Pheidole indica

Fischer and Fisher (2013) - Pheidole teneriffana (=P. indica) is an invasive species with collection records scattered over several continents and islands across the globe. Described from the Canary Islands and found widely distributed throughout the greater Mediterranean region, its native range and origin are unknown (Wilson 2003, Wetterer 2011). Probably introduced to the Malagasy region, it was described from Madagascar as P. voeltzkowii, only one year after the publication of the senior synonym. Morphologically, P. teneriffana can be grouped together with, and is possibly related to, Pheidole fervens, Pheidole oceanica and Pheidole sinaitica. In the New World, Pheidole teneriffana has been introduced to California (Martinez 1992, Snelling 1992), Cuba, Peru (Wilson 2003), and the West Indian islands (Wetterer 2011). It seems to be common in dry habitats (Wetterer 2011), especially along coasts and in urban areas (Collingwood et al. 1997, Gómez & Espalader 2006), and has been described as aggressive toward other ant species, locally abundant, and spreading in urban areas (Collingwood 1985, Gómez & Espalader 2006). In the Malagasy region P. teneriffana was collected on the Comoros, Mauritius, the Seychelles, and from coastal towns in Madagascar, usually from under stones, ground nests, or foraging on the ground or lower vegetation in urban or garden habitats at elevations between 2 and 296 m, on Mayotte in native littoral and secondary forest (7 m elevation). Recently in Saudi Arabia it was collected from soil, under stones, and foraging on the ground on a few farms at elevations between 570 and 1620 m.

Identification
Fischer and Fisher (2013) - Moderately small species (WL major 1.18–1.44 mm, WL minor 0.79–0.97 mm), minor workers with relatively long scapes and legs (SI minor 120–149, FI minor 134–170) and major workers with moderately long legs (FI major 73–89), both with a well-developed promesonotal process. Major with frontal carinae and antennal scrobe reaching posterior quarter of head, face with longitudinal rugae often very oblique posteriorly, curved toward posterolateral lobes, eyes moderately large (EI 13–16), submedian hypostomal teeth small, median process reduced and often very shallow, promesonotal process prominent and well developed, in profile metanotal groove shallow and dorsal propodeum usually level, postpetiole in dorsal view trapezoidal, often with rounded lateral corners, but sometimes with acute lateral dents, postpetiole between 1.9 and 2.5 times wider than petiole, postpetiolar ventral process reduced and very shallow. Minor head oval, posteriorly rounded, scapes and legs relatively long (SI 120–149, FI 134–170), with decumbent to subdecumbent pilosity, eyes relatively large (EI 25– 30), metanotal groove not or barely impressed in profile.

Distribution
An invasive species that has been reported occurring in the Canary Islands, the Mediterranean area, the Malagasy region, California, Cuba, Peru, and West Indian islands.

Distribution based on Regional Taxon Lists
Afrotropical Region: Comoros, Eritrea, Kenya, Saint Helena, Saudi Arabia, Socotra Archipelago, United Arab Emirates, Yemen. Malagasy Region: Madagascar, Mauritius, Mayotte, Seychelles. Nearctic Region: United States. Neotropical Region: Antigua and Barbuda, Barbados, Colombia, Cuba, Dominican Republic, Greater Antilles, Guadeloupe, Peru. Oriental Region: Bangladesh, India, Nepal, Pakistan. Palaearctic Region: Balearic Islands, Canary Islands, China, Egypt, Greece, Iberian Peninsula, Iran, Israel, Japan, Kuwait, Malta, Oman, Republic of Korea, Spain.

Biology
Wilson (2003) - Evidently a native of North Africa and possibly also the Canary Islands. It has turned up in disturbed habitats in Cuba, Peru, and California, but as of this writing is evidently still sporadic and local. In 1989 Martinez (1992) found a population occupying about two hectares of Admiral Kidd Park, evidently consisting of a single continuous, polydomous colony. The separate nests contained large numbers of workers and multiple inseminated queens, as many as 23 in one instance. Nest sites included lawns and open ground, where nest entrances were surmounted by mounds of excavated soil; crevices of sidewalks and curbs; and the bases of trees. New nest sites were occupied by budding from occupied sites. The workers were aggressive toward other ant species; they preyed on insects and harvested seeds. By 1998, according to Gulmahamad and Martinez (1999), the population was extinct. It had been weakened by attempts to exterminate it and changes in the nest habitat, and given the coup de grâce by encroaching Argentine ants (Linepithema humile).

