The Japanese pavement ant, Tetramorium tsushimae, is one of the most commonly encountered ants in southern and central Japan. There are also introduced populations of this species found in a number of US states. In Japan this species is found in open land, nesting under stones (Japanese Ant Image Database).
- 1 Photo Gallery
- 2 Identification
- 3 Distribution
- 4 Biology
- 5 Castes
- 6 Nomenclature
- 7 References
- 8 References based on Global Ant Biodiversity Informatics
Identification Keys including this Taxon
Latitudinal Distribution Pattern
Latitudinal Range: 45.98333333° to 30.30894°.
- Source: AntMaps
Distribution based on Regional Taxon Lists
Distribution based on AntMaps
Distribution based on AntWeb specimens
Check data from AntWeb
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.
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.
This image is Tetramorium tsushimae, Japanese pavement ant, feeding on mooshy cracker fragment, in a play area, in a suburb of St. Louis, Missouri. Light colored individuals among the dark ones are typical of this species, which is a bit smaller than Tetramorium immigrans, (2.5-2.8 mm vs. 2.9-3.2 mm). Colonies are polygyne, polycalic. The species is native to Japan, the Korean Peninsula, and temperate China. USA population originated in Tokyo area, so says its DNA (James Trager).
Sanada-Morimura et al. (2006): Polygynous colonies of T. tsushimae are very common in central Japan where it builds subterranean nests in open grassland. Large colonies may consist of several hundred queens and millions of workers, with multiple nests that are probably connected underground with each other (Sanada-Morimura, pers. obs.). Their main food resources are seeds of grass plants and root aphids (Abe, 1971; Yamaguchi, pers. comm.). This study examined reproductive success and territoriality in a study population in a Mitaka, Japan chestnut field. Neighboring colonies were observed having large battles with many workers massed in a small area. These battles lasted for hours and would end with a general retreat by both colonies. Few dead workers were found once the battle ceased. In 9 out of 15 cases territory boundaries shifted as a result of the battle. Reproductive output was correlated with colony size, with the average territory size 32m2. Reproductive pupae growth-rate was strongly correlated with temperature, in lab assays being highest at temperature between 30 and 35°C. In field colonies, under artificial boards as nesting cover, reproductive brood was preferentially kept in areas with the highest field temperatures, from 27.5 to 30.0°C. It was also observed that larger colonies, with larger territories, had more aggregated nest entrances than those with small territories and a more evenly distributed set of nest entrances.
Sakata et al. (2017) examined how this ant uses cuticular hydrocarbons (CHCs) in recognizing aphid mutualists. Workers of the ants were obtained from a colony found on the campus of Chiba University in Matsudo, Japan. This colony had ~1,000 workers and 3 queens. Tetramorium tsushimae tends the aphid species Aphis craccivora in Japan, and it has been shown there is a correlation in decreasing aggressiveness of the ants towards the aphids with increasing familiarity, i.e., via interactions when collecting honeydew (Hayashi et al. 2015). Another aphid species, Acyrthosiphon pisum, that are also in the ant's environment are never tended. Using a combination of extracted aphid CHCs and synthetic standards, worker reactions towards whole aphid CHCs, methylalkane fractions, and n-alkanes were tested. The ants were less aggressive toward the CHC profile of A. craccivora and methylalkane fractions from this aphid. The ants reacted negatively toward CHC profiles from A. pisum and n-alkane. This suggests T. tsushimae is learning and recognizing aphid mutualists using CHCs and, in particular, methylalkanes.
Watanabe et al. (2019) conducted an observational study of the aphid species Macrosiphoniella yomogicola (Matsumura) and the ants that tended their colonies. This aphid has been found to be an obligate ant mutualist. Nine aphid colonies were monitored in Sapporo, Japan over a multi-week period in August and September. Lasius japonicus, Tetramorium tsushimae and Pheidole fervida were found attending the aphids, with each ant species exclusively tending the aphid colony where each was observed. The only aphid colonies to survive through the monitoring period were those tended by Lasius japonicus. This suggests, but the he number of observed colonies was not sufficient to conclude, that T. tsushimae is not a mutualist of the aphid and is only opportunistically tending the colonies where it was found.
