Hypoponera punctatissima

A widespread ponerine tramp-species, this small ant can be found nesting in settings that vary from litter, soil, and rotten wood in forests to disturbed sites such as gardens and fields. It is polygynous and polydomous. Two types of queen are present, winged and apterous. Males are also dimorphic, they may be large, apterous and dark brown, or small, apterous and yellow (Japanese Ant Image Database).

Identification
Reddish yellow to dark brown; mesosoma and gaster thickly pubescent, finely and closely punctured. Antennae with 12 segments gradually broadening to an indefinite club; scapes do not reach posterior border of head. Frontal furrow continued as a fine line to near occipital margin. Eyes minute, set forward close to mandibular insertions. Mandibles with 3-4 strong teeth towards apex and numerous smaller denticles posteriorly. Ventral lobe of petiole simple without tooth-like process: Length: 2.5-3.2 mm. (Collingwood 1979)

A member of the punctatissima group.

Distribution
Bolton and Fisher (2011) - H. punctatissima is without doubt the world’s most accomplished ponerine tramp-species. Its range incorporates all tropical and subtropical zoogeographical regions, including most oceanic islands, and it also penetrates well into the temperate zones of both hemispheres where it is frequently synanthropic; for a brief synopsis of world distribution see Delabie & Blard (2002); European distribution is summarised in Seifert (2003).

It is a common introduced species in south and central Florida. First published Florida record: Smith 1933. (Deyrup, Davis & Cover, 2000.)

Distribution based on Regional Taxon Lists
Afrotropical Region: Angola, Benin, Botswana, Cape Verde, Central African Republic, Comoros, Democratic Republic of Congo, Ethiopia, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Nigeria, Rwanda, Saint Helena, Saudi Arabia, Socotra Archipelago, South Africa, Sudan, São Tomé & Principe, Togo, Uganda, United Arab Emirates, United Republic of Tanzania, Yemen, Zambia, Zimbabwe. Australasian Region: Australia, New Caledonia, New Zealand, Norfolk Island. Indo-Australian Region: Fiji, Guam, Hawaii, Indonesia, Marshall Islands, Micronesia (Federated States of), New Guinea, Niue, Northern Mariana Islands, Palau, Philippines, Samoa, Solomon Islands, Tonga, Wallis and Futuna Islands. Malagasy Region: Madagascar, Mauritius, Mayotte, Réunion, Seychelles. Nearctic Region: Canada, United States. Neotropical Region: Bahamas, Barbados, Bermuda, Costa Rica, Cuba, Dominican Republic, Galapagos Islands, Greater Antilles, Guadeloupe, Guatemala, Haiti, Martinique, Mexico, Netherlands Antilles, Paraguay, Puerto Rico, Saint Lucia, Saint Vincent and the Grenadines, United States Virgin Islands. Oriental Region: Sri Lanka. Palaearctic Region: Andorra, Armenia, Austria, Balearic Islands, Belgium, Bulgaria, Canary Islands, China, Czech Republic, Denmark, Finland, France, Germany , Greece, Hungary, Iberian Peninsula, Ireland, Israel, Italy, Japan, Luxembourg, Malta, Morocco, Netherlands, Norway, Oman, Poland, Portugal, Romania, Russian Federation, Slovakia, Spain, Sweden, Switzerland, Tunisia, Ukraine, United Kingdom of Great Britain and Northern Ireland.

Africa
Bolton and Fisher (2011) - H. punctatissima is very widely distributed in sub-Saharan Africa and appears relatively common. It is often retrieved from litter and topsoil samples in forests, and also from rotten wood, but it seems particularly frequent in areas disturbed by human activities, especially in gardens, plantations and crop fields.

Florida
Deyrup, Davis and Cover (unpublished) - This species occurs in small colonies in a variety of moist or well-drained habitats. In southern Florida it is often found in rotten logs and large grass tussocks, and especially in accumulations of organic matter, such as grass clippings or compost, often in open or disturbed sites. The key to the housefly-like success of H. punctatissima lies in its predilection for dung and various kinds of middens, which humans and their livestock supply in inexhaustible quantities. Delabie and Bland (2002) have compiled a whole series of breeding sites, ranging from earthworm cultures to heaps of chicken manure, and suggest that this ant emerged out of Asia following the domestication and extensive use of horses.

