Dorylus orientalis

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Dorylus orientalis
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Dorylinae
Genus: Dorylus
Species: D. orientalis
Binomial name
Dorylus orientalis
Westwood, 1835

Dorylus orientalis casent0217468 p 1 high.jpg

Dorylus orientalis casent0217468 d 1 high.jpg

Specimen Labels

Subspecies
Synonyms

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.


Photo Gallery

  • Workers from Kerala, India. Photo by Kalesh Sadasivan.
  • Dorylus orientalis workers hunting termites, India. Photo by Manoj Vembayam.
  • Dorylus orientalis major worker. Dehradun, Uttarakhand, India. Photo by Yathumon M A.
  • 'Dorylus orientalis major worker. Dehradun, Uttarakhand, India. Photo by Yathumon M A.
  • 'Dorylus orientalis major and minor (background) workers. Dehradun, Uttarakhand, India. Photo by Yathumon M A.

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

Latitudinal Distribution Pattern

Latitudinal Range: 32.812778° to 8.033333333°.

   
North
Temperate
North
Subtropical
Tropical South
Subtropical
South
Temperate

Distribution based on Regional Taxon Lists

Indo-Australian Region: Borneo, Brunei Darussalam, Indonesia, Malaysia.
Oriental Region: Bangladesh, India (type locality), Laos, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand, Vietnam.
Palaearctic Region: China.

Distribution based on AntMaps

AntMapLegend.png

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
pChart

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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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."

Castes

  • Liu, C. et al. 2020. Ants of the Hengduan Mountains, Figure 9, Dorylus orientalis.
  • Liu, C. et al. 2020. Ants of the Hengduan Mountains, Figure 10, Dorylus orientalis, solder.

Worker

Images from AntWeb

Dorylus orientalis casent0249443 h 1 high.jpgDorylus orientalis casent0249443 p 1 high.jpgDorylus orientalis casent0249443 d 1 high.jpgDorylus orientalis casent0249443 l 1 high.jpg
Worker. Specimen code casent0249443. Photographer Will Ericson, uploaded by California Academy of Sciences. Owned by PSWC, Philip S. Ward Collection.

Male

Images from AntWeb

Dorylus orientalis casent0901942 h 1 high.jpgDorylus orientalis casent0901942 p 1 high.jpgDorylus orientalis casent0901942 p 2 high.jpgDorylus orientalis casent0901942 p 3 high.jpgDorylus orientalis casent0901942 d 1 high.jpgDorylus orientalis casent0901942 l 1 high.jpg
Type of Dorylus orientalisMale (alate). Specimen code casent0901942. Photographer Ryan Perry, uploaded by California Academy of Sciences. Owned by OUM, Oxford, UK.
Dorylus orientalis casent0901951 h 1 high.jpgDorylus orientalis casent0901951 p 1 high.jpgDorylus orientalis casent0901951 p 2 high.jpgDorylus orientalis casent0901951 p 3 high.jpgDorylus orientalis casent0901951 d 1 high.jpgDorylus orientalis casent0901951 l 1 high.jpg
Type of Dorylus longicornisMale (alate). Specimen code casent0901951. Photographer Ryan Perry, uploaded by California Academy of Sciences. Owned by OUM, Oxford, UK.
Dorylus orientalis casent0903710 h 1 high.jpgDorylus orientalis casent0903710 p 1 high.jpgDorylus orientalis casent0903710 p 2 high.jpgDorylus orientalis casent0903710 p 3 high.jpgDorylus orientalis casent0903710 d 1 high.jpgDorylus orientalis casent0903710 l 1 high.jpg
Syntype of Dorylus fuscusMale (alate). Specimen code casent0903710. Photographer Will Ericson, uploaded by California Academy of Sciences. Owned by MSNG, Genoa, Italy.
Wilson 1964 Army Ant fig 1-14

Phylogeny

Relationships among selected Dorylus species based on Kronauer et al. (2007).

Dorylus

Dorylus laevigatus

Dorylus conradti

Dorylus orientalis

Dorylus fimbriatus laevipodex

Dorylus fulvus

Dorylus spininodis

Dorylus mayri

Dorylus nigricans rubellus

Dorylus nigricans molestus

Dorylus nigricans terrificus

Dorylus wilverthi

Dorylus nigricans burmeisteri

Dorylus nigricans sjostedti

Dorylus nigricans arcens

Dorylus nigricans

Dorylus emeryi

Dorylus gerstaeckeri

Dorylus gribodoi

Dorylus kohli

Dorylus emeryi opacus

Dorylus braunsi

Dorylus affinis

Dorylus helvolus

Nomenclature

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

  • orientalis. 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

References based on Global Ant Biodiversity Informatics

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  • 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.
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  • 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.
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