Regional Notes
Heterick (2009) - WA Australia: Currently only confirmed for the Fremantle area, but ants of similar appearance have been seen in Claremont (a Perth suburb) and the wheatbelt town of York. Unlike the case with most nests of Pheidole megacephala, this species has diurnally active workers. Pheidole indica had not previously been reported from Australia.

Espadaler (2007) - Canary Islands: A nest with winged males was detected on cracks in the pavement. Ants rushed out after a small air blow with the aspirator.

Nomenclature

 *  indica. Pheidole indica Mayr, 1879: 679 (s.w.q.) INDIA. Forel, 1902c: 199 (m.); Imai, Baroni Urbani, et al. 1984: 6 (k.). Senior synonym of himalayana, rotschana, striativentris and material of the unavailable name formosae referred here: Eguchi, 2004: 198. See also: Bingham, 1903: 263; Menozzi, 1939a: 298; Ogata, 1982: 196.
 * striativentris. Pheidole striativentris Mayr, 1879: 678 (s.) INDIA. Forel, 1902c: 195 (w.q.). Junior synonym of indica: Eguchi, 2004: 199.
 * himalayana. Pheidole indica r. himalayana Forel, 1902c: 185 (s.), 199 (w.) INDIA. [Also described as new by Forel, 1902f: 546.] Raised to species: Bingham, 1903: 265. Subspecies of indica: Emery, 1921f: 91; Menozzi, 1939a: 298; Pisarski, 1967: 385. Junior synonym of indica: Eguchi, 2004: 198.
 * rotschana. Pheidole indica r. rotschana Forel, 1902c: 185 (s.), 199 (w.m.) INDIA. Imai, Baroni Urbani, et al. 1984: 6 (k.). [Also described as new by Forel, 1902f: 546.] Raised to species: Bingham, 1903: 264. Subspecies of indica: Forel, 1909e: 394; Forel, 1911i: 222. Junior synonym of indica: Eguchi, 2004: 199.
 * teneriffana. Pheidole teneriffana Forel, 1893d: 465 (s.w.) SPAIN (Canary Is). [Also described as new by Forel, 1894a: 160.] Santschi, 1908: 521 (q.). Gómez & Espadaler, 2006: 229 (m.). Status as species: Santschi, 1908: 521; Wheeler, W.M. 1922a: 821; Wheeler, W.M. 1927g: 105; Aguayo, 1932: 219; Finzi, 1936: 164; Baroni Urbani, 1968b: 438; Baroni Urbani, 1971c: 73; Barquin Diez, 1981: 114; Schembri & Collingwood, 1981: 428; Collingwood, 1985: 255; Agosti & Collingwood, 1987b: 271 (in key); Snelling, R.R. 1992b: 121; Mei, 1995: 762; Collingwood & Agosti, 1996: 324; Wilson, 2003: 640 (redescription); Collingwood, et al. 2004: 478; Gómez & Espadaler, 2006: 229; Heterick, 2009: 169; Fischer & Fisher, 2013: 346 (redescription). Senior synonym of taina: Wilson, 2003: 640. Senior synonym of voeltzkowii: Fischer & Fisher, 2013: 346. Junior synonym of indica: Sarnat, et al. 2015: 46.
 * taina. Pheidole teneriffana subsp. taina Aguayo, 1932: 219 (s.) CUBA. Junior synonym of teneriffana: Wilson, 2003: 640.
 * voeltzkowii. Pheidole voeltzkowii Forel, 1894e: 227 (s.w.m.) MADAGASCAR. Forel, 1897c: 207 (q.). Junior synonym of teneriffana: Fischer & Fisher, 2013: 340.

Type Material
Tenerife, Canary Islands. - as reported in Wilson (2003)

Description
Wilson (2003) - DIAGNOSIS Major: unique in the possession of a broad, convex metanotum and a four-lobed mesosomal profile in dorsal-oblique view (2 on pronotum, one each on mesonotum and metanotum); also, presence of a weak antennal scrobe; carinulae cover all the dorsal head surface except for the frontal triangle and midclypeus; carinulae originating laterad to antennal scrobes circle outward and downward again to travel to the eye and behind it; postpetiole elliptical from above.

Minor: occiput slightly narrowed, no nuchal collar.