The tendency for H. punctatissima to produce disproportionately large numbers of dispersing females is appropriate for a species that colonizes resource-rich sites that are often ephemeral. This dispersal, in turn, leads to problems with stinging alates. Queens often fly in large numbers, stinging when they land on human skin if they are touched, trapped under clothing, or stuck in sweat. We have had a number of complaints about this species, mostly from west-central Florida (Deyrup, Davis & Cover, 2000).

New Mexico
Mackay and Mackay (2002) - Habitat. Riparian zones in arid habitats. Biology. This species nests under stones or in the soil.

Fennoscandia, Denmark and the British Isles
This species is often imported with plant material. However, it has long been resident in North Europe and head capsules presumed to be of this species have been recorded from sewage mud deposited about 1500 years ago in North England. Most recorded occurrences are from heated premises such as bakehouses and conservatories. However, colonies have been recorded outside in England, Ireland, Denmark, Norway and Finland from fermenting rubbish dumps, waste tips, sawdust heaps and deep mines away from buildings. Queens and sometimes workers have also been captured individually by general herbage sweeping or in woodlands. Occurrences in Denmark and Fennoscandia have been summarised by Skott (1971). Colonies are often populous and many alate queens may be produced to fly out during August and September. The apterous males remain in the nest. This species, as with most Ponerini, is mainly carnivorous on small arthropods (Collingwood 1979).

Yemen
Sharaf et al. (2017) - This genus is recorded for the first time from Socotra; based on its distribution, this species has probably been introduced. This species was collected from moist soil and leaf litter under mango and date palm trees where the soil was rich in decaying organic material. It was also observed nesting under a stone next to a dragon blood tree. This species was also nesting directly in moist soil under a dead palm tree where organic materials are found. Another nest was found among old moist leaf sheaths surrounding the trunk base of a date palm tree.

Castes
Bolton and Fisher (2011) - As well as the usual workers and alate queens, it also produces ergatoid queens and dimorphic large and small ergatoid males, but never the usual alate male form. Yamauchi et al. (1996) studied reproductive behavior in a japanese population

