Dorylus orientalis

My Ceylon collection was made in disturbed rain forest in the Udawaddatekele Sanctuary (Garden of the Kandy kings), at Kandy. Workers were found dispersed in the soil to a depth of at least 12.5 cm. None were found above ground. Our limited records indicate that the species is generally subterranean and occurs in a wide range of habitats, including cultivated land. (Wilson 1964) Heterick & Kitching (2022) collected this species in a pitfall trap within a lowland dipterocarp forest in Brunei.

Identification
A member of the Dorylus orientalis-group.

Wilson (1964) - A single series from Kandy, Ceylon varied in HW 0.55-1.37mm, or as much as all the other series together. Antenna always 9-segmented. The species is only moderately polymorphic. As in other Oriental Dorylus, the mandibular dentition shows great variation which is in part allometric. Concolorous yellowish brown.

Eguchi et al. (2014): The mode of polymorphism observed has the following interesting aspects: (1) workers are clearly subdivided into two series by a set of qualitative characters; (2) the “typical series” is numerically much more dominant than the “atypical series” (the latter occupied less than 1% of the whole of the workers collected); (3) a wider size variation was observed in the former (HW, 0.48–1.41 mm; ML, 0.42–1.12 mm) than in the latter (HW, 0.44–1.13 mm; ML, 0.35–0.79 mm); and (4) within the atypical series, smaller workers are numerically much dominant. The atypical worker mentioned above is characterized by a set of the following features: (1) head narrowed anteriorly, (2) median portion of clypeus strongly projecting anteriad, and (3) antenna 8-segmented. The word “typical” used means “to agree well with the habitus of the major worker (Meitan, Kweichow, G. Liu leg.) which was determined as D. orientalis by Wilson (1964: 442; see also Figs. 1–4)

Distribution based on Regional Taxon Lists
Indo-Australian Region: Borneo, Brunei Darussalam, Indonesia, Malaysia. Oriental Region: Bangladesh, India, Laos, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand, Vietnam. Palaearctic Region: China.

Biology
Roonwal (1976) - A pest species that damages crops and other plants. In India, Burma and Sri Lanka Dorylus orientalis is a plant-pest, sometimes a serious one, of several plants including vegetables, tubers, bulbs, shrubs, trees and also including cash crops such as sugarcane, coconut palm, citrus and groundnut. Tubers of potatoes and bulbs are eaten through hollow, while in other cases the roots and root-collars, especially of seedlings in gardens and nurseries, are eaten. The damage is entirely underground and is done by the workers. Workers are not entirely herbivorous; they also eat insects and earthworms. Other publications have variously stated (see the publication for a summary of previously published records) this species is insectivorous, carnivorous (insects and earthworms), and omnivorous (including plants). They have also been observed preying on Pheidole indica.

Dorylus orientalis is a subterranean ant. Their nests are not known but they have been found in tunnels and galleries in the soil. Males swarm at night and are attracted to light. Females are unknown. In north India swarms occur at the end of the cold weather, in late February (Lefroy 1909).

Wilson (1964) - Green (1903) claimed that orientalis is vegetarian, attacking potatoes, dahlias, and roots of the common sunflower. But Mukerji (1933) doubted this. He studied the worker mouthparts in detail and found them " better adapted for feeding on animal food than on plants. The mandible is similar to that of the species Dorylus labiatus, which is carnivorous in habit. The sharp-pointed bristles, spines, and setae on the 1st and 2nd maxillae, can well pierce the skin of the victim, and draw out the nutritive fluid from the body of their prey, which they suck by their mobile tongue." This conclusion is supported by certain limited observations. "Examples of this species were found within the college compound at Ballygunge, Calcutta, engaged in feeding on a dead earthworm, underneath an earthern seedling pot. On digging the adjoining turf a large number of these blind ants were found below the surface of the earth, a few being busy in attacking a live grub of a beetle. Evidently they reached the bottom of the seedling pot by tunneling through the ground to hunt the prey, as none of them were seen on the surface of the lawn. These specimens were then collected in live condition, and were kept in an earthern vessel half filled with moist earth. They settled down there, and made nests in the earth. I fed them with small live earthworms, but they did not partake of any vegetable food which was given to them."

