species
- Key to Aphaenogaster of the southwestern Australian Botanical Province
- Key to Aphaenogaster gibbosa group workers
- Key to Arabian Aphaenogaster
- Key to Mediterranean Aphaenogaster graeca complex
Distribution is notably concentrated in the northern hemisphere. In the southern hemisphere, they are extremely sparse in the Neotropical and Afrotropical regions, although relatively rich in the Australian region.
Distribution and Richness based on AntMaps
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), Enspel Formation, Rhineland-Palatinate, Germany (Oligocene), Florissant, Colorado, United States (Late Eocene), Mexican amber, Chiapas, Mexico (Middle Miocene), Quesnel, British Columbia, Canada (Early Miocene?), Rott, Westphalia, Germany (Late Oligocene), Rovno amber (Priabonian, Late Eocene), Sakhalin amber, Ukraine (Thanetian, Paleocene).
Notes on Australian species: The distinctive nests of Australian Aphaenogaster ants are often the first indication of their presence. These nests can be very dense and when in sandy soils, individual entrances can be large, deep cones or bores (up to 4 cm in diameter and 30 cm deep) with large mounds of loose dirt. This style of nest has resulted in these ants being known as "funnel ants." In some cases nests can be so dense and extensive that they severely affect soil structure, resulting in a loose and fragile surface which easily collapses under foot. When this occurs in situations such as golf courses, pastures and unsealed airstrips damage can be severe and these ants can become a serious problem. Although not aggressive, workers will defend their nests when disturbed, emerging from entrances in small numbers to attack intruders.
While nests can contain large numbers of workers, few workers are usually seen on the surface, and then most are found near the entrance; they are rarely seen foraging any distance from nests. It is known that these ants tend aphids on the roots of plants and that arthropod fragments are often found in the upper portions of their nests. It is possible that the tended aphids provide much of the food needed by the nest, and that the funnel-shaped entrances act as traps for surface foraging arthropods. These factors may combine to reduce or eliminate the need to forage on the surface of the ground.
Notes on North American species: These are elongate, slender ants which are very fast and agile in the field. Most species nest in the soil under stones or logs while some of the desert species nest in the soil with the nest entrance surrounded by pebbles. They are carnivorous, and collect dead insects, as well as tend Homoptera or collect nectar. Colonies are moderately large to very large. This is a common genus and occurs in all habitats, but is especially common in forested ecosystems.
Notes on Vietnamese species: Eguchi, Bui and Yamane (2011) - The majority of species inhabit well-developed forests but some occur in sparse forests, dwarf forests and areas with low bushes. Nests are usually found in the soil, under stones and in rotting logs (Bui & Eguchi 2003, Eguchi et al. 2004).
Csősz et al. (2021): Across the distribution of Aphaenogaster in Madagascar, there are species of Camponotus that mimic the general form and behavior of Aphaenogaster: Camponotus imitator, Camponotus jodina, Camponotus karaha and Camponotus longicollis. These Camponotus mimics are often seen foraging with workers of Aphaenogaster, but potential beneﬁts for Camponotus have not been studied.
Association with Other Organisms
All Associate Records for Genus
|Taxon||Relationship||Associate Type||Associate Taxon||Associate Relationship||Locality||Source||Notes|
|Aphaenogaster baronii||host||fungus||Aegeritella maroccana||pathogen||Espadaler & Santamaria, 2012|
|Aphaenogaster fulva||host||ant||Aphaenogaster tennesseensis||temporary parasite|
|Aphaenogaster gibbosa||mutualist||butterfly||Lampides boeticus||Obregon et al. 2015|
|Aphaenogaster picea||host||ant||Aphaenogaster tennesseensis||temporary parasite|
|Aphaenogaster rudis||host||ant||Aphaenogaster tennesseensis||temporary parasite|
|Aphaenogaster tennesseensis||temporary parasite||ant||Aphaenogaster fulva||host|
|Aphaenogaster tennesseensis||temporary parasite||ant||Aphaenogaster picea||host|
|Aphaenogaster tennesseensis||temporary parasite||ant||Aphaenogaster rudis||host|
|Aphaenogaster texana||host||phorid fly||Apocephalus portalensis||parasite||phorid.net||attacked|
|Aphaenogaster treatae||host||cricket||Myrmecophilus pergandei||myrmecophile||United States|
- Antennal segment count: 12
- Antennal club: absent, gradual, 4 weak
- Palp formula: 5,3; 4,3
- Total dental count: 7-16 (-6)
- Spur formula: 1 simple, 1 simple; 0, 0
- Eyes: present
- Scrobes: absent
- Caste: most monomorphic, at least one species polymorphic
- Sting: present
- 2n = 30 (Malaysia) (Goni et al., 1982).
