Vollenhovia emeryi

Vollenhovia emeryi has colonies of two types: Type 1 colonies are usually monogynous with alate females, the reproductive offspring of which are also alate females; Type 2 colonies are usually polygynous and have short-winged reproductive females, the reproductive female offspring of which are also short-winged. Type 1 colonies reproduce sexually and are found mainly in montain forests, nesting in decaying wood. Type 2 colonies reproduce clonally and are found in or near riverside forests, also nesting in decaying wood. In both types the alate females and males hibernate within the nests. Vollenhovia nipponica is a parasite frequently found in Type 2 colonies of V. emeryi.

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Oriental Region: Taiwan, Thailand. Palaearctic Region: China, Democratic Peoples Republic of Korea, Japan, Republic of Korea.

Biology
Miyakawa et al. (2018) studied the details of genetic control of sex deterimination in this species. They concluded the doublesex (dsx) gene appears to integrate signal from a multi-locus complementary sex determination system to control if females or males develop from an egg.

Reproduction
This species has two modes of reproduction. Clonally produced, homozygous queens have short malformed wings. Long-winged dispersing queens, with heterozygous genomes and so not produced clonally, can be produced facultatively. Queens eclose before fall hibernation. Clonal queens mate in their natal nest before hibernation. In the spring new colonies can be formed by budding. Laboratory colonies collected from field sites in Japan and provided with more food than a second set of colonies produced long-winged queens, non-dispersing queens and more queens overall. The colonies that were fed half as much food only produced clonal short-winged queens. Laboratory produced long-winged queens were not inseminated after eclosing while the short-winged queens were all found to be inseminated. Presumably the long-winged queens mate after hibernation in the spring (Okomoto et al. 2015, Ohkawara et al. 2002).

Genetics
The genome of V. emeryi was sequenced for a study examining how parasitic ants with no worker caste may have altered their genome to arrive at a workerless state. In comparison to ants with a full complement of castes, there appeared to be no loss of genes in the parasitic ants. This suggests regulatory differences and not sequence differences predominate in gains and losses of castes (phenotypes). (Smith et al. 2015)

Castes
Both winged and brachypterous queens are known (Ohkawara et al. 2006)

Nomenclature

 *  emeryi. Vollenhovia emeryi Wheeler, W.M. 1906c: 312, pl. 41, figs. 10, 11 (w.q.) JAPAN. Wheeler, W.M. 1928d: 112 (m.); Imai, 1966: 119 (k.). Senior synonym of chosenica: Radchenko, 2005b: 154.
 * chosenica. Vollenhovia emeryi subsp. chosenica Wheeler, W.M. 1928d: 113 (w.) KOREA. Junior synonym of emeryi: Radchenko, 2005b: 154.

References based on Global Ant Biodiversity Informatics

 * Anonymous. A list of ants collected at Bandai, Fukushima Prefecture by the members of the Myrmecological Society of Japan in 1992. ARI Reports of the Myrmecologists Society (Japan) 18: 31
 * 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
 * Choi B.-M. 1987. Taxonomic study on ants (Formicidae) in Korea (1). On the genus Monomorium. Journal of the Institute of Science Education (Cheongju National Teachers' College) 11:17-30.
 * Choi B.M. 1985. Study on distribution of ants (Formicidae) from Korea (1). Formic fauna in Mt. Songni. Cheongju Sabom Taehak Nonmunjip (Journal of Cheongju National Teachers' College) 22:401-437.
 * Choi B.M. 1986. Studies on the distribution of ants (Formicidae) in Korea. Journal of Chongju National Teacher College 23: 317-386.
 * Choi B.M. 1988. Studies on the distribution of ants (Formicidae) in Korea (5) Ant fauna in Is. Kanghwado. Chongju Sabom Taehak Nonmunjip (Journal of Chongju National Teacher' College) 25: 217-231.
 * Choi B.M. 1996. Distribution of ants (Formicidae) in Korea (16) - Ant fauna from Chollabukdo. Korean J. Soil. Zoology 1(1): 5-23.
 * Choi B.M. 1996. Distribution of ants (Formicidae) in Korea (16): Ant fauna from Chollabukdo. Korean Journal of Soil Zoology 1(1): 5-23.