Nomenclature

 * . Dorylus orientalis Westwood, 1835: 72 (m.) INDIA.
 * Type-material: holotype male.
 * Type-locality: India: (“India Orientali”, no further data).
 * [Note: type-locality Bengal, after Shuckard, 1840c: 321, Westwood, 1842: 80.]
 * Type-depository: OXUM.
 * Forel, 1901a: 463 (w.).
 * Combination in D. (Alaopone): Emery, 1895j: 731.
 * Status as species: Shuckard, 1840c: 320; Westwood, 1842: 80; Smith, F. 1859b: 3; Mayr, 1863: 408; Roger, 1863b: 41; Smith, F. 1871a: 335; Emery, 1889b: 487; Dalla Torre, 1893: 12; Emery, 1895j: 731; Emery, 1895k: 453; Forel, 1901a: 463; Emery, 1901c: 196 (in key); Emery, 1901f: 113; Rothney, 1903: 96; Bingham, 1903: 4; Forel, 1906b: 90; Forel, 1907e: 17; Emery, 1910b: 15; Forel, 1913k: 20; Wheeler, W.M. 1913e: 233; Santschi, 1924c: 97; Karavaiev, 1926d: 422; Wheeler, W.M. 1927b: 42; Wheeler, W.M. 1928c: 3; Donisthorpe, 1929a: 445; Mukerjee, 1930: 149; Wheeler, W.M. 1930h: 57; Menozzi, 1939a: 326; Santschi, 1939a: 147; Chapman & Capco, 1951: 9; Collingwood, 1962: 224; Wilson, 1964a: 442; Collingwood, 1970: 372; Wang, M. 1992: 678; Wu, J. & Wang, 1992: 1303; Xu, 1994a: 118; Bolton, 1995b: 180; Wu, J. & Wang, 1995: 52; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16; Mathew & Tiwari, 2000: 268; Zhou, 2001b: 56; Jaitrong & Nabhitabhata, 2005: 21; Pfeiffer, et al. 2011: 35; Guénard & Dunn, 2012: 26; Eguchi, Bui, et al. 2014: 33 (redescription); Bharti, Guénard, et al. 2016: 22; Jaitrong, Guénard, et al. 2016: 26.
 * Senior synonym of curtisii: Forel, 1901a: 463; Bingham, 1903: 4; Emery, 1910b: 15; Wilson, 1964a: 442; Bolton, 1995b: 180; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16; Zhou, 2001b: 56.
 * Senior synonym of fuscus: Wilson, 1964a: 442; Bolton, 1995b: 180; Tang, J., Li, et al. 1995: 44; Zhou, 2001b: 57.
 * Senior synonym of longicornis: Bingham, 1903: 4; Wilson, 1964a: 442; Bolton, 1995b: 180; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16; Zhou, 2001b: 57.
 * Senior synonym of oberthueri: Forel, 1901a: 463; Bingham, 1903: 4; Emery, 1910b: 15; Wheeler, W.M. 1911f: 158; Wilson, 1964a: 442; Bolton, 1995b: 180; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16.
 * Distribution: China, India, Laos, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam.
 * Current subspecies: nominal plus obscuriceps.
 * curtisii. Labidus (Typhlopone) curtisii Shuckard, 1840b: 265 (w.) (no state data).
 * Type-material: holotype(?) worker.
 * [Note: no indication of number of specimens is given.]
 * Type-locality: none given.
 * [Note: Smith, F. 1858b: 112, gives the locality as Ceylon (= Sri Lanka).]
 * Type-depository: BMNH.
 * Combination in Typhlopone: Smith, F. 1858b: 112;
 * combination in Dorylus: Dalla Torre, 1893: 10;
 * combination in D. (Alaopone): Emery, 1895j: 730.
 * Status as species: Smith, F. 1858b: 112; Roger, 1861a: 45; Roger, 1863b: 20; Mayr, 1863: 457; Motschoulsky, 1863: 15; Dalla Torre, 1893: 10; Emery, 1895j: 730; Mayr, 1897: 420.
 * Junior synonym of orientalis: Forel, 1901a: 463; Bingham, 1903: 4; Emery, 1910b: 15; Wilson, 1964a: 442; Bolton, 1995b: 178; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16; Zhou, 2001b: 55.
 * fuscus. Dorylus fuscus Emery, 1889b: 487 (m.) MYANMAR.
 * Type-material: 2 syntype males.
 * Type-locality: Myanmar (“Birmania”): Rangoon (L. Fea).
 * Type-depository: MSNG.
 * Combination in D. (Alaopone): Emery, 1895j: 731.
 * Subspecies of orientalis: Dalla Torre, 1893: 13; Emery, 1895j: 731; Emery, 1895k: 453; Forel, 1901a: 463; Bingham, 1903: 5; Emery, 1910b: 15; Santschi, 1939a: 147; Chapman & Capco, 1951: 9.
 * Junior synonym of orientalis: Wilson, 1964a: 442; Bolton, 1995b: 179; Tang, J., Li, et al. 1995: 44; Zhou, 2001b: 57.
 * longicornis. Dorylus longicornis Shuckard, 1840c: 321 (m.) INDIA (West Bengal).
 * Type-material: holotype male.
 * Type-locality: India: Bengal.
 * Type-depository: OXUM.
 * Combination in D. (Alaopone): Emery, 1895j: 731.
 * Status as species: Westwood, 1842: 80; Smith, F. 1859b: 3; Mayr, 1863: 408; Roger, 1863b: 41; Smith, F. 1871a: 335; Rothney, 1889: 371.
 * Subspecies of orientalis: Emery, in Dalla Torre, 1893: 13; Emery, 1895j: 731; Emery, 1895k: 453; Forel, 1901a: 463; Forel, 1907e: 17; Emery, 1910b: 15; Santschi, 1920h: 160 (footnote); Santschi, 1924c: 97; Santschi, 1939a: 147; Chapman & Capco, 1951: 9.
 * Junior synonym of orientalis: Bingham, 1903: 4; Wilson, 1964a: 442; Bolton, 1995b: 179; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16; Zhou, 2001b: 57.
 * oberthueri. Alaopone oberthuri Emery, 1881a: 274 (footnote), figs. (w.) INDIA (West Bengal).
 * [Note: name is spelled oberthuri in species heading, but oberthüri in figure captions; hence correct spelling taken as oberthueri.]
 * Type-material: syntype workers (number not stated).
 * Type-locality: India: Calcutta (R. Oberthür).
 * Type-depository: MSNG.
 * Combination in Dorylus: Dalla Torre, 1893: 12.
 * Status as species: Emery, 1889b: 488; Dalla Torre, 1893: 12; Emery, 1895k: 453.
 * Junior synonym of curtisii: Emery, 1895j: 730.
 * Junior synonym of orientalis: Forel, 1901a: 463; Bingham, 1903: 4; Emery, 1910b: 15; Wheeler, W.M. 1911f: 158; Wilson, 1964a: 442; Bolton, 1995b: 178; Tang, J., Li, et al. 1995: 44; Tiwari, 1999: 16.