All Karyotype Records for Genus
|Aphaenogaster||30||Malaysia||Goni et al., 1982|
|Aphaenogaster beccarii||30 • 46||India; Malaysia||Imai et al., 1983; Imai et al., 1984|
|Aphaenogaster depilis||34||Tunisia||Hauschteck-Jungen & Jungen, 1983|
|Aphaenogaster famelica||17||34||Japan||Imai & Yosida, 1964; Imai, 1966; Imai, 1969; Imai, 1971|
|Aphaenogaster fulva||36||USA||Crozier, 1977|
|Aphaenogaster gibbosa||11 • 16 • 17||34||4M+2SM+4ST+6T; 6M+4SM+8ST+16T||Switzerland||Hauschteck-Jungen & Jungen, 1983; Lorite et al., 2000; Palomeque et al., 1993b||this karyotype is reported as fusion of two telocentric chromosomes|
|Aphaenogaster iberica||17||34||6M+4SM+8ST+16T||Spain||Palomeque et al., 1993a; Palomeque et al., 1993b; Lorite et al., 2000||ocurrence of supernumery chromosome segment|
|Aphaenogaster japonica||22||Japan||Imai, 1969|
|Aphaenogaster lamellidens||38||USA||Crozier, 1977; Taber & Cokendolpher, 1988|
|Aphaenogaster longiceps||45 • 46||Australia||Imai et al., 1977||a metacentric pair is single and two telocentric corresponds the arms|
|Aphaenogaster miamiana||36||USA||Crozier, 1977|
|Aphaenogaster osimenseis||16||32||Japan||Imai & Yosida, 1964; Imai & Yosida, 1966; Imai, 1966; Imai, 1969; Imai, 1971|
|Aphaenogaster rudis||18 • 20||40 • 41 • 42 • 44||USA||Crozier, 1977||suggested to be due supernumerary chromosome|
|Aphaenogaster sardoa||34||Tunisia||Hauschteck-Jungen & Jungen, 1983|
|Aphaenogaster senilis||16||32||6M+6SM+20ST||Spain||Palomeque et al., 1993a; Palomeque et al., 1993b; Lorite et al., 2000|
|Aphaenogaster smythiesii||11||22 • 34||20SM+2M; 16M+18A||India; Japan||Imai, 1969; Imai, 1971; Imai et al., 1984|
|Aphaenogaster subterranea||11||22||Germany; Switzerland||Hauschteck, 1962; Hauschteck-Jungen & Jungen, 1983|
|Aphaenogaster testaceopilosa||17||34||Croatia; Spain; Tunisia||Hauschteck-Jungen & Jungen, 1983||Lorite and Palomeque 2010 states some issues about the identification/sampling location|
|Aphaenogaster tipuna||34||Taiwan||Hung et al., 1972|
|Aphaenogaster treatae||42||USA||Crozier, 1977|
The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.
- APHAENOGASTER [Myrmicinae: Stenammini]
- Aphaenogaster Mayr, 1853b: 107. Type-species: Aphaenogaster sardoa, by subsequent designation of Bingham, 1903: 270.
- Aphaenogaster junior synonym of Atta: Mayr, 1863: 395.
- Aphaenogaster subgenus of Stenamma: Emery, 1895c: 298.
- Aphaenogaster revived status as genus: Emery, 1908c: 309.
- Aphaenogaster senior synonym of Novomessor: Brown, 1974b: 47.
- Aphaenogaster senior synonym of Attomyrma, Deromyrma, Novomessor, Nystalomyrma, Planimyrma: Smith, D.R. 1979: 1359; Bolton, 1982: 364.
- Aphaenogaster senior synonym of Brunella: Bolton, 1982: 364.
- Aphaenogaster senior synonym of †Sinaphaenogaster: Bolton, 2003: 230, 273.
- ATTOMYRMA [junior synonym of Aphaenogaster]
- Attomyrma Emery, 1915d: 70 [as subgenus of Aphaenogaster]. Type-species: Formica subterranea, by original designation.
- Attomyrma junior synonym of Aphaenogaster: Bolton, 1982: 364.
- BRUNELLA [junior synonym of Aphaenogaster]
- Brunella Forel, 1917: 234. Type-species: Aphaenogaster belti, by monotypy.
- Brunella junior synonym of Atopula: Emery, 1924d: 242; Donisthorpe, 1943f: 629.
- Brunella junior synonym of Aphaenogaster: Bolton, 1982: 364; Bolton, 1994: 106.
- DEROMYRMA [junior synonym of Aphaenogaster]
- Deromyrma Forel, 1913b: 350 [as subgenus of Aphaenogaster]. Type-species: Aphaenogaster (Ischnomyrmex) swammerdami, by original designation.
- [Deromyrma also described as new by Forel, 1913k: 49.]
- Deromyrma junior synonym of Aphaenogaster: Bolton, 1982: 364.