 * Choi B.M. 1996. Studies on the distribution of ants (Formicidae) in Korea (15) -Ant fauna islands Ullungdo and Dokdo. Journal of Chongju National University of Education 33: 201-219.
 * Choi B.M. 1998. Distribution of ants (Formicidae) in Korea (19) Ants fauna from Chungcheongbukdo Province. Cheongju Tea. Coll. 35: 213-266.
 * Choi B.M., Bang, J.R. 1992. Studies on the distribution of ants (Formicidae) in Korea (9). Ant fauna in Mt. Togyusan. Korean Journal of Applied Entomology 31:101-112.
 * Choi B.M., I. H. Lee. 1995. Studies on the distribution of ants (Formicidae) in Korea (14). Ant fauna in island Sohuksando. Korean Journal of Applied Entomology 34(3): 191-197.
 * Choi B.M., K. Ogata, and M. Terayama. 1993. Comparative studies of ant faunas of Korea and Japan. 1. Faunal comparison  among islands of Southern Korean and northern Kyushu, Japan. Bull. Biogeogr. Soc. Japan 48(1): 37-49.
 * Choi B.M., Kim, C.H., Bang, J.R. 1993. Studies on the distribution of ants (Formicidae) in Korea (13). A checklist of ants from each province (Do), with taxonomic notes. Cheongju Sabom Taehakkyo Nonmunjip (Journal of Cheongju National University of Education) 30: 331-380.
 * Choi B.M., and H.S. Lee. 1999. Studies on the distribution ants in Korea (21) - Ant fauna in Kwanaksan. Korean J. Soil Zoology 4(1): 1-4.
 * Choi B.M., and J. R. Bang. Studies on the distribution of ants (Formicidae) in Korea (12): the analysis of ant communities in 23 islands. Journal of Cheongju National University of Education 30:317-330.
 * Eto S., and K. Ogata. 1983. Ants of Hirado Island, Kyushu. Bulletin of the Nagasaki Prefecture Biological Group 25: 7-11.
 * Fontanilla A. M., A. Nakamura, Z. Xu, M. Cao, R. L. Kitching, Y. Tang, and C. J. Burwell. 2019. Taxonomic and functional ant diversity along tropical, subtropical, and subalpine elevational transects in southwest China. Insects 10, 128; doi:10.3390/insects10050128
 * Forel A. 1912. H. Sauter's Formosa-Ausbeute. Formicidae (Hym.) (Schluss). Entomol. Mitt. 1: 45-61.
 * Fukumoto S., Jaitrong W. and Yamane S.K. 2013. Ant Fauna of Kuro-shima, Iwo-jima and Take-shima islands, Kagoshima Prefecture, southwestern Japan. Nature of Kagoshima 39: 119-125
 * Fukumoto S., W. Jaitrong, and S. Yamane. 2013. Ant fauna of Take-shima, Iwo-jima and Kuro-shima islands, Kagoshima Prefecture, southwestern Japan. Nature of Kagoshima 39: 99-105.
 * Guenard et al. 2013 - Collection XTBG, Yunnan, China
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Ha S.J, S.J. Park, and B.J. Kim. 2002. Comparative ant faunas between Seonyudo and seven other islands of West Sea in Korea. Korean Journal of Entomology 32(2): 75-79.
 * Harada Y. 1997. Ants from the Koshiki islands, Kagoshima-ken, southern Japan. Ari 21: 1-4.
 * Harada Y. 2000. Ant fauna of the forest floor of the Koshikijima Islands, Kagoshima-ken, southern Japan. Ari 24: 4-11.
 * Harada Y. S. Koto, N. Kawaguchi, K. Sato, T. Setoguchi, R. Muranaga, H. Yamashita, A. Yo, and S. Yamane. 2012. Ants of Jusso, Isa City, Kagoshima Prefecture, southwestern Japan. Bull. biogeogr. Soc. Japan 67: 143-152.
 * Harada Y., H. Yadori, M. Yoneda, R. Takinami, K. Nagahama, Y. Matsumoto, A. Oyama, S. Maeda, and S. Yamane. 2009. Ant fauna of Tanegashima (Hymenoptera, Formicidae). Nankiseibutu, the Nanki Biological Society 51(1): 15-21.