Description
Worker

Eguchi et al. (2014): (1) workers are clearly subdivided into two series by a set of qualitative characters (Table 1); (2) the “typical series” is numerically much more dominant than the “atypical series” in foraging columns (the latter occupied less than 1% of the whole of the workers collected); (3) a wider size variation was observed in the former (HW, 0.48–1.41 mm; ML, 0.42–1.12 mm) than in the latter (HW, 0.44–1.13 mm; ML, 0.35–0.79 mm); and (4) within the atypical series, smaller workers are numerically much dominant.

Typical series workers: Head in full-face view distinctly longer than broad, with sides parallel or slightly convergent posteriad from the level of mandibular insertions, with posterior margin weakly concave (Figs. 1, 5, 7); a short median longitudinal furrow present running anteriad from the posterior concavity; anterior clypeal margin weakly produced and distinctly truncated in the middle, with a very long and thick median clypeal seta and a pair of thick paracarinal setae; frontal lobes closely approximated (often confluent posteriorly in smaller workers), extending anteriad close to anterior clypeal margin (the lobes reaching anterior margin in largest workers); mandibular blade falcate; apical tooth very large; preapical tooth distinct but much smaller than apical tooth, followed by long edentate or indistinctly serrate margin and small third (basal) tooth; basal margin edentate; antenna 9-segmented; scape not reaching midlength of head when it is laid backward; relative lengths of antennal segments II–IX as in Figs. 5 and 7; IX much longer than broad, weakly flattened apically. Mesosoma in lateral view relatively long and low, with a straight dorsal outline (Figs. 2, 9), in dorsal view slightly narrowed between promesonotum and propodeum. Petiolar node in dorsal view trapezoidal, broader than long, with weakly concave anterior margin, almost straight lateral margin, and almost straight or slightly convex posterior margin; petiolar node in lateral view roundly convex apically and weakly leaned posteriad; subpetiolar process rectangular, leaned anteriad. Abdominal posttergites IV, V and VI each almost as long as high.