Hosoishi et al. (2019) examined winter foraging activity in western Japan. They found foraging activity was generally low, except during relatively warm periods when the surface ground temperature was above 6–7°C or soil temperature was above 4–5°C. Tetramorium tsushimae, Messor aciculatus and Pheidole nodus were the most abundant in the open land type, whereas Nylanderia flavipes, Pheidole nodus and Crematogaster osakensis were the most abundant in the forest type.
Association with Other Organisms
- This species is a host for the ant Strongylognathus koreanus (a slave maker) in Japan (Collingwood, 1976).
- This species is a host for the cricket Myrmecophilus tetramorii (a myrmecophile) in Japan.
- This species is a host for the cestode Raillietina echinobothrida (a parasitoid) (Quevillon, 2018) (encounter mode secondary; indirect transmission; transmission outside nest; as Tetramorium caespitum jacoti).
- This species is a host for the cestode Raillietina tetragona (a parasitoid) (Quevillon, 2018) (encounter mode secondary; indirect transmission; transmission outside nest; as Tetramorium caespitum jacoti).
The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.
- tsushimae. Tetramorium caespitum subsp. tsushimae Emery, 1925c: 187, fig. C (w.q.) JAPAN. Synonym of jacoti: Wheeler, W.M. 1927d: 7. [Wheeler gives jacoti as senior synonym, but tsushimae has priority and is the first available name for this taxon: Bolton, 1995b: 415.] Raised to species: Bolton, 1995b: 415. [For current synonymy of tsushimae see under its junior synonym jacoti.]
- jacoti. Tetramorium caespitum subsp. jacoti Wheeler, W.M. 1927d: 7 (w.) CHINA. [First available use of Tetramorium caespitum subsp. semilaeve var. jacoti Wheeler, W.M. 1923b: 3; unavailable name.] Synonym of tsushimae: Wheeler, W.M. 1927d: 7. [Wheeler gives jacoti as senior synonym but tsushimae has priority and is first available name for this taxon: Bolton, 1995b: 415.] Raised to species: Pisarski, 1969b: 303; Radchenko, 1992b: 51. Senior synonym of tsushimae and material of the unavailable names chinensis, geei referred here: Radchenko, 1992b: 51.
- Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 415, [Wheeler gives jacoti as senior synonym, but tsushimae has priority and is the first available name for this taxon: Bolton, 1995: 415.] )
- Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 415, Raised to species)
- Chalissery, J.M., Gries, R., Alamsetti, S.K., Ardiel, M.J., Gries, G. 2021. Identification of the trail pheromone of the Pavement Ant Tetramorium immigrans (Hymenoptera: Formicidae). Journal of Chemical Ecology (doi:10.1007/s10886-021-01317-3).
- de la Mora, A., Sankovitz, M., Purcell, J. 2020. Ants (Hymenoptera: Formicidae) as host and intruder: recent advances and future directions in the study of exploitative strategies. Myrmecological News 30: 53-71 (doi:10.25849/MYRMECOL.NEWS_030:053).
- Emery, C. 1925d. Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum 183: 1-302 (page 187, fig. C worker, queen described)
- Hisasue, Y. 2018. Ant fauna of Matsuyama Castle. ARI 39: 18-36.
- Hisasue, Y. 2019. Ants collected from Akita Prefecture (Japan) in August 2017. ARI 40: 15-18.
- Hisasue, Y., Tsuji, Y. 2020. Records of Nylanderia amia (Forel) in Shikoku, with notes of range expansion of Japan. ARI 41: 18-36.
- Hosoishi, S., Rahman, M.M., Murakami, T., Park, S.-H., Kuboki, Y., Ogata, K. 2019. Winter activity of ants in an urban area of western Japan. Sociobiology 66(3): 414-419 (doi:10.13102/sociobiology.v66i3.4374).
- Imai, H.T., Kihara, A., Kondoh, M., Kubota, M., Kuribayashi, S., Ogata, K., Onoyama, K., Taylor, R.W., Terayama, M., Yoshimura, M., Ugawa, Y. 2003. Ants of Japan. 224 pp, Gakken, Japan.
- Iwata, K., Eguchi, K., Yamane, S. 2005. A case study on urban ant fauna of southern Kyusyu, Japan, with notes on a new monitoring protocol (Insecta, Hymenoptera, Formicidae). Journal of Asia-Pacific Entomology 8, 263-272.