Nomenclature

 *  punctatissima. Ponera punctatissima Roger, 1859: 246, pl. 7, fig. 7 (w.q.) POLAND and GERMANY. Combination in Hypoponera: Taylor, 1967a: 12. Senior synonym of androgyna: Emery & Forel, 1879: 455, Seifert, 2003c: 69; of tarda: Dalla Torre, 1893: 41, Seifert, 2003c: 69; of kalakauae, mina, mumfordi: Wilson & Taylor, 1967: 29 (in text); of ergatandria: Smith, D.R. 1979: 1343; of mina: Taylor, 1987a: 30; of exacta: Atanassov & Dlussky, 1992: 71, Seifert, 2003c: 69; of jugata: Seifert, 2003c: 69; of aemula, argonautorum, bondroiti, breviceps, brevis, cognata, durbanensis, incisa, mesoepinotalis, petri, schauinslandi, sordida, sulcitana, ursoidea: Bolton & Fisher, 2011: 87. Current subspecies: nominal plus indifferens. See also: Emery, 1916b: 110; Wheeler, W.M. 1937c: 59; Collingwood, 1979: 30.
 * androgyna. Ponera androgyna Roger, 1859: 246 (footnote) (ergatoid m.) GERMANY. Junior synonym of punctatissima: Emery & Forel, 1879: 455; Seifert, 2003c: 69.
 * argonautorum. Ponera argonautorum Arnol'di, 1932b: 66, figs. 6, 7 (q.) RUSSIA. Combination in Hypoponera: Bolton, 1995b: 213. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * breviceps. Ponera breviceps Bernard, 1953b: 202, fig. 3 (q.) (m. excluded, see Bolton & Fisher, 2011: 88) GUINEA. Combination in Hypoponera: Bolton, 1995b: 213. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * brevis. Ponera brevis Santschi, 1921c: 113, fig. 1 (w.) BENIN. Combination in Hypoponera: Bolton, 1995b: 213. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * cognata. Ponera ergatandria st. cognata Santschi, 1912b: 153 (w.) ANGOLA. [Unresolved junior primary homonym of Ponera cognata Emery, 1896g: 56 (now in Pachycondyla).] Bernard, 1953b: 202 (q.). Combination in Hypoponera: Bolton, 1995b: 213. Raised to species: Bernard, 1953b: 202. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87. See also: Arnold, 1915: 81.
 * petri. Ponera ergatandria r. petri Forel, 1916: 397 (ergatoid q., not w.) DEMOCRATIC REPUBLIC OF CONGO. Combination in Hypoponera: Bolton, 1995b: 216. Raised to species: Santschi, 1938b: 78. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * incisa. Ponera incisa Santschi, 1914d: 320, fig. 8 (ergatoid m., not w.) NIGERIA. Combination in Hypoponera: Bolton, 1995b: 214. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * mina. Ponera mina Wheeler, W.M. 1927i: 131, fig. 2 (w.q. ergatoid m.) AUSTRALIA. Combination in Hypoponera: Taylor & Brown, D.R. 1985: 31. Junior synonym of gleadowi: Wilson, 1958d: 329. Revived from synonymy: Taylor & Brown, D.R. 1985: 31. Junior synonym of punctatissima: Wilson & Taylor, 1967: 29 (in text); Taylor, 1987a: 30. See also: Wheeler, W.M. 1937c: 59.
 * mesoepinotalis. Ponera mesoepinotalis Weber, 1942a: 44, fig. 4 (w.) SUDAN. Combination in Hypoponera: Bolton, 1995b: 215. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * mumfordi. Ponera mumfordi Wheeler, W.M. 1933f: 141 (w.) FRENCH POLYNESIA (Marquesas Is). Wheeler, W.M. 1936f: 4 (q.). Junior synonym of gleadowi: Wilson, 1958d: 328; of punctatissima: Wilson & Taylor, 1967: 29 (in text).
 * exacta. Ponera punctatissima var. exacta Santschi, 1923a: 134 (w.) TUNISIA. Junior synonym of punctatissima: Atanassov & Dlussky, 1992: 71; Seifert, 2003c: 69.
 * jugata. Ponera punctatissima r. jugata Forel, 1892l: 251 (q.) MADAGASCAR. Combination in Hypoponera: Bolton, 1995b: 215. Raised to species: Dalla Torre, 1893: 39. Subspecies of punctatissima: Emery, 1899f: 268. Junior synonym of punctatissima: Seifert, 2003c: 69.