Body covered by short decumbent to appressed hairs as shown in Figs. 1–3, 5, 7, 9 and 12, with a few long hairs on the dorsum of petiole, and gaster. Body largely smooth except for hair-pits, but anterior and posterolateral faces of promesonotum, part of mesopleuron and metapleuron, and posterolateral face of propodeum shagreened. Body reddish-brown to yellowish-brown; smaller worker lighter and rather yellowish in color

Measurements and indices (30 workers): HL, 0.57–1.69 mm; HW, 0.48–1.41 mm; ClyL, 0.03–0.07 mm; SL, 0.26–0.58 mm; ML, 0.42–1.12 mm; PW, 0.29–0.87 mm; HFL, 0.36–0.99 mm; PtNL, 0.19–0.54 mm; PtW, 0.21–0.60 mm; PtNH, 0.20–0.63 mm; CI, 80–97; ClyI, 3–7; SI, 40–55; MI, 128–145; PtI1, 80–97; PtI2, 81–99; HFI 70–78.

Atypical series workers: Atypical series consists of small to medium-sized workers only (HW ranges from 0.44 to1.13 mm in the atypical series, but from 0.48 to 1.41 mm in the typical series). Head in full-face view as long as or longer than broad, with sides more or less divergent posteriad from the level of mandibular insertions to midlength of head or further (Figs. 6, 8), with posterior margin distinctly concave; head in lateral view slightly more thickened dorsoventrally in medium-sized workers than in smaller workers (Figs. 10, 11); a short median longitudinal furrow present running anteriad from posterior concavity; anterior clypeal margin distinctly convex medially, with rounded apex in medium-sized workers (Fig. 6) or with subtriangular apex in smaller workers (Fig. 8); median clypeal seta short and thin in medium-sized workers (Fig. 6), and vestigial or completely absent in smaller workers (Fig. 8); paracarinal setae almost completely absent; frontal lobes vertical, approximated anteriorly, and often confluent posteriorly, not reaching anterior clypeal margin; mandibular blade elongate-triangular in medium-sized workers (Fig. 6), or somewhat linear in smaller workers (Fig. 8) but not falcate as seen in the typical workers (Figs. 5, 7); apical tooth very large; preapical tooth also developed well but smaller than apical tooth, followed by several denticles and distinct third (basal) tooth; basal margin edentate or with several denticles; antenna 8-segmented; scape not reaching midlength of head when it is laid backward; relative lengths of antennal segments II–VIII as in Figs. 6 and 8; VIII much longer than broad, weakly flattened apically. Mesosoma in lateral view relatively short and high, with a straight dorsal outline (Figs. 10, 13), in dorsal view narrowed behind promesonotum; humeral angle of promesonotum expanded laterad in medium-sized workers more than in smaller workers (Figs. 13, 14). Petiolar node in dorsal view trapezoidal, broader than long, with weakly concave anterior margin, almost straight lateral margin, and almost straight or slightly convex posterior margin; petiolar node in lateral view roundly convex and weakly leaned posteriad; subpetiolar process rectangular or round, leaned anteriad. Abdominal posttergite IV in lateral view almost as long as or a little shorter than high; posttergites V and VI a little shorter than high.

Body covered by short decumbent to appressed hairs as shown in Figs. 6, 8, 10, 11 and 13, with a few long hairs on the dorsum of petiole, and gaster. Body largely smooth with hair-pits, but anterior and posterolateral faces of promesonotum, part of mesopleuron and metapleuron, and posterolateral face of propodeum shagreened. Body reddish-brown to yellowish-brown; smaller worker lighter and rather yellowish in color.

Measurements and indices (39 workers). HL, 0.52–1.13 mm; HW, 0.44–1.13 mm; ClyL, 0.07–0.11 mm; SL, 0.20–0.45 mm; ML, 0.35–0.79 mm; PW, 0.28–0.72 mm; HFL, 0.31–0.79 mm; PtNL, 0.18–0.38 mm; PtW, 0.23–0.46 mm; PtNH, 0.21–0.46 mm; CI, 85–102; ClyI, 9–15; SI, 39–46; MI, 109–131; PtI1, 74–87; PtI2, 77–93; HFI 67–73.