MEASUREMENTS (mm) Major (Oriente, Cuba): HW 1.34, HL 1.34, SL 0.82, EL 0.20, PW 0.64. Minor (Oriente, Cuba): HW 0.62, HL 0.70, SL 0.74, EL 0.14, PW 0.34.

COLOR Major and minor: light yellowish brown, with head, mandibles, and gaster a slightly darker shade.

Fischer and Fisher (2013) - Major (n=14): HW 1.22–1.67 (1.47), HL 1.31–1.71 (1.51), SL 0.73– 0.84 (0.80), MDL 0.70–0.91 (0.82), EL 0.20–0.23 (0.22), WL 1.18–1.44 (1.28), PNH 0.43–0.66 (0.55), PNW 0.61–0.80 (0.70), MNH 0.76–0.99 (0.86), PDH 0.39–0.56 (0.45), PTL 0.36–0.45 (0.42), PPL 0.20–0.28 (0.25), PTH 0.24–0.30 (0.26), PPH 0.22–0.26 (0.24), PTW 0.16–0.22 (0.20), PPW 0.35–0.48 (0.42), PSL 0.17–0.24 (0.20), MFL 1.08–1.28 (1.18), MTL 0.66–0.95 (0.87), CI 93–101 (98), SI 49–62 (55), MDI 52–60 (56), EI 13–17 (15), FI 73–89 (80), PSLI 11–16 (14), LPpI 87–123 (104), DPpI 146–235 (172), PpWI 190–247 (213), PpLI 51–67 (59), PpHI 83–108 (90).

Head usually slightly longer than wide (CI 93–101), sides convex to strongly convex. Mandibles smooth and relatively long (MDI 52–60). Clypeus smooth, median carina inconspicuous or absent, usually two pairs of lateral carinae present. Frontal carinae well developed and reaching 3/4 of the way to posterior head margin, antennal scrobe conspicuous and weakly to superficially punctate. Frons longitudinally rugose, sides of head rugoreticulate, rugulae on posterolateral lobes variable, from straight longitudinal or longitudinal with scarce, short, reticulate rugulae in between, to obliquely curved toward posterolateral lobes, to transversal, interspaces smooth to superficially punctate; posterior head margin in dorsal view usually smooth to superficially sculptured. Scape moderately long (SI 49–62) with decumbent to subdecumbent pilosity and usually three erect longer hairs on outer edge. Eyes relatively large (EI 13–17). Submedian hypostomal teeth small to medium-sized, median process small to inconspicuous. Promesonotum in profile high-domed, convex, promesonotal process prominently produced, with deeply concave to angulate transverse groove, set almost at a right angle toward posterior declivity. Dorsum of promesonotum transversely rugulose, lateropronotum weakly rugulose to largely smooth, sides of mesonotum and propodeum densely punctate. Metanotal groove impressed to very shallow in profile, cross-ribs reduced to inconspicuous. Dorsum of propodeum weakly to superficially sculptured, in profile usually straight, not sloped toward spines, and about as long as posterior declivity. Propodeal spines acute, moderately short, slightly shorter than distance between their bases (PSI 11–16). Metatibia moderately long (FI 73–89), pilosity on inner edge decumbent, on outer edge subdecumbent. Postpetiole on average 1.7 times wider than long (DPpI 146–235) and 2.1 times wider than petiole (PpWI 190–247), sides in dorsal view roundly convex, or subangulate or with small denticle laterally, ventral process very small or inconspicuous. Dorsum of waist segments partly smooth, partly superficially punctate, remainder punctate. Gaster smooth to micropunctate. Standing hairs on mesosoma moderately scarce, yellowish, of short to moderate length, acute, with more abundant shorter decumbent pilosity. Color light to darker reddish brown, with dark brown to blackish gaster.

Minor (n=16): HW 0.50–0.65 (0.58), HL 0.60–0.74 (0.68), SL 0.64– 0.81 (0.74), MDL 0.35–0.46 (0.42), EL 0.14–0.17 (0.16), WL 0.79–0.97 (0.88), PNH 0.27–0.33 (0.30), PNW 0.31–0.42 (0.38), MNH 0.50–0.62 (0.58), PDH 0.25–0.31 (0.28), PTL 0.23–0.29 (0.26), PPL 0.15–0.18 (0.16), PTH 0.15–0.18 (0.16), PPH 0.13–0.16 (0.15), PTW 0.10–0.12 (0.11), PPW 0.19–0.26 (0.22), PSL 0.08–0.10 (0.09), MFL 0.71–0.95 (0.85), MTL 0.53–0.75 (0.66), CI 72–88 (85), SI 120–149 (129), MDI 66–83 (72), EI 25– 30 (28), FI 134–170 (147), PSLI 14–19 (16), LPpI 100–124 (112), DPpI 112–160 (133), PpWI 177–236 (199), PpLI 58–70 (63), PpHI 81–100 (94).