 * sordida. Ponera ragusai var. sordida Santschi, 1914b: 54 (w.) KENYA. Combination in Hypoponera: Bolton, 1995b: 216. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * durbanensis. Ponera sulcatinasis r. durbanensis Forel, 1914d: 213 (w.) SOUTH AFRICA. Arnold, 1926: 206 (q.). Combination in Hypoponera: Bolton, 1995b: 214. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * sulcitana. Ponera sulcitana Stefani, 1970: 1, figs. 1-12 (w.q. ergatoid m., l.) ITALY. Combination in Hypoponera: Bolton, 1995b: 216. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * tarda. Ponera tarda Charsley, 1877: 162 (w.q.) GREAT BRITAIN. Junior synonym of punctatissima: Dalla Torre, 1893: 41; Seifert, 2003c: 69.
 * ursoidea. Ponera ursoidea Bernard, 1953b: 203, fig. 3 (ergatoid q., not w.) GUINEA. Combination in Hypoponera: Bolton, 1995b: 216. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87.
 * ergatandria. Ponera ergatandria Forel, 1893g: 365 (w.q. ergatoid m.) ST VINCENT & THE GRENADINES (St Vincent I.).
 * Type-material: syntype workers, syntype queens (numbers not stated), 1 syntype ergatoid male.
 * Type-localities: St Vincent & the Grenadines: St Vincent I., 41a, 41b, 41c, and 41d (H.H. Smith).
 * [Note: 41a = Bowwood Valley, nr Kingston, 800 ft, 15.x., 41b = Islet fronting Châteaubelais Bay, 31.v., 41c = Richmond Estate, 31.x., 41d = Villa Estate, south end of island, 14.x.]
 * Type-depositories: BMNH, MHNG.
 * Combination in Hypoponera: Kempf, 1972a: 122.
 * Status as species: Emery, 1896g: 54 (in key); Forel, 1899c: 16; Wheeler, W.M. 1905b: 82, 121; Forel, 1907a: 2; Wheeler, W.M. 1908a: 125; Wheeler, W.M. 1908e: 405; Wheeler, W.M. 1910g: 561; Emery, 1911d: 92; Wheeler, W.M. 1913b: 481; Wheeler, W.M. 1913d: 239; Santschi, 1914b: 53; Forel, 1914d: 213; Wheeler, W.M. & Mann, 1914: 13; Wheeler, W.M. 1922a: 781; Menozzi, 1927c: 267; Menozzi, 1929a: 2; Menozzi & Russo, 1930: 151; Menozzi, 1931b: 261; Weber, 1934a: 23; Smith, M.R. 1936: 425; Wheeler, W.M. 1936b: 195; Smith, M.R. 1937: 825; Wheeler, W.M. 1937c: 59; Wheeler, W.M. 1938: 251; Weber, 1948b: 79; Creighton, 1950a: 48; Smith, M.R. 1951a: 786; Smith, M.R. 1958c: 112; Bernard, 1967: 89 (redescription); Kempf, 1972a: 122; Alayo, 1974: 7 (in key); Bolton, 1995b: 214; Seifert, 2013: 198; Borowiec, L. 2014: 82; Lebas, et al. 2016: 404; Seifert, 2018: 149.
 * Junior synonym of punctatissima: Taylor, 1968a: 65; Smith, D.R. 1979: 1343; Bolton & Fisher, 2011: 86; Radchenko, 2016: 72; Lubertazzi, 2019: 123.
 * aemula. Ponera dulcis var. aemula Santschi, 1911c: 351 (w.) TANZANIA. Combination in Hypoponera: Bolton, 1995b: 213. Junior synonym of schauinslandi: Seifert, 2003c: 69; of punctatissima: Bolton & Fisher, 2011: 87; of ergatandria: Seifert, 2013: 198.
 * bondroiti. Ponera ergatandria subsp. bondroiti Forel, 1911c: 285 (w.q. ergatoid m.) BELGIUM. Combination in Hypoponera: Onoyama, 1989b: 2. Raised to species: Santschi, 1937h: 364. Junior synonym of schauinslandi: Seifert, 2003c: 69; of punctatissima: Bolton & Fisher, 2011: 87; of ergatandria: Seifert, 2013: 198.
 * kalakauae. Ponera kalakauae Forel, 1899a: 115 (ergatoid q. (not w.), q.) HAWAII. Junior synonym of gleadowi: Wilson, 1958d: 328; of punctatissima: Wilson & Taylor, 1967: 29 (in text); of ergatandria: Seifert, 2013: 198.
 * schauinslandi. Ponera punctatissima subsp. schauinslandi Emery, 1899d: 439 (q.) HAWAII. Junior synonym of punctatissima: Wilson & Taylor, 1967: 28. Revived from synonymy and raised to species: Seifert, 2003c: 69. Junior synonym of punctatissima: Bolton & Fisher, 2011: 87; of ergatandria: Seifert, 2013: 198.