Major

Bingham (1903): Castaneous brown, with the abdomen generally lighter in colour than the head and thorax; head and thorax densely, abdomen more lightly punctured; pubescence almost entirely wanting, a few yellow erect hairs on the front of the head, at the apex of and beneath the abdomen. Head rectangular, occiput deeply emarginate; the head slightly wider anteriorly than posteriorly, a deeply impressed medial line or furrow down the front. Thorax elongate, somewhat rectangular, but rounded anteriorly, depressed and flat above, constricted at the pro-mesonotal suture; legs short and robust. Node of pedicel broader than long, transverse anteriorly and posteriorly; abdomen depressed above, about as long as the thorax and node of pedicel united.

Length: 5 - 6 mm

Minor

Bingham (1903): Similar, very much smaller and very much lighter in colour, being of a pale honey-yellow; occiput not emarginate, front of the head not furrowed; node of the pedicel rounded above.

Length: 2.5 - 3 mm

Male

Bingham (1903): Brownish yellow, the mandibles and legs castaneous brown, pubescent and very hairy, the hairs erect, yellow, and most dense under the head, on the sides and beneath the thorax, and at the apex of the abdomen; pubescence pale, dense, line and silky; wings flavo-hyaline. Head narrow, transverse, eyes and ocelli large; mandibles short, broad, porrect, blunt at apex and with a blunt tooth on the inner margin at base. Thorax massive, gibbous in front; scutellum compressed; postscutellum narrow, sunk between the scutellum and metanotum, basal portion of the latter depressed, flat, apex truncate; legs short and robust. Node of pedicel somewhat square, convex above, slightly contracted anteriorly and posteriorly; abdomen long, cylindrical, massive, clavate towards the apex, and curved downwards.