- Kim, G., Lyu, D. 2012. Distribution of Ants (Insecta, Hymenoptera) in Chiaksan Mountain, Prov. Gangweon, Korea. Journal of Korean Nature 5, 127–129 (doi:10.7229/jkn.2012.5.2.127).
- Meurville, M.-P., LeBoeuf, A.C. 2021. Trophallaxis: the functions and evolution of social fluid exchange in ant colonies (Hymenoptera: Formicidae). Myrmecological News 31: 1-30 (doi:10.25849/MYRMECOL.NEWS_031:001).
- Moss, A.D., Swallow, J.G., Greene, M.J. 2022. Always under foot: Tetramorium immigrans (Hymenoptera: Formicidae), a review. Myrmecological News 32: 75-92 (doi:10.25849/MYRMECOL.NEWS_032:075).
- Nakabayashi, Y., Mochioka, Y., Tokuda, M., Ohshima, I. 2020. Mutualistic ants and parasitoid communities associated with a facultative myrmecophilous lycaenid, Arhopala japonica, and the effects of ant attendance on the avoidance of parasitism. Entomological Science 23, 233–244 (doi:10.1111/ENS.12417).
- Nakajima, T. 2018. Evaluation of ant diversity in urban areas and research on IPM. Japanese Journal of Environmental Zoology 29, 149–158.
- Park, S.-H., Hosoishi, S., Ogata, K. 2014. Long-term impacts of Argentine ant invasion of urban parks in Hiroshima, Japan. Journal of Ecology and Environment 37, 123–129 (doi:10.5141/ecoenv.2014.015).
- Park, S.-H., Hosoishi, S., Ogata, K., Kasuya, E. 2014. Changes of species diversity of ants over time: A case study in two urban parks. Journal of the Faculty of Agriculture, Kyushu University 59(1), 71–76.
- Park, S.-H., Hosoishi, S., Ogata, K., Kuboki, Y. 2014. Clustering of ant communities and indicator species analysis using self-organizing maps. Comptes Rendus Biologies 337, 545–552 (doi:10.1016/j.crvi.2014.07.003).
- Saito-Morooka, F., Fukuhara, K., Suda, K. 2015. The ant fauna of Rissho University at Kumagaya, Saitama Prefecture (Insecta, Hymenoptera, Formicidae). Bulletin of geo-environmental science (17), 35-39.
- Sakata, I., M. Hayashi, and K. Nakamuta. 2017. Tetramorium tsushimae ants Use Methyl branched hydrocarbons of aphids for partner recognition. Journal of Chemical Ecology. 43:966-970. doi:10.1007/s10886-017-0891-3
- Sanada-Morimura, S., T. Satoh, and Y. Obara. 2006. Territorial behavior and temperature preference for nesting sites in a pavement ant Tetramorium tsushimae. Insectes Sociaux. 53(2):141-148. doi:/10.1007/s00040-005-0849-2
- Seifert, B. 2021. Surviving the winter: Tetramorium sibiricum n. sp., a new Central Siberian ant species (Hymenoptera: Formicidae). Osmia 9, 15–24 (doi:10.47446/osmia9.3).
- Steiner, F.M., Schlick-Steiner, B.C., Trager, J.C., Moder, K., Sanetra, M., Christian, E., StauVer, C., 2006. Tetramorium tsushimae, a new invasive ant in North America. Biol. Invasions 8, 117–123.
- Sunamura, E., Tamura, S., Urano, T., Shoda-Kagaya, E. 2020. Predation of invasive red-necked longhorn beetle Aromia bungii (Coleoptera: Cerambycidae) eggs and hatchlings by native ants in Japan. Applied Entomology and Zoology 55, 291–298 (doi:10.1007/S13355-020-00681-Y).
- Takahashi, N., Touyama, Y., Kameyama, T., Ito, F. 2018. Effect of the Argentine ant on the specialist myrmecophilous cricket Myrmecophilus kubotai in western Japan. Entomological Science 21, 343–346 (doi:10.1111/ENS.12314).
- Tanaka, K., Tokuda, M. 2020. Negative correlation between dispersal investment and canopy openness among populations of the ant-dispersed sedge, Carex lanceolata. Plant Ecology. (doi:10.1007/S11258-020-01065-6).