As noted above, this species has accumulated a large number of synonyms.

Worker
Bolton and Fisher (2011) - Measurements: HL 0.56–0.72, HW 0.46–0.60, HS 0.515–0.660, SL 0.35–0.48, PrW 0.33–0.43, WL 0.70–0.90, HFL 0.36–0.48, PeNL 0.14–0.20, PeH 0.30–0.39, PeNW 0.22–0.29, PeS 0.223–0.290 (60 measured). Indices: CI 79–87, SI 75–84, PeNI 63–74, LPeI 43–53, DPeI 140–165.

Eyes small but conspicuous, of 1–6 often poorly defined or partially fused ommatidia, located far forward on the side of the head. Impression that extends back along midline of head from the frontal lobes reaches at least the midlength of the vertex and usually further. Apex of scape, when laid straight back from its insertion, usually fails to reach or less commonly just touches the midpoint of the posterior margin in full-face view; SL/HL 0.62–0.70. Minute punctulate sculpture of cephalic dorsum extremely fine and superficial. Mesonotal-mesopleural suture absent. Metanotal groove deeply incised across dorsum of mesosoma; mesonotum with a well-defined posterior margin. Propodeal declivity rounds into the sides, without sharp marginations or carinae. In profile the anterior margination of the mesopleuron rounded to very bluntly angular behind base of anterior coxa; without a prominent sharp angle or tooth. Mesopleuron smooth and shining, at most with a few scattered punctures; usually with a small reticulate patch in the extreme posteroventral corner above the mesocoxa. Petiole in profile with the anterior and posterior faces of the node weakly convergent dorsally; node longer just above the anterior tubercle than at the dorsum. Sternite of petiole in profile a rounded lobe, without angles anteriorly or posteriorly. Maximum width of first gastral tergite in dorsal view less than the width of the second tergite at its midlength. Base of cinctus of second gastral tergite smooth and shining in dorsal view, without cross-ribs. Posttergite of second gastral segment, from posterior margin of cinctus to apex, much broader than long. Disc of second gastral tergite minutely punctulate, appearing weakly microreticulate in places. Full adult colour varies from dull brownish yellow to very dark brown or almost black. In general, workers at the bottom end of the size range given above tend to be lighter and the colour becomes darker with increasing size, but this is by no means an entirely consistent rule.

Queen
Fusca, pube adpressa cinereo-micans, ore, antennis pedibus et apice abdominis pallide rufis, oculis margini anteriori capitis proximis. - Long. 3 – 3 1/2 Mm.

Bolton and Fisher (2011) - True queens are alate, with much larger eyes (obviously >50 ommatidia) that have short setae projecting between the ommatidia, and have a full complement of flight sclerites on the mesosoma. Ergatoid queens are very worker-like but have larger eyes (usually 10 to about 20 ommatidia) and a developed mesonotal-mesopleural suture that is absent in the worker.

Male
Bolton and Fisher (2011) - Ergatoid males fall into major (larger, brown and with small eyes present) and minor (smaller, yellow and eyeless) categories (Yamauchi, et al. 1996) and are again very worker-like, especially in head structure, but have shorter scapes (SI 68–72), only 12-segmented antennae (as do workers, intercastes and queens), and of course fully developed but retractile male genitalia. Alate males have never been seen and it is probably now safe to say are never produced.