Head shape oval, distinctly longer than wide (CI 72–88), sides strongly convex, posterior head margin rounded and occipital carina conspicuous in full-face view. Mandibles moderately long (MDI 66–83), weakly longitudinally rugulose. Clypeus smooth, sometimes with short median and lateral carinae present. Face smooth, except for concentric rugulae around antennal insertion and usually two malar carinae reaching posterior eye level. Scapes distinctly longer than head (SI 120–149), with decumbent to subdecumbent pilosity and longer suberect hairs along outer edge. Pronotum in profile flatly convex, posterior promesonotal process well developed, angulate, and prominently produced. Metanotal groove not or barely impressed, with weak to inconspicuous cross-ribs. Propodeum in profile about as long as high or slightly longer with highest point immediately behind metanotal groove, declining smoothly toward spines. Propodeal spines short-triangular and acute, much shorter than distance between their bases (PSLI 14–19). Promesonotum largely smooth, except for superficial punctures anteriorly near the neck, remainder of mesosoma punctate to weakly punctate, often with scattered superficially sculptured to smooth areas. Metafemur relatively long (FI 134–170), metatibia pilosity mostly decumbent with longer suberect hairs along outer edge. Postpetiole in profile without ventral process, on average 1.1 times longer than high (LPpI 100–124), and on average 2 times wider and significantly shorter than petiole (PpWI 177–236, PpLI 58–70). Dorsum of petiole and postpetiole mostly smooth, rest weakly to superficially punctate. Gaster smooth and shiny. Standing hairs moderately scarce, short to moderately long, thin, and acute, suberect, some hairs subapically branched. Short decumbent to subdecumbent pilosity comparatively scarce. Color reddish to darker brown, with significantly darker head and gaster.



'''Wilson 2003. Upper: major. Lower: minor. CUBA: Hotel Telegrafo, Holguin, Oriente (lectotype and paralectotype of P. teneriffana subsp. taina Aguayo). Scale bars = 1 mm.'''

Etymology
Named after the place of origin of the types. (Wilson 2003)