- Watanabe, S., Y. Murakami, and E. Hasegawa. 2019. Effects of Attending Ant Species on the Fate of Colonies of an Aphid, Macrosiphoniella yomogicola (Matsumura) (Homoptera: Aphididae), in an Ant-Aphid Symbiosis. 128(4):325-332.
- Wheeler, W. M. 1927d. Chinese ants collected by Professor S. F. Light and Professor N. Gist Gee. Am. Mus. Novit. 255: 1-12 (page 7, Synonym of jacoti)
- Yamane, S., Hosoishi, S., Ito, F. 2022. Japanese Tetramorium queens: identiﬁcation key and species diagnoses (Hymenoptera, Formicidae, Myrmicinae). ZooKeys 1084: 43–64 (doi:10.3897/zookeys.1084.69767).
- Yu, Y. 2016. Risk of alien species introduction to Ogasawara Islands : Case study of ants at Tokyo Port. World Heritage Studies 1, 86-89.
- Zhu, W., Wu, L., Duan, L., Xu, S. 2022. A checklist of ants (Hymenoptera: Formicidae) in northern Shaanxi Province, China, with one new species of genus Proformica Ruzsky, 1902, Journal of Asia-Pacific Entomology 25, 101875 (doi:10.1016/j.aspen.2022.101875).
References based on Global Ant Biodiversity Informatics
- Azuma M. 1938. A list of ants found in Osaka Prefecture, Japan. Entomological World. Tokyo 6:238-243.
- Azuma M. 1951. On the Myrmecological fauna of Osaka Prefecture, Japan with description of new species (Formicidae, Hymenoptera). Hyogo Biology 1(5): 1-5.
- Azuma M. 1953. On the myrmecological fauna of Mt. Rokko, Hyogo Prefecture. Warera 2:1-7.
- 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
- Collingwood C. A. 1962. Some ants (Hym. Formicidae) from north-east Asia. Entomologisk Tidskrift 83: 215-230.
- Fukumoto S. 2017. Records of ants (Hymenoptera: Formicidae) from Uji-shima, Uji Islands, Kagoshima, Japan. Nature of Kagoshima 43: 295–296.
- Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
- Harada Y., D. Fukukura, R. Kurisu, and S. Yamane. 2013. Ants of Ports, monitoring of alien ant species. Bull. Biogeogr. Soc. Japan 68: 29-40.
- 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. Asai, M. Araba, T. Higasayama, and N. Saito. 2019. Ant fauna at ports on the Goto Islands – monitoring of alien ant species –. Nature of Kagoshima 46: 27–32.
- 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: 111121.
- 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.
- 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.
- Hosaka T., L. Di, K. Eguchi, and S. Numata. 2019. Ant assemblages on littered food waste and food removal rates in urban–suburban parks of Tokyo. Basic and Applied Ecology 37: 1–9.
- Hosoichi S., M. M. Rahman, T. Murakami, S. H. Park, Y. Kuboki, and K. Ogata. 2019. Winter activity of ants in an urban area of western Japan. Sociobiology 66(3): 414-419.
- Hosoichi S., W. Tasen, S. H. Park. A. Le Ngoc, Y. Kuboki, and K. Ogata. 2015. Annual fire resilience of ground-dwelling ant communities in Hiraodai Karst Plateau grassland in Japan. Entomological Science 18: 254–261.
- 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.
- Huang Jian-hua, and Zhou Shan-yi. 2007. Checklist of Family Formicidae of China - Myrmicinae (Part III). (Insecta; Hymenoptera). Journal of Guangxi normal University: Natural Science Edition 25(3): 88-96.
- 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.
- Iwata, Kouki, Eguchi, Katsuyuki and Yamane, Seiki. 2005. A Case Study on Urban Ant Fauna of Southern Kyusyu, Japan, with Notes on a New Monitoring Protocol (Insecta, Hymenoptera, Formicidae). Asia-Pacific Entomol. 8(3):263-272.
- Kuboki Y., K. Ogata, and E. Kasuya. 2006. Note on the bait trapping method. ANeT Newsletter 8: 16-19
- 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.