 * Harada Y., K. Nishikubo, K. Matsumoto, M. Matsuda, Y. Inazawa, Y. Ozono, S. Koto, N. Kawaguchi, and S. Yamane. 2011. Ant fauna of Japanese beech (Fagus crenata) forests in southwestern Japan. Bull. Biogeogr. Soc. Japan 66: 115-127.
 * Harada Y., K. Tashiro, K. Ebihara, H. Yadori, M. Yoneda, R. Takinami, K. Nagahama, and K. Hayashi. 2008. Ant fauna of the lavas of Sakurajima Volcano, Southern Japan. Bull. Biogeogr. Soc. Japan 63: 205-215.
 * Harada Y., M. Enomoto, N. Nishimata, and K. Nishimuta. 2014. Ants of the Tokara Islands, northern Ryukyus, Japan. Nature of Kagoshima 40: 111121.
 * Harada Y., S. Haruguchi, T. Iwasaki, K. Onishi, Y. Tashiro, and Sk Yamane. 2010. Ants from Japanese cherry trees, Prunus x yedoensis, in public parks in Kagoshima, southwestern Japan. Bull. Biogeogr. Soc. Japan 65: 169-179.
 * Harada Y., Y. Matsumoto, S. Maeda, A. Oyama, and S. Yamane. 2009. Comparison of ant fauna among different habitats of Yaku-shima Island, southern Japan. Bull. Biogeogr. Soc. Japan 64: 125-134.
 * Harada Y., Yadori H., Takinami R., Nagahama K., Matsumoto Y., Oyama A., Maeda S. and Yamane S.K. 2013. Ants of the southernmost Fagus crenata forest in Japan. Nature of Kagoshima 39: 113-118
 * Hayashida K. 1957. Ecological distribution of ants in Sapporo and vicinity. (Preliminary report.). Journal of the Faculty of Science, Hokkaido University. Series VI. Zoology 13:173-177.
 * Hayashida K. 1961. Studies on the ecological distribution of ants in Sapporo and its vicinity (1 et 2). Insectes Sociaux 7: 125-162.
 * Hisamatsu M. 2004. List of Hymenoptera Recorded in Ibaraki Prefecture. Bulletin of Ibaraki Nature Museum 7: 125-164.
 * Hosoichi S., M. Yoshimura, Y. Kuboki, and K. Ogata. 2007. Ants from Yakushima Island, Kagoshima Prefecture. Ari 30: 47-54.
 * Hosoishi S. 2006. Ant fauna of Noko Island. pp99-107. In: The floristic and faunistic surveys of the Noko Island.
 * Hosoishi S., M. Yoshimura, Y. Kuboki, and K. Ogata. 2007. Ants from Yakushima Island, Kagoshima Prefecture. Ari 30: 47-54.
 * Hua Li-zhong. 2006. List of Chinese insects Vol. IV. Pages 262-273. Sun Yat-sen university Press, Guangzhou. 539 pages.
 * 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.
 * Ichikawa A. 1999. Records of ants observed from several localities of Osaka Prefecture, Japan, -1. Ari 23: 1-3.
 * Ikeshita Y., A. Gotoh, K. Yamamoto, N. Taniguchi, and F. Ito. 2007. Ants collected in Mt. Linoyama, Marugame, Kagawa Prefecture (Hymenoptera, Formicidae). Kagawa Seibutsu 34: 59-62.
 * Kawahara Y., S. Hosoyamada, and S. Yamane. 1999. Ant fauna of the Terayama Station for Education and Research on Nature, Kagoshima University. Bulletin of the Faculty of Education, Kagoshima University. Natural Science 50: 147-156.
 * Kim B.J. 1996. Synonymic list and distribution of Formicidae (Hymenoptera) in Korea. Entomological Research Bulletin Supplement 169-196.
 * Kim B.J., K.G. Kim, J.H. Kim, S.J. Park. Ants from Mt. Mirok. Korean J. Soil Zoology 2(2): 115-128.
 * Kim B.J., S.J. Park, and J.H. Kim. 1996. Ants from Naejangsan national park (Hymenoptera: Formicidae). Korean J. Soil. Zoology &(2): 120-133.
 * Kim K.I., C.H. Kim, and B. Choi. 1989. The ant fauna of the southern shore in Gyeongsangnamdo, Korea. Journal of Gyeongsang Nat. Univ. 28(2): 213-226.