References based on Global Ant Biodiversity Informatics

 * Azuma, S. and M. Kinjo. 1987. Family Formicidae, In Checklist of the insects of Okinawa. The Biological Society of Okinawa, Nishihara. Pages 310-312.
 * 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.
 * Bharti H., and A. Gill. 2011. SEM studies on immature stages of Pheidole indica Mayr, 1879 (Hymenoptera: Formicidae) from India. Halteres 3: 38-44.
 * 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
 * Chavhan A., and S.S. Pawar. 2011. Distribution and diversity of ant species (Hymenoptera: Formicidae) in and around Amravati City of Maharashtra, India. World Journal of Zoology 6(4): 395-400.
 * Chen Y. Q., Q. Li, Y. L. Chen, Z. X. Lu, X. Y. Zhou. 2011. Ant diversity and bio-indicators in land management of lac insect agroecosystem in Southwestern China. Biodivers. Conserv. 20: 3017-3038.
 * Collingwood C.A. 1970. Formicidae (Hymenopter: Aculeata) of Nepal. Himalaya Khumbu Himal, 3: 371-388.
 * Dias R. K. S. 2002. Current knowledge on ants of Sri Lanka. ANeT Newsletter 4: 17- 21.
 * Eguchi K. 2004. Taxonomic revision of two wide-ranging Asian ants, Pheidole fervens and P. indica (Insecta: Hymenoptera, Formicidae), and related species. Annalen des Naturhistorischen Museums in Wien. B, Botanik, Zoologie 105: 189-209
 * Eguchi, K. 2004. Taxonomic revision of two wide-ranging Asian ants, Pheidole fervens and P. indica (Insecta: Hymenoptera, Formicidae) and related species. Ann. Naturhist. Mus. Wien. 105B:189-209.
 * Emery C. 1901. Ameisen gesammelt in Ceylon von Dr. W. Horn 1899. Deutsche Entomologische Zeitschrift 1901: 113-122.
 * Forel A. 1885. Indian ants of the Indian Museum, Calcutta. J. Asiat. Soc. Bengal Part II Nat. Sci. 54: 176-182.
 * Forel A. 1902. Les Formicides de l'Empire des Indes et de Ceylan. Part IX. J. Bombay Nat. Hist. Soc. 14: 520-546.
 * Forel A. 1902. Myrmicinae nouveaux de l'Inde et de Ceylan. Rev. Suisse Zool. 10: 165-249.
 * Forel A. 1903. Les fourmis des îles Andamans et Nicobares. Rapports de cette faune avec ses voisines. Rev. Suisse Zool. 11: 399-411.
 * Forel A. 1906. Les fourmis de l'Himalaya. Bulletin de la Société Vaudoise des Sciences Naturelles 42: 79-94.
 * Forel A. 1909. Études myrmécologiques en 1909. Fourmis de Barbarie et de Ceylan. Nidification des Polyrhachis. Bull. Soc. Vaudoise Sci. Nat. 45: 369-407.
 * Forel A. 1911. Ameisen aus Ceylon, gesammelt von Prof. K. Escherich (einige von Prof. E. Bugnion). Pp. 215-228 in: Escherich, K. Termitenleben auf Ceylon. Jena: Gustav Fischer, xxxii + 262 pp.
 * Forel A. 1912. H. Sauter's Formosa-Ausbeute. Formicidae (Hym.) (Schluss). Entomol. Mitt. 1: 45-61.
 * Forel A. 1913. H. Sauter's Formosa-Ausbeute: Formicidae II. Arch. Naturgesch. (A)79(6): 183-202
 * Forel, A. 1908. Fourmis de Ceylan et d'Égypte récoltées par le Prof. E. Bugnion. Lasius carniolicus. Fourmis de Kerguelen. Pseudandrie? Strongylognathus testaceus. Bull. Soc. Vaudoise Sci. Nat. 44: 1-22
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Imai H. T., C. Baroni Urbani, M. Kubota, G. P. Sharma, M. H. Narasimhanna, B. C. Das, A. K. Sharma, A. Sharma, G. B. Deodikar, V. G. Vaidya, and M. R. Rajasekarasetty. 1984. Karyological survey of Indian ants. Japanese Journal of Genetics 59: 1-32.
 * Kaleeswaran B., S. Ezil, B. P. Ganesh, and S. Bhavatarini. 2008. Biodiversity and niches of Ants in Alagar hills, Tamil nadu Wildlife Biodiversity Conservation Published by Day publishing house pp 188-208.
 * Li Q., B. D. Hoffmann, Z. X. Lu, and Y. Q. Chen. 2017. Ants show that the conservation potential of afforestation efforts in Chinese valley-type savanna is dependent upon the afforestation method. Journal of Insect Conservation DOI 10.1007/s10841-017-0005-0
 * Li Qiao, Chen You-qing, Guo Xiao, Duan Yan, Chen Yan-lin, and Xu Zheng-hui. 2007. Diversity of ants in differents habitats in Yuanmou arid-hot valley, Yunnan. Journal of Fujian College of Forestry 27(3): 272-277.
 * 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.
 * Lu Y-F., and S. Y. Zhou. 2005. Karyotype analysis of four species in Formicidae. Guangxi Shifan Daxue Xuebao Ziran Kexue Ban 23: 81-84.
 * Lu Z., K. Li, N. Zhang, and Y. Chen. 2017. Diversity and indicator species of leaf-litter ants in Eucalyptus grandis plantations and secondary natural forests. Forest Research 29(4): 576-580