- NYSTALOMYRMA [junior synonym of Aphaenogaster]
- Nystalomyrma Wheeler, W.M. 1916j: 215 [as subgenus of Aphaenogaster]. Type-species: Myrmica longiceps, by original designation.
- Nystalomyrma junior synonym of Aphaenogaster: Bolton, 1982: 364.
- PLANIMYRMA [junior synonym of Aphaenogaster]
- Planimyrma Viehmeyer, 1914d: 604 [as subgenus of Aphaenogaster]. Type-species: Stenamma (Ischnomyrmex) loriai, by original designation.
- Planimyrma junior synonym of Aphaenogaster: Bolton, 1982: 364.
- †SINAPHAENOGASTER [junior synonym of Aphaenogaster]
- †Sinaphaenogaster Zhang, J. 1989: 266 [as subgenus of Aphaenogaster]. Type-species: †Paraphaenogaster shanwangensis, by original designation.
- †Sinaphaenogaster junior synonym of Aphaenogaster: Bolton, 2003: 230, 273.
Unpublished evidence has suggested Aphaenogaster is not monophyletic. A molecular phylogenetic study is needed to resolve this problem.
- Aupanun, S., Kulsarin, J., Jaitrong, W. Ito, F. 2018. Colony composition and nesting habits of six species of Aphaenogaster in Thailand (Hymenoptera; Formicidae). Asian Myrmecology 10: e010003 (DOI 10.20362/am.010003).
- Bingham, C. T. 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. London: Taylor and Francis, 506 pp. (page 270, Type-species: Aphaenogaster sardoa; by subsequent designation)
- 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 364, Aphaenogaster senior synonym of Attomyrma, Deromyrma, Novomessor, Nystalomyrma, Planimyrma, and Brunella)
- Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 229, Aphaenogaster as genus)
- Brown, W. L., Jr. 1949a. Synonymic and other notes on Formicidae (Hymenoptera). Psyche (Camb.) 56: 41-49.
- 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 178,180, 183, 184, Aphaenogaster provisional senior synonym of Attomyrma, Deromyrma, Nystalomyrma and Planimyrma)
- Brown, W. L., Jr. 1974b. Novomessor manni a synonym of Aphaenogaster ensifera (Hymenoptera: Formicidae). Entomol. News 85: 45-47 (page 47, Aphaenogaster senior synonym of Novomessor)
- Chapman, J. W.; Capco, S. R. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia. Monogr. Inst. Sci. Technol. Manila 1: 1-327 (page 131, Aphaenogaster in Myrmicinae, Pheidolini)
- Crawley, W. C. (1922) Notes on some Australian ants. Biological notes by E. B. Poulton, D.Sc., M.A., F.R.S., and notes and descriptions of new forms by W. C. Crawley, B.A., F.E.S., F.R.M.S. [concl.]. Entomologist's Monthly Magazine, 58: 121–126.
- Cresson, E. T. 1887. Synopsis of the families and genera of the Hymenoptera of America, north of Mexico, together with a catalogue of the described species, and bibliography. Trans. Am. Entomol. Soc., Suppl. Vol. 1887: 1-351 (page 260, Aphaenogaster in Myrmicinae [Myrmicidae])
- Csősz, S., Loss, A.C., Fisher, B.L. 2021. Taxonomic revision of the Malagasy Aphaenogaster swammerdami group (Hymenoptera: Formicidae). PeerJ 9, e10900 (doi:10.7717/peerj.10900).