Type Material
Bolton and Fisher (2011):
 * Syntype workers and queen, POLAND: Rauden (now Rudy, Opole Prov.) and GERMANY: Berlin, in hothouses (Roger) (MNHN) [examined].
 * Ponera androgyna: Syntype ergatoid males (not workers), POLAND: Rauden (now Rudy, Opole Prov.) (Roger) (ZMHB) [not seen].
 * Ponera tarda: Syntype workers and queen, GREAT BRITAIN: Oxford, 1877 (R.S. Charsley) (OXUM) [not seen].
 * Holotype queen'': MADAGASCAR: Prov. d’Imerina (Sikora) (MHNG) [not seen].
 * Ponera ergatandria: Syntype workers, queens and ergatoid male, SAINT VINCENT I. (Antilles): 41b (H.H. Smith). (MHNG, BMNH) [BMNH ergatoid male examined].
 * Ponera kalakauae: Syntype worker-queen intercaste (not worker) and queen, HAWAIIAN IS: Kauai I., Lahue, 2000 ft, vii.1896 (R.C.L. Perkins) (intercaste), and Honolulu (Oahu I.), xi.1896 (R.C.L. Perkins) (queen) (BMNH) [examined].
 * Syntype queens, HAWAIIAN IS: Laysan I. (now Laycan I.) (Schauinsland) (MSNG) [not seen].
 * Ponera ergatandria subsp. bondroit: Syntype workers, queen and ergatoid male, BELGIUM: Bruxelles, Jardin Bot., xi.1909, 5.xi.1909, 15.xi.1909 (Bondroit) (MHNG) [not seen].
 * Ponera dulcis var. aemula: Lectotype and paralectotype workers (designated by Seifert, 2003: 68), TANZANIA: Kilimandjaro, zone des cultures, Kiboscho, 1400 m, 1904 (Ch. Alluaud) (NHMB) [examined].
 * Ponera ergatandria st. cognate: Syntype workers, ANGOLA: Benguela, Cucula (J. Cruchet) (NHMB) [examined].
 * Ponera ragusai var. sordida: Holotype worker, KENYA: région côtière, Shimoni, st. no. 9, xi.1911 (Alluaud & Jeannel) (NHMB) [examined].
 * Ponera incise: Holotype ergatoid male (not worker), NIGERIA: Lagos (F. Silvestri) (not in NHMB, presumably in DEUN) [not seen].
 * Ponera sulcatinasis r. durbanensis: Syntype workers, SOUTH AFRICA: Natal, Durban, 16.i.1914, no. 323 (G. Arnold) (MHNG) [examined].
 * Ponera ergatandria r. petri: Holotype worker-queen intercaste (not worker), DEMOCRATIC REPUBLIC OF CONGO: St Gabriel (Kohl) (MHNG) [examined].
 * Ponera brevis: Holotype worker, BENIN (Dahomey on label): Porto Novo (Le Moult) (NHMB) [examined].
 * Ponera punctatissima var. exacta: Syntype workers, TUNISIA: Hammamet (Santschi) (NHMB, BMNH) [BMNH syntype examined].
 * Ponera mina: Syntype workers, queens and ergatoid male, AUSTRALIA: Norfolk I., 1915 (A.M. Lea) (MCZC and possibly SAMA) [not seen].
 * Ponera argonautorum: Holotype queen, RUSSIA: Black Sea Coast, Anapa, 26.viii.1925, Nr. 422 (K.V. Arnol’di) (probably in ZMUM) [not seen].
 * Ponera mumfordi: Syntype workers, FRENCH POLYNESIA: Marquesas Is, Uapou; Kohepu (Tekohepu) summit, alt. 3000 ft, 3.xi.1931 (LeBronnec) (probably in MCZC) [not seen].
 * Ponera mesoepinotalis: Holotype worker, SUDAN: Imatong Mts, 2.viii.1939, 6400 ft, no. 1395 (N.A. Weber) (not in MCZC, presumed lost).
 * Ponera breviceps: LECTOTYPE dealate queen (by present designation) and two paralectotype dealate queens, GUINEA: Nimba (Lamotte) (MNHN) [examined].
 * Ponera ursoidea: LECTOTYPE worker-queen intercaste (not worker) (by present designation), GUINEA: Crête de Nion, 1300 m. (no collector’s name, probably Lamotte) (MNHN) [examined].
 * Ponera sulcitana: Holotype worker; paratype workers, queens and ergatoid male, ITALY: Sardinia, Grotta dei Fiori presso Carbonia (A. Serra) (IZUC) [not seen].