- Lelej A. S. 2012. Annotated catalogue of the Insects of Russian Far East. Volume 1. Hymenoptera. Dalnauka: Vladivostok. 635 p.
- 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.
- Matsumura S. and Yamane Sk. 2012. Species composition and dominant species of ants in Jigenji Park, Kagoshima City, Japan. Nature of Kagoshima 38: 99107
- 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.
- Nishimuta K., T. Yamaguchi, and Y. Harada. 2013. Ants of ports-monitoring of alien ant species. 9th International ANeT Conference, Kota Kinabalu, Malaysia, 28th October - 1st November 2013.
- Park S. H., S. Hosoishi, K. Ogata, and E. Kasuya. 2014. Changes of Species Diversity of Ants Over Time: a Case Study in Two Urban Parks. J. Fac. Agr., Kyushu Univ., 59 (1), 7176.
- 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
- Pfeiffer M., L. Chimedredzen, and K. Ulykpan. 2003. Community organization and species richnees of ants (Hymenoptera/ Formicidae) in Mongolia along an ecological gradient from Steepe to Gobi desert. Journal of Biogeography 30: 1921-1935.
- Pisarski B. 1969. Fourmis (Hymenoptera: Formicidae) de la Mongolie. Fragmenta Faunistica (Warsaw). 15: 221-236.
- 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.
- Santschi F. 1925. Contribution à la faune myrmécologique de la Chine. Bulletin de la Société Vaudoise des Sciences Naturelles 56: 81-96.
- Santschi F. 1928. Nouvelles fourmis de Chine et du Turkestan Russe. Bulletin et Annales de la Société Entomologique de Belgique. 68: 31-46.
- Shimana Y., and S. Yamane. 2009. Geogrpahical distribution of Technomyrmex brunneus Forel (Hymenoptera, Formicidae) in the western part of the mainland of Kagoshima, South Kyushu, Japan. Ari 32: 9-19.
- Stitz H. 1934. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sü Ping-chang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927-1930. 25. Hymenoptera. Ark. Zool. 27A(1) 11: 1-9.
- 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. 2005. Ants from the Akasaka Imperial Gardens, Tokyo. Mem. Natn. Sci. Mus., Tokyo 39: 239-243.
- Terayama M., S. Kubota, and K. Eguchi. 2014. Encyclopedia of Japanese ants. Asakura Shoten: Tokyo, 278 pp.
- Terayama M., and R. Sonobe. 2002. Ants from the Nasu Imperial Villa, Tochigi Prefecture, Japan. Flora and fauna of the Tochigi Prefectural Museum research report Nasu Imperial Villa 157-161.
- Terayama M., and S. Kubota. 2002. Ants of Tokyo, Japan. ARI 26: 1-32.
- Terayama. M. 2004. Geological and ecological distribution of Japanese ants communities. (translated from Japanese) Reports of the Saitama Prefecture Animal Research Association. 48:30
- Terayama. M. 2004. Geological and ecological distribution of Japanese ants communities. (translated from Japanese) Reports of the Saitama Prefecture Animal Research Association. 48:31
- Togashi I., and M. Terayama. 2010. Ant fauna of Hegurajima and Nanatsujima islands, Ishikawa Prefecture. Bull. Biogeoger. Soc. Japan 65: 195-196.
- Wheeler W. M. 1927. Chinese ants collected by Professor S. F. Light and Professor N. Gist Gee. American Museum Novitates 255: 1-12.
- 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. 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. Some ants from China and Manchuria. American Museum Novitates 361: 1-11.
- Wheeler W. M. 1930. A list of the known Chinese ants. Peking Natural History Bulletin 5: 53-81.
- Yamane S. 2007. Ants of Mongolia. The Nature and Insects. 42: 20-25
- Yamane S. 2019. Seasonal change in the foraging activity of ants in a residential area of mainland Kagoshima, Southwest Japan (Insecta, Hymenoptera, Formicidae). Nature of Kagoshima 45: 361–366.
- Yamane S. 2019. Seasonal change in the foraging activity of ants in a residential area of mainland Kagoshima, Southwest Japan (Insecta, Hymenoptera, Formicidae). Nature of Kagoshima 45: 361–366.
- 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 K. Eguchi. 2013. Classification and ecology of ants. Natural history of ants in Southern Kyushu. 200 pages
- 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.