Length: 17 - 23 mm

References based on Global Ant Biodiversity Informatics

 * Bharti H., Y. P. Sharma, M. Bharti, and M. Pfeiffer. 2013. Ant species richness, endemicity and functional groups, along an elevational gradient in the Himalayas. Asian Myrmecology 5: 79-101.
 * Bharti H., Y. P. Sharma, and A. Kaur. 2009. Seasonal patterns of ants (Hymenoptera: Formicidae) in Punjab Shivalik. Halteres 1(1): 36-47.
 * Bhoje P. M., K. Shilpa, and T. V. Sathe. 2014. Diversity of ants (Hymenoptera: Formicidae) from Kolhapur district of Maharashtra, India. Uttar Pradesh J. Zool. 34(1): 23-25.
 * Chapman, J. W., and Capco, S. R. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia. Monogr. Inst. Sci. Technol. Manila 1: 1-327
 * Chapman, J.W. and S.R. Capco. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia. Monographs of the Institute of Science and Technology (Manila) 1: 1- 327
 * Cheng D., Z. Chen, and S. Zhou. 2015. An analysis on the ant fauna of Jinzhongshan Nature Reserve in Gunagxi, China. Journal of Guangxi Normal University: Natural Science Edition 33(3): 129.137.
 * Chhotani O. B., and K. K. Ray. 1976. Fauna of Rajasthan, India, Hymenoptera. Records of the Zoological Survey of India 71: 13-49.
 * Collingwood C. A. 1962. Some ants (Hym. Formicidae) from north-east Asia. Entomologisk Tidskrift 83: 215-230.
 * Collingwood C.A. 1970. Formicidae (Hymenopter: Aculeata) of Nepal. Himalaya Khumbu Himal, 3: 371-388.
 * Dad J. M., S. A. Akbar, H. Bharti, and A. A. Wachkoo. 2019. Community structure and ant species diversity across select sites ofWestern Ghats, India. Acta Ecologica Sinica 39: 219–228.
 * Dias R. K. S. 2002. Current knowledge on ants of Sri Lanka. ANeT Newsletter 4: 17- 21.
 * Dias R. K. S. 2006. Current taxonomic status of ants (Hymenoptera: Formicidae) in Sri Lanka. The Fauna of Sri Lanka: 43-52. Bambaradeniya, C.N.B. (Editor), 2006. Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. The World Conservation Union, Colombo, Sri Lanka & Government of Sri Lanka. viii + 308pp.
 * Dias R. K. S. 2013. Diversity and importance of soil-dweeling ants. Proceedings of the National Symposium on Soil Biodiversity, chapt 4, pp 19-22.
 * Dias R. K. S., K. R. K. A. Kosgamage, and H. A. W. S. Peiris. 2012. The Taxonomy and Conservation Status of Ants (Order: Hymenoptera, Family: Formicidae) in Sri Lanka. In: The National Red List 2012 of Sri Lanka; Conservation Status of the Fauna and Flora. Weerakoon, D.K. & S. Wijesundara Eds., Ministry of Environment, Colombo, Sri Lanka. p11-19.
 * Dias R. K. S., and K. R. K. Anuradha Kosgamage. 2012. Occurrence and species diversity of ground-dwelling worker ants (Family: Formicidae) in selected lands in the dry zone of Sri Lanka. J. Sci. Univ. Kelaniya 7: 55-72.
 * Dias, R.K.S. 2006. Overview of ant research in Sri Lanka: 2000-2004. ANeT Newsletter 8:7-10
 * Donisthorpe H. 1929. The Formicidae (Hymenoptera) taken by Major R. W. G. Hingston, M.C., I.M.S. (ret.), on the Mount Everest Expedition, 1924. Annals and Magazine of Natural History (10)4: 444-449.
 * Eguchi K., B. T. Viet, and S. Yamane. 2014. Generic Synopsis of the Formicidae of Vietnam (Insecta: Hymenoptera), Part IICerapachyinae, Aenictinae, Dorylinae, Leptanillinae, Amblyoponinae, Ponerinae, Ectatomminae and Proceratiinae. Zootaxa 3860: 001-046.
 * Eguchi K., T. V. Bui, S. Yamane, H. Okido, and K. Ogata. 2004. Ant faunas of Ba Vi and Tam Dao, North Vietnam (Insecta: Hymenoptera: Formicidae). Bull. Inst. Trop. Agr. Kyushu Univ. 27: 77-98.
 * Eguchi K., V. T. Bui, E. Oguri, M. Maruyama, and S. Yamane. 2014. A new data of worker polymorphism in the ant genus Dorylus (Hymenoptera: Formicidae: Dorylinae). Journal of Asia-Pacific Entomology 17: 31-36.
 * Emery C. 1895. Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 8: 685-778.
 * Emery C. 1901. Ameisen gesammelt in Ceylon von Dr. W. Horn 1899. Deutsche Entomologische Zeitschrift 1901: 113-122.
 * Emery C. 1910. Hymenoptera. Fam. Formicidae. Subfam. Dorylinae. Genera Insectorum 102: 1-34.
 * Field Museum Collection, Chicago, Illinois (C. Moreau)