- Dalla Torre, K. W. von. 1893. Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann, 289 pp. (page 98, Aphaenogaster in Myrmicinae)
- 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, Aphaenogaster in Myrmicinae, Pheidolini [Pheidolidae])
- Emery, C. 1895d. Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. (Schluss). Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 8: 257-360 (page 298, Aphaenogaster subgenus of Stanamma)
- Emery, C. 1895l. Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 8: 685-778 (page 769, Aphaenogaster in Myrmicinae, Myrmicini; Aphaenogaster as subgenus of Stenamma)
- Emery, C. 1908d. Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes. (Hym.) (Fortsetzung.) III. Die mit Aphaenogaster verwandte Gattungengruppe. Dtsch. Entomol. Z. 1908: 305-338 (page 309, Aphaenogaster revived status as genus)
- Emery, C. 1914b. Cephalotes et Cryptocerus. Le type du genre Crematogaster. Ann. Soc. Entomol. Belg. 58: 37-39 (page 40, Aphaenogaster in Myrmicinae, Pheidolini)
- Emery, C. 1921c. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [part]. Genera Insectorum 174A:1-94 94: 1-94 + 7 (page 55, Aphaenogaster in Myrmicinae, Pheidolini [subtribe Stenammini])
- Emery, C.; Forel, A. 1879. Catalogue des Formicides d'Europe. Mitt. Schweiz. Entomol. Ges. 5: 441-481 (page 461, Aphaenogaster in Myrmicinae [Myrmicidae])
- Enzmann, J. 1947b. New forms of Aphaenogaster and Novomessor. J. N. Y. Entomol. Soc. 55: 147-152 (page 147, Aphaenogaster in Myrmicinae, Aphaenogastrini)
- Forel, A. 1891c. Les Formicides. [part]. In: Grandidier, A. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (28e fascicule). Paris: Hachette et Cie, v + 237 pp. (page 166, Aphaenogaster in Myrmicinae, Myrmicini)
- Forel, A. 1895b. A fauna das formigas do Brazil. Bol. Mus. Para. Hist. Nat. Ethnogr. 1: 89-139 (page 129, Aphaenogaster in Myrmicinae, Myrmicini)
- Forel, A. 1899e. Formicidae. [part]. Biol. Cent.-Am. Hym. 3: 57-80 (page 58, Aphaenogaster in Myrmicinae, Myrmicini)
- Forel, A. 1903a. Les Formicides de l'Empire des Indes et de Ceylan. Part X. J. Bombay Nat. Hist. Soc. 14: 679-715 (page 693, Aphaenogaster subgenus of Stanamma)
- Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 241, Aphaenogaster in Myrmicinae, Pheidolini)
- Hitchcock, B. E. (1958) The funnel ant. Cane Growers Quart. Bull. (Queensland), 21: 104–105.
- Hitchcock, B. E. (1962) Funnel ant flights in 1961. Cane Growers Quart. Bull. (Queensland), 26: 65–66.
- Hitchcock, B. E. (1968) Funnel ant control in Queensland cane fields. Int Soc Sugar Cane Technol Proc Congr, 1 . pp. 1389–1396.
- Kempf, W. W. 1972b. Catálogo abreviado das formigas da regia~o Neotropical. Stud. Entomol. 15: 3-344 (page 22, Aphaenogaster in Myrmicinae, Myrmicini (anachronism))
- Mayr, G. 1853c. Beiträge zur Kenntniss der Ameisen. Verh. Zool.-Bot. Ver. Wien 3: 101-114 (page 107, Aphaenogaster as genus)
- Mayr, G. 1855. Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate aufgefundenen Ameisen, nebst Hinzufügung jener in Deutschland, in der Schweiz und in Italien vorkommenden Arten. Verh. Zool.-Bot. Ver. Wien 5: 273-478 (page 466, Aphaenogaster in Myrmicinae [Myrmicidae])
- Mayr, G. 1863a. Formicidarum index synonymicus. Verh. K-K. Zool.-Bot. Ges. Wien 13: 385-460 (page 395, Aphaenogaster as junior synonym of Atta)
- Murray, D. A. H. (1982) Insecticidal control of funnel ants in turf. Qld J. Agric. Anim. Sci. 39: 147–148.
- Nalini, T., Jayanthi, R. 2020. First record on interaction between ants, other insect visitors and extrafioral nectaries (EFNS) in selected crops from India. Plant Archives 20, 2105-2113.
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- Santschi, F. 1932d. Résultats scientifiques du voyage aux Indes orientales néerlandaises de LL. AA. RR. le Prince et la Princesse Léopold de Belgique. Hymenoptera. Formicidae. Mém. Mus. R. Hist. Nat. Belg. 4: 11-29 (page 13, Aphaenogaster in Myrmicinae, Myrmicini)
- Saunders, G. W. (1961) Defeating the funnel ant. Queensland Agr. J. 87: 751–753.
- Saunders, G. W. (1967) Funnel ants (Aphaenogaster spp., Formicidae) as pasture pests in North Queensland: I. Ecological background, status and distribution. Bulletin of Entomological Research, 57: 419–432.
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- Saunders, G. W. (1970) An investigation of funnel ants (Aphaenogaster spp: Hym., Formicidae) as pasture pests in North Queensland. (Australia). J. Aust. Inst. Agr. Sci. 36: 54–55.
- Shattuck,S.O. (2008) Australian ants of the genus Aphaenogaster (Hymenoptera: Formicidae). Zootaxa, 1677: 25–45.
- Wheeler, W. M. (1916) The Australian ants of the genus Aphaenogaster Mayr. Transactions of the Royal Society of South Australia, 40: 213–223.
- Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 140, Aphaenogaster in Myrmicinae, Myrmicini)
- Wheeler, W. M. 1915i . The ants of the Baltic Amber. Schr. Phys.-Ökon. Ges. Königsb. 55: 1-142 (page 53, Aphaenogaster in Myrmicinae, Myrmicini)
- 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 661, Aphaenogaster in Myrmicinae, Pheidolini)
- Wilson, G. (1962) Progress report on funnel ant control. Proceedings of the Queensland Society for Sugar Cane Technology, 29: 127–130.