 * Mohanraj P., M. Ali, and K. Veerakumari. 2010. Formicidae of the Andaman and Nicobar Islands (Indian Ocean: Bay of Bengal). Journal of Insect Science 10: Article 172
 * Mohanraj, P., M. Ali and K. Veenakumari. 2010. Formicidae of the Andaman and Nicobar Islands (Indian Ocean: Bay Of Bengal). Journal of Insect Science 10:172.
 * Ogata K. 1982. Taxonomic study of the ant genus Pheidole Westwood of Japan, with a deszcription of a new species (Hymenoptera, Formicidae). Kontyu, Tokyo 50(2): 189-197.
 * Pan Y.S. 2007. Systematic Study on the Ant Genera Pheidole Westwood and Aphaenogaster Mayr (Hymenoptera: Formincidae : Myrmicinae) In China. Guangxi Normal University, Guangxi, China. 73 pages.
 * Pisarski B. 1967. Fourmis (Hymenoptera: Formicidae) d'Afghanistan récoltées par M. Dr. K. Lindberg. Annales Zoologici (Warsaw) 24: 375-425.
 * Presty J., and K. A. Karmaly. 2014. New records of three species of genus Pheidole Westwood (Hymenoptera: Formicidae) from Kerala. Journal of Zoology Studies 1(6): 01-10.
 * Rasheed M. T., I. Bodlah, A. G. Fareen, A. A. Wachkoo, X. Huang, and S. A. Akbar. 2019. A checklist of ants (Hymenoptera: Formicidae) in Pakistan. Sociobiology 66(3): 426-439.
 * Song Y., Z. H. Xu, C. L. Li, Y. Q. Hao, and H. B. Li. 2014. Ant communities of Nangun River Nature Reserve in Yunnan. Journal of West China Forestry Science 45(5): 93-100.
 * 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.
 * Terayama M. 2009. A synopsis of the family Formicidae of Taiwan (Insecta: Hymenoptera). Research Bulletin of Kanto Gakuen University. Liberal Arts 17:81-266.
 * Terayama Mamoru. 2009. A synopsis of the family Formicidae of Taiwan (Insecta, Hymenoptera). The Research Bulletin of Kanto Gakuen University 17: 81-266.
 * Terayama, M. 2009. A synopsis of the family Formicidae of Taiwan (Insecta; Hymenoptera). The Research Bulletin of Kanto Gakuen University 17: 81-266.
 * Thapa V. K. 2000. An Inventory of Nepal's Insects, Vol. III. IUCN Nepal, Kathmandu, xi + 475 pp.
 * 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.
 * Wang W. 2006. A taxonomic study of Formicidae ants from Hujiaping forest farm in Laifeng County. Journal of Hubei Institute for Nationalities (Natural Science Edition). 24(3): 298-300.
 * 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. 1929. Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 24: 27-64.
 * Wheeler W. M. 1930. A list of the known Chinese ants. Peking Natural History Bulletin 5: 53-81.
 * Wilson E. O. 1984. Tropical social parasites in the ant genus Pheidole, with an analysis of the anatomical parasitic syndrome (Hymenoptera: Formicidae). Insectes Sociaux 31: 316-334.
 * Wu Z. W., X. D. Bi, X. He, Z. X. Lu, L. J. Wei, and Y. Q Chen. 2015. Impact of continuous fire disturbance on ground-dwelling ant communities in arid-hot valleys of Panzhihua, Sichuan. Journal of Yunnan University 37(3): 467-474.
 * Yamane S.; Ikudome, S.; Terayama, M. 1999. Identification guide to the Aculeata of the Nansei Islands, Japan. Sapporo: Hokkaido University Press, xii + 831 pp. pp138-317.
 * Zhang N. N., Y. Q. Chen, Z. X. Lu, W. Zhang, and K. L. Li. 2013. Species diversity, community structure difference and indicator species of leaf-litter ants in rubber plantations and secondary natural forests in Yunnan, southwestern China. Acta Entomologica Sinica 56(11): 1314-1323.
 * Zhang R. J., L. W. Liang, and S. Y. Zhou. 2014. An analysis on the ant fauna of Nonggang Nature Reserve in Guangxi, China. Journal of Guangxi Normal university: Natural Science Edition 32(3): 86-93.
 * Zhang W., G. Liu, P. Zhong, and S. Zhang. 2014. Investigation of Formicidae in Luofushan Mountain. Journal of Huizhou University 34(3): 46-50.
 * Zhang Z., M. Cao, X. Yang, X. Deng, and Y. She. 2000. A study on species diversity of ant in fragments of seasonal rain forest of Xishuangbanna, China. Zoological Research 21(1): 70-75.
 * Zhou S. Y., and Z. M. Zheng. 1999. Taxonomic study of the ant genus Pheidole Westwood from Guangxi, with descriptions of three new species (Hymenoptera: Formicidae). Acta Zootaxonomica Sinica 24: 83-88.
 * Zhou S.-Y. and Zheng Z. 1999. Taxonomic study of the ant genus Pheidole Westwood from Guangxi, with descriptions of three new species (Hymenoptera: Formicidae). Acta Zootaxonomica Sinica24(1): 83-88.