References based on Global Ant Biodiversity Informatics

 * Arnold G. 1926. A monograph of the Formicidae of South Africa. Appendix. Annals of the South African Museum. 23: 191-295.
 * Bernard F. 1953. La réserve naturelle intégrale du Mt Nimba. XI. Hyménoptères Formicidae. Mémoires de l'Institut Français d'Afrique Noire 19: 165-270.
 * Bolton B. and B. L. Fisher. 2011. Taxonomy of Afrotropical and West Palaearctic ants of the ponerine genus Hypoponera Santschi (Hymenoptera: Formicidae). Zootaxa 2843: 1-118
 * Bolton, B., and B. L. Fisher. "Taxonomy of Afrotropical and West Palaearctic ants of the ponerine genus Hypoponera Santschi (Hymenoptera: Formicidae)." Zootaxa 2843 (2012): 1-118.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Delabie J. H. C., and F. Blard. 2002. The Tramp Ant Hypoponera punctatissima (Roger) (Hymenoptera: Formicidae: Ponerinae): New Records from the Southern Hemisphere. Neotropical Entomology 31(1): 149-151.
 * Emery C. 1911. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125.
 * Forel A. 1914. Formicides d'Afrique et d'Amérique nouveaux ou peu connus. Bulletin de la Société Vaudoise des Sciences Naturelles 50: 211-288.
 * Forel A. 1916. Fourmis du Congo et d'autres provenances récoltées par MM. Hermann Kohl, Luja, Mayné, etc. Revue Suisse de Zoologie 24: 397-460.
 * Garcia F.H., Wiesel E. and Fischer G. 2013.The Ants of Kenya (Hymenoptera: Formicidae)Faunal Overview, First Species Checklist, Bibliography, Accounts for All Genera, and Discussion on Taxonomy and Zoogeography. Journal of East African Natural History, 101(2): 127-222
 * IZIKO South Africa Museum Collection
 * Koen J. H., and W. Breytenbach. 1988. Ant species richness of fynbos and forest ecosystems in the Southern Cape. South Afr. Tydskr. Dierk. 23(3): 184-188.
 * Medler J. T. 1980: Insects of Nigeria - Check list and bibliography. Mem. Amer. Ent. Inst. 30: i-vii, 1-919.
 * Nwosu L. C., and I. A. Lawal. 2010. Assesment of insect invaders of decaying banana and plantain (Musa spp.) pseudo stems in Umuagwo, Ohaji-Egbema local government area, Imo state, Nigeria. Journal of Tropical Agriculture, Food, Environment and Extension 9(2): 84-89.
 * Ross S. R. P. J., F. Hita Garcia, G. Fischer, and M. K. Peters. 2018. Selective logging intensity in an East African rain forest predicts reductions in ant diversity. Biotropica 1-11.
 * Santschi F. 1912. Fourmis d'Afrique et de Madagascar. Annales de la Société Entomologique de Belgique 56: 150-167.
 * Santschi F. 1914. Formicides de l'Afrique occidentale et australe du voyage de Mr. le Professeur F. Silvestri. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 8: 309-385.
 * Santschi F. 1933. Contribution à l'étude des fourmis de l'Afrique tropicale. Bulletin et Annales de la Société Entomologique de Belgique. 73: 95-108.
 * Santschi F. 1938. Notes sur quelques Ponera Latr. Bulletin de la Société Entomologique de France 43: 78-80.
 * Taylor B. 1976. Ants of the Nigerian Forest Zone (Hymenoptera: Formicidae). I. Ponerinae, Cerapachyinae, Pseudomyrmecinae. Cocoa Research Institute of Nigeria Technical Bulletin Series 4: 1-41.
 * Weber N. A. 1942. New doryline, cerapachyine and ponerine ants from the Imatong Mountains, Anglo-Egyptian Sudan. Proc. Entomol. Soc. Wash. 44: 40-49.
 * Weber N. A. 1943. The ants of the Imatong Mountains, Anglo-Egyptian Sudan. Bulletin of the Museum of Comparative Zoology 93: 263-389.
 * Wetterer J. K., X. Espadaler, A. L. Wetterer, D. Aguin-Pombo, and A. M. Franquinho-Aguiar. 2006. Long-term impact of exotic ants on the native ants of Madeira. Ecological Entomology 31: 358-368.
 * Wetterer J. K., X. Espadaler, A. L. Wetterer, D. Aguin-Pombo, and A. M. Franquinho-Aguiar. 2007. Ants (Hymenoptera: Formicidae) of the Madeiran archipelago. Sociobiology 49: 265-297.
 * Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bulletin of the American Museum of Natural History 45: 711-1004