 * Kim et al. 1993. Systematic study of ants from Chejudo Province. Koran Journal of Entomology 23(3): 117-141.
 * Kobayashi K., E. Hasegawa, and K. Ohkawara, K. 2008. Clonal reproduction by males of the ant Vollenhovia emeryi (Wheeler). Entomological Science 11: 167-172.
 * Kobayashi K., H. Tamura, M. Okamoto, E. Hasegawa, K. Ohkawara. 2012. Phylogenetic relationships among populations of Vollenhovia ants, with particular focus on the evolution of wing morphology. Annals of the Entomological Society of America 105(3):454-461.
 * Kondoh M., and Y. Kitazawa. 1984. Ant communities of the campus of UOEH and in an adjacent natural forest. Journal of UOEH 6(3): 221-234.
 * Kubota. S., and M. Terayama. 1988. Ant fauna of Tokyo. (1) A list of ants collected at the parks. ARI Reports of the Myrmecologists Society (Japan) 16: 14-16
 * Kwon T. S. 2012. Korean ant atlas. Korea Forest Research Institute 162 pages.
 * Kwon T. S. 2015. Ant assemblages along the Baekdudaegan Mountain Range in South Korea: Human roads and temperature niche. Journal of Asia-Pacific Biodiversity 8: 152-157.
 * Kwon T. S. 2018. High competition between ant species at intermediate temperatures. Journal of Thermal Biology 72: 59-66.
 * Kwon T. S., S. S. Kim, and J. H. Chun. 2014. Pattern of ant diversity in Korea: An empirical test of Rapoport's altitudinal rule. Journal of Asia-Pacific Entomology 17: 161167.
 * Kwon T.S., C. M. Lee, J. H. Chun, J. H. Sung, and S. K. Kim. 2011. Ants in Hongneung forest. Korea Forest Research Institute, 92 pages.
 * Li Q., Z. Lu, Z. Wei, M. Yanyan, and F. Ping. 2015. Communities of ground-dwelling ants in different plantation forest in arid-hot valleys of Jinsha river, Yunnan Province, China. Scientia Silvae Sinicae 51(8): 134-142.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * Lyu D.P. 2003. Systematics of Myrmicinae from Korea (Hymenoptera: Formicidae). PhD thesis Faculty of the Graduate School of Chungbuk National University 330 pages.
 * Maeto K. and S. Sato. 2004. Impacts of forestry on ant species richness and composition in warm-temperate forests of Japan. Forest Ecology and Management 187: 213223.
 * Manabe K. 1994. Ants of the shrine forest in Fukuoka Prefecture (First report; Ants of lowland). Ari 17: 6.
 * Manabe K. 1994. Ants of the shrine forest in Fukuoka Prefecture (first report; ants of lowland). Ari 18: 6.
 * Manabe K. 1994. Ants of the shrine forest in Fukuoka Prefecture (second report; ants of mountain zone). Ari 17: 8.
 * Masuko K. 1982. A data on the myrmecofauna of Miyake Island. Ari 10: 1-2.
 * Matsumura S. and Yamane Sk. 2012. Species composition and dominant species of ants in Jigenji Park, Kagoshima City, Japan. Nature of Kagoshima 38: 99107
 * Matsumura S., and S. Yamane. 2012. Species composition and dominant species of ants in Jigenji Park, Kagoshima City, Japan. Nature of Kagoshima 38: 99-107.
 * Miyama D., S. Yamane, T. Hishida, T. Kyono, T. Saito, T. Kuwahara, and N. Inoue. 2007. Ant Fauna in a Coppice Area around Shishitsuka-Ohike in Tsuchiura, Central Japan (Hymenoptera, Formicidae). Bulletin of Ibaraki Nature Museum 10: 1-10.
 * Morisita M. 1945. Ants of the southern part of Hokkaido, Japan. [In Japanese.] Mushi 16:21-28.
 * Ochi K. 1983. Distribution pattern of ants in pine stands, with special reference to Monomorium nipponense Wheeler (Hymenoptera: Formicidae). Gensei 44:1-6.
 * Ogata K. 1981. The ant fauna of the Goto islands, Natural history of the Goto? Islands, Japan : Iki Tsushima to no taihi (Danjo Gunto? Ko?rai Sone o fukumu Japan: 347-351.