 * Forel A. 1901. Les Formicides de l'Empire des Indes et de Ceylan. Part VIII. J. Bombay Nat. Hist. Soc. 13: 462-477
 * Forel A. 1906. Les fourmis de l'Himalaya. Bulletin de la Société Vaudoise des Sciences Naturelles 42: 79-94.
 * Forel A. 1907. Formiciden aus dem Naturhistorischen Museum in Hamburg. II. Teil. Neueingänge seit 1900. Mitt. Naturhist. Mus. Hambg. 24: 1-20.
 * Forel A. 1909. Études myrmécologiques en 1909. Fourmis de Barbarie et de Ceylan. Nidification des Polyrhachis. Bull. Soc. Vaudoise Sci. Nat. 45: 369-407.
 * [[Media:Forel 1913l.pdf|Forel A. 1913k. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien ausgeführt im Auftrage der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen. II. Ameisen aus Sumatra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren 1911-1912. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 36:1-148.]]
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Huong N. T. T., P. V. Sang, and B. T. Viet. 2015. A preliminary study on diversity of ants (Hymenoptera: Formicidae) at Hon Ba Nature Reserve. Environmental Scientific Conference 7: 614-620.
 * Jaitrong W., B. Guenard, E. P. Economo, N. Buddhakala, and S. Yamane. 2016. A checklist of known ant species of Laos (Hymenoptera: Formicidae). Asian Myrmecology 8: 1-32. DOI: 10.20362/am.008019
 * Karavaiev V. 1926. Ameisen aus dem Indo-Australischen Gebiet. Treubia 8: 413-445.
 * Karmaly K. A.; S. Sumesh, T. P. Rabeesh, and L. Kishore. 2010. A checklist of ants of Thirunelli in Wayanad, Kerala. J. of the Bombay Natural History Society 107(1): 64-67.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * Liu X. 2012. Taxonomy, diversity and spatial distribution characters of the ant family Formicidae (Insecta: Hymenoptera) in southeastern Tibet. PhD Thesis 139 pages
 * Liu X., Z. Xu, N. Yu, and C. Zhang. 2016. Distribution patterns of ant species ( Hymenoptera: Formicidae) in Galongla Mountains and Medog Valley of Southeastern Tibet. Scientia Silvae Sinicae 52(11): 88-95.
 * Mathew R., and R. N. Tiwari. 2000. Insecta: Hymenoptera: Formicidae. Pp. 251-409 in: Director; Zoological Survey of India (ed.) 2000. Fauna of of Meghalaya. Part 7. [State Fauna Series 4.] Insecta 2000. Calcutta: Zoological Survey of India, 621 pp.
 * Mukerjee D. 1930. Report on a collection of ants in the Indian Museum, Calcutta. J. Bombay Nat. Hist. Soc. 34: 149-163.
 * Musthak Ali T. M. 1982. Ant fauna (Hymenoptera: Formicidae) of Bangalore with observations on their nesting and foraging habits. Thesis Abstracts. Haryana Agricultural University 8: 370-371.
 * Musthak Ali T. M. 1991. Ant Fauna of Karnataka-1. Newsletter of IUSSI Indian Chapter 5(1-2): 1-8.
 * Ogata K. 2005. Asian ant inventory and international networks. Report on Insect inventory Project in Tropic Asia TAIIV: 145-170.
 * Pajni H. R., and R. K. Suri. 1978. First report on the Formicid fauna (Hymenoptera) of Chandigarh. Res. Bull. (Science) Punjab University 29: 5-12.
 * Pfeiffer M.; Mezger, D.; Hosoishi, S.; Bakhtiar, E. Y.; Kohout, R. J. 2011. The Formicidae of Borneo (Insecta: Hymenoptera): a preliminary species list. Asian Myrmecology 4:9-58
 * Rajan P. D., M. Zacharias, and T. M. Mustak Ali. 2006. Insecta: Hymenoptera: Formicidae. Fauna of Biligiri Rangaswamy Temple Wildlife Sanctuary (Karnataka). Conservation Area Series, Zool. Surv. India.i-iv,27: 153-188.
 * Roonwal M. L. 1976. Plant-pest status of root-eating ant, Dorylus orientalis, with notes on taxonomy, distribution and habits (Insecta: Hymenoptera). Journal of the Bombay Natural History Society 72: 305-313.
 * Santschi F. 1924. Fourmis d'Indochine. Opuscules de l'Institut Scientifique de l'Indochine 3: 95-117
 * Sheela S. 2008. Handbook of Hymenoptera, Formicidae. Zoological Survey of India, 56 pages
 * Sheikh A. H., M. Manzoor, Y. A. Rather, and T. Jobiraj. 2019. Taxonomic study of ant (Formicidae : Hymenoptera) fauna of Dumna Nature Park, Jabalpur, Madhya Pradesh, India. Journal of Entomological Research 43(2): 203-212.
 * Song Y., Z. Xu, C. Li, N. Zhang, L. Zhang, H. Jiang, and F. Mo. 2013. An Analysis on the Ant Fauna of the Nangun river Nature Reserve in Yunnan, China. Forest Research 26(6): 773-780.