 * Ogata. K., Touyama, Y. and Choi, B. M. 1994. Ant fauna of Hiroshima Prefecture, Japan. ARI Reports of the Myrmecologists Society (Japan) 18: 18-25
 * Ohkawara K., and Y. Fukushima. 1998. A Preliminary report of ant fauna in Ishikawa Prefecture, Central Japan. Ari 22: 6-9.
 * Paik W.H. 1984. A checklist of Formicidae (Hymenoptera) of Korea. Korean J. Plant Prot. 23(3): 193-195.
 * Park S. H., S. Hosoishi, K. Ogata, and Y. Kuboki. 2014. Clustering of ant communities and indicator species analysis using self-organizing maps. Comptes Rendus Biologies http://dx.doi.org/10.1016/j.crvi.2014.07.003
 * Park S.J., and B.J. Kim. 2002. Faunal comparison of ants among Cheongsando and other islands of South Sea in Korea. Korean Journal of Entomology 32(1): 7-12.
 * Park, Seong, Joon and Byung, and Kim, Jin. 2002. Faunal Comparison of Ants among Cheongsando and Other Islands of South Sea in Korea. Korean Jornal of Entomology. 32(1):7-12.
 * Radchenko, A. 2005. Monographic revision of the ants (Hymenoptera: Formicidae) of North Korea. Annales Zoologici (Warsaw) 55: 127-221.
 * Radchenko, A. 2005. Monographic revision of the ants (Hymenoptera, Formicidae) of North Korea. Annales Zoologici 55(2): 127-221.
 * Sakai H. 2002. Reproductive flight season of Japanese ants. Ari 26: 33-39.
 * Skarbek C. J., M. Noack, H. Bruelheide, W. Hardtle, G. von Oheimb, T. Scholten, S. Seitz, M. Staab. 2019. A tale of scale: plot but not neighbourhood tree diversity increases leaf litter ant diversity. Journal of Animal Ecology DOI: 10.1111/1365-2656.13115
 * So, Ha, Seong, Jin, Park, Joon and Byung, and Kim, Jin. 2002. Comparitive Ant Faunas between Seonyudo and Seven other Islands of West Sea in Korea. Korean Journal of Entomology. 32:75-79.
 * Sonobe R. 1977. Ant fauna of Miyagi prefecture, Japan. Japanese Journal of Ecology 27: 111-116.
 * Takechi F. 1960. A list of ants unrecorded from Mt. Ishizuchi and Omogo Valley, Iyo, Shikoku (Hymenoptera: Formicidae). Transactions of the Shikoku Entomological Society 6:91.
 * Tanaka B., N. Kanie, T. Mano, R. Arita, and A. Shiragane. 1999. The insect fauna between Takabashi bridge and Nomi-kouen Park, margins of the Yahagigawa river. ????? 3: 35-79.
 * Terayama M. 1977. Checklist of the known ants of Saitama Prefecture. Insects and nature 12(4): 26-27
 * Terayama M. 1983. Kagoshima-ken-hondo no ari. Kanagawa-chucho (Journal of the Kanagawa Entomologists Association): 13-24.
 * Terayama M. 1992. Structure of ant communities in East Asia. A. Regional differences and species richness. Bulletin of the Bio-geographical Society of Japan 47: 1-31.
 * Terayama M. 1994. Ants of Okushiri-to Island, Hokkaido. Ari 18: 28-29.
 * Terayama M. 2001. Ants of the Institute for Nature study in Minato-ku, Tokyo: species composition, relative nest abundance, and nest density. Rept. Inst. Nat. Stu. 33: 289-300.
 * Terayama M., K. Ogata, and B.M. Choi. 1994. Distribution records of ants in 47 prefectures of Japan. Ari (report of the Myrmecologists Society of Japan) 18: 5-17.
 * Terayama M., S. Kubota, and K. Eguchi. 2014. Encyclopedia of Japanese ants. Asakura Shoten: Tokyo, 278 pp.
 * Terayama M., and K. Murata. 1987. Relation between ant communities and vegetations of Toshima island, the Izu Islands. Bull. Biogeogr. Soc. Japan 42(9): 57-63.
 * Terayama M., and K. Murata. 1990. Effects of area and fragmentation of forests for nature conservation: Analysis by ant communities. Bull. Biogeogr. Soc. Japan 45(2): 11-17.
 * Terayama M., and K. Murata. 2000. Ants on Saru-shima Island, Kanagawa Pref., Japan. Ari 24: 1-3.
 * Terayama M., and S. Kubota. 1994. Ants from Aogashima Island. Ari 17: 11.
 * Terayama M., and S. Kubota. 2002. Ants of Tokyo, Japan. ARI 26: 1-32.
 * Terayama M., and S. Yamane. 1984. Ants of Yaku-shima Island, the northern Ryukyus, with reference to their altitudinal distribution (Insecta: Hymenoptera). Cons. Rep. Yaku-shima Wildness Area, Kyushu, Japan, pp. 643-667. Nat. Cons. Bureau, Env. Agency, Japan.
 * Terayama. M. 1994. Ants of Okushiri-to Island, Hokkaido. ARI Reports of the Myrmecologists Society (Japan) 18: 28-29
 * Terayama. M. 2004. Geological and ecological distribution of Japanese ants communities. (translated from Japanese) Reports of the Saitama Prefecture Animal Research Association. 48:34
 * Teruyama. M. 1988. Ant fauna of Saitama Prefecture, Japan. ARI Reports of the Myrmecologists Society (Japan) 16: 4-13
 * Tian M., L. Deharveng, A. Bedos, Y. Li, Z. Xue, B. Feng, and G. Wei. 2011, Advances of cave biodiversity survey: a result based mainly on invertebrates. Proceedings of the 17th National Congress of Speleology, Yinshuidong, Hubei, 1-3 Nov 20111, p 149-163.
 * Wheeler W. M. 1906. The ants of Japan. Bulletin of the American Museum of Natural History 22: 301-328.
 * Wheeler W. M. 1928. Ants collected by Professor F. Silvestri in Japan and Korea. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 22: 96-125.
 * 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.
 * Xu Z., Zeng G., Liu T.-Y. and He Y.-F. 1999. [A study on communities of Formicidae ants in different subtypes of vegetation in Xishuangbanna District of China.] Zoological Research 20: 118-125
 * Yamane S. 2016. How many species of Ants in Amami Islands? (in Japanese). Part 2, chapter 1 in How many species of Ants in Amami Islands? Pp. 92-132.
 * Yamane S., S. Ikudome, and M. Terayama. 1999. Identification guide to the Aculeata of the Nansei Islands, Japan. Sapporo: Hokkaido University Press, xii + 831 pp. pp, 138-317.
 * Yamane S., Y. Harada and M. Yano. 1985. Ant fauna of Tanega-shima Island, the northern Ryukyus (Hymenoptera, Formicidae). Mem. Kagoshima Univ. Res. Center S. Pac. 6(1): 166-173.
 * Yamane S., Y. Harada, and K. Eguchi. 2013. Classification and ecology of ants. Natural history of ants in Southern Kyushu. 200 pages
 * Yamane S., and S. Ikudome. 2008. Ants, wasps and bees of Kuro-shima Northern Ryukyus, Japan (Hymenoptera, Aculeata). Bull. Inst. Minami-Kyushu Reg. Sci. (Kagoshima Women's Jr. Coll.) 24: 1-9.
 * 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.
 * Yamazaki Y., S. Yamane, T. Hishida, T. Kuwahara, and N. Inoue. 2009. Ant fauna on the grounds of the Kashima-Jingu Shrine, Ibaraki, Central Japan (Hymenoptera, Formicidae). Bull. Ibaraki Nat. Mus. 12: 5-14.
 * Yoshitomi H., and S. Matsuno. 2012. List of species of Hymenoptera and Diptera in Matsuyama City, Ehime Prefecture, Shikoku, Japan. pp. 167-176. In: Committee for Surveys of Natural Environment of Matsuyama City (Chief Editor: Kazuo ISHIKAWA) (ed.) Checklist of the Wild Animals, Fungi, and Plants of Matsuyama City, 2012. Published by the Department of Environment, Matsuyama City, 404 pp.
 * Zhou S.-Y. 2001. Ants of Guangxi. Guangxi Normal University Press, Guilin, China, Guilin, China. 255 pp.