 * Tak N. 2009. Ants Formicidae of Rajasthan. Records of the Zoological Survey of India, Occasional Paper No. 288, iv, 46 p
 * Tak N., N. S. Rathore, and S. Kumar. 2007. Insecta: Hymenoptera. Fauna of Pichhola lake (Rajasthan). Wetland Ecosystem series. Zool. Surv. India. 8 : 127-130.
 * Tak N., and N. S. Rathore. 1996. Ant (Formicidae) fauna of the Thar Desert. Pp. 271-276 in: Ghosh, A. K.; Baqri, Q. H.; Prakash, I. (eds.) 1996. Faunal diversity in the Thar Desert: gaps in research. Jodhpur: Scientific Publishers, xi + 410 pp.
 * Tak N., and N. S. Rathore. 2004. Insecta: Hymenoptera. Rathore, N.S. Fauna of Desert National Park Rajasthan (proposed biosphere reserve). Conservation Area Series 19,Zool. Surv. India. 1-135. Chapter pagination: 81-84.
 * Tak N., and N. S. Rathore. 2004. Insecta: Hymenoptera: Formicidae. State Fauna Series 8: Fauna of Gujarat. Zool. Surv. India. Pp. 161-183.
 * Tak N., and S. L. Kazmi. 2011. On a collection of Insecta: Hymenoptera: Formicidae from Uttarakhand. Rec. zool. Surv. India : 111(2) : 39-49.
 * Tak, N. 2009. Ants (Hymenoptera: Formicidae) of the Thar Desert of Rajasthan and Gujarat. in C. Sivaperuman et al. (eds.), Faunal Ecology and Conservation of the Great Indian Desert
 * Tang J., Li S., Huang E., Zhang B. and Chen Y.. 1995. Hymenoptera: Formicidae (1). Economic Insect Fauna of China 47: 1-133.
 * Thapa V. K. 2000. An Inventory of Nepal's Insects, Vol. III. IUCN Nepal, Kathmandu, xi + 475 pp.
 * Tiwari R. N. 1997. Hymenoptera: Formicidae. Pp. 441-451 in: Zoological Survey of India; Director (ed.) 1997. Fauna of Delhi. Calcutta: Zoological Survey of India, vi + 903 pp.
 * Tiwari R. N. 1999. Taxonomic studies on ants of southern India (Insecta: Hymenoptera: Formicidae). Memoirs of the Zoological Survey of India 18(4): 1-96.
 * Tiwari R. N., B. G. Kundu, S. Roy Chowdhury, and S. N. Ghosh. 2003. Insecta: Hymenoptera: Formicidae. Fauna of Sikkim. Part 4. State Fauna Series. 9.Zool.Surv.India. i-iii, 1-512. Chapter pagination: 467-506.
 * Tiwari R.N., B.G. Kundu, S. Roychowdhury, S.N. Ghosh. 1999. Insecta: Hymenoptera: Formicidae. Pp. 211-294 in: Director; Zoological Survey of India (ed.) 1999. Fauna of West Bengal. Part 8. Insecta (Trichoptera, Thysanoptera, Neuroptera, Hymenoptera and Anoplura). Calcutta: Zoological Survey of India, iv + 442 pp.
 * Tiwari, R.N. 1999. Taxonomic studies on ants of southern India (Insecta: Hymenoptera: Formicidae). Memoirs of the Zoological Survey of India 18(4):1-96
 * Viehmeyer H. 1912. Ameisen aus Deutsch Neuguinea gesammelt von Dr. O. Schlaginhaufen. Nebst einem Verzeichnisse der papuanischen Arten. Abhandlungen und Berichte des Königlichen Zoologischen und Anthropologische-Ethnographischen Museums zu Dresden 14: 1-26.
 * Wang C. and Wu J.. 1992. Ants of the Jianfengling forest region in Hainan Province (Hymenoptera: Formicidae). Scientia Silvae Sinicae 28: 561-564.
 * Wheeler W. M. 1928. Ants collected by Professor F. Silvestri in China. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 22: 3-38.
 * Wheeler W. M. 1930. A list of the known Chinese ants. Peking Natural History Bulletin 5: 53-81.
 * Wilson E. O. 1964. The true army ants of the Indo-Australian area (Hymenoptera: Formicidae: Dorylinae). Pacific Insects 6: 427-483.
 * Wu B., Y. Lu, L. Zeng, and G. Liang. 2008. Influences of Solenopsis invicta Buren invasion on the native ant communities in different habitats in Guangdong. Chin. J. Appl. Ecol. 19(1): 151-156.
 * Xu Z. 1994. A taxonomic study of the ant subfamily Dorylinae China (Hymenoptera Formicidae Ponerinae). Journal of Southwest Forestry College 14(2): 115-122
 * Yamane S.; Bui T. V.; Ogata K.; Okido H.; Eguchi K. 2002. Ant fauna of Cuc Phuong National Park, North Vietnam (Hymenoptera: Formicidae). Bulletin of the Institute of Tropical Agriculture Kyushu University 25: 51-62.
 * Zryanin V. A. 2011. An eco-faunistic review of ants (Hymenoptera: Formicidae). In: Structure and functions of soil communities of a monsoon tropical forest (Cat Tien National Park, southern Vietnam) / A.V. Tiunov (Editor). – M.: KMK Scientific Press. 2011. 277 р.101-124.