Ponera scabra

AntWiki: The Ants --- Online
Jump to navigation Jump to search
Ponera scabra
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Ponerinae
Tribe: Ponerini
Genus: Ponera
Species: P. scabra
Binomial name
Ponera scabra
Wheeler, W.M., 1928

Ponera scabra P casent0281908.jpg

Ponera scabra D casent0281908.jpg

Specimen Label

Common Name
Language: Japanese
(as Ponera yakushimensis)
Language: Japanese

This species nests in the soil as well as under moss. The larvae spin cocoons (Okamoto, 1972). Not rare in some localities within Japan. (Japanese Ant Image Database)

At a Glance • Facultatively polygynous  



Leong et al. (2019) - Worker: This species is characterized by a long antennal scape; a clypeus with an acute median tooth; a petiolar node in lateral view thick with straight anterior and posterior margins. Ponera scabra presents similarities with Ponera rishen, Ponera kohmoku, and Ponera takaminei. Ponera scabra can be distinguished from Ponera takaminei by its indistinct metanotal groove, while distinct in Ponera takaminei. Ponera scabra is distinct from Ponera kohmoku by its small eyes consisting of 5–7 facets, but about 20 facets in Ponera kohmoku. Ponera scabra is distinct from Ponera takaminei by the straight posterior margin of petiolar node in lateral view, while convex in Ponera takaminei.

Taylor (1967) - A Japanese species closely related to, and possibly cognate with, Ponera chapmani. Distinguished in general from other Ponera species by the combination of moderately large size (head width 0.61-0.64 mm in worker, 0.71-0.74 mm in queen), heavy sculpturing, and long scapes, which almost exactly reach the median occipital border. Readily differentiated from the only other known Japanese species, Ponera japonica, by its larger size (head width 0.42-0.50 in japonica) and heavier sculpturation. P. scabra may be differentiated from the related chapmani by characters stated below in the worker description.

Keys including this Species


Latitudinal Distribution Pattern

Latitudinal Range: 38.30444444° to 30.359°.

Tropical South

Distribution based on Regional Taxon Lists

Palaearctic Region: China, Democratic Peoples Republic of Korea, Japan (type locality), Republic of Korea.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb


The general biology of species in the genus was summarized by Taylor (1967): Ponera are small ants that nest in rotting logs in forested areas or under stones in nonforested situations. In the tropical areas specimens are rarely encountered away from rain forest. In temperate areas, however, species may occur in relatively lightly forested areas. This appears to be the case with Ponera japonica, Ponera pennsylvanica and especially with Ponera coarctata. The Australian Ponera leae is essentially limited to rain forest in the northern parts of its range, but further south it may be found in dry, lightly forested areas.

Foraging is probably cryptobiotic, though some New Guinea species have been taken straying on the ground surface. Little information is available concerning feeding. However, most species are probably insectivorous. I have conducted feeding experiments with some of the New Guinea and Samoan species, including Ponera xenagos, Ponera elegantula, Ponera tenuis, Ponera incerta and Ponera woodwardi. These were unsuccessful with the larger species, except elegantula, which accepted moderately large (8-12 mm) campodeid and japygid Diplura. Tenuis and incerta accepted smaller (4-6 mm) campodeids, isotomid and sminthurid Collembola, and small newly hatched spiders (2 mm long). Negative feeding response was obtained with eggs and larvae of various ants, small crushed insects of various orders, and small myriapods. Stray workers were never observed carrying prey, and distinct middens of insect or other remains were not located near nests.

Colonies usually contain about 30 workers. Larvae and pupae are not segregated in most cases, but occasionally aggregations of pupae were observed. These may have included the total brood of the colonies involved. Larvae are attached to the floor or walls of the nest galleries by the glutinous abdominal tubercles described above, and the ants move them high up on the walls or ceilings of artificial nests, if they are flooded. Details of nuptial behavior of pennsylvanica were given by Wheeler (1900), and Haskins & Enzmann (1938). The flights appear to be of a pattern typical for ants, with the alates meeting in the air and mating there or on the ground. Colony foundation is non-claustral and independent in pennsylvanica (Kannowski 1959); judging from my observations this is typical for the genus.


Leong et al. 2019. Figure 65. P. scabra and P. wui mesosoma detail.


Leong et al. 2019. Figure 45. Worker, CMPon170.


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

  • scabra. Ponera scabra Wheeler, W.M. 1928d: 99 (w.q.) JAPAN. Santschi, 1937h: 364 (gynecoid w.); Santschi, 1941: 273 (q.); Ogata, 1987: 121 (m.); Imai & Kubota, 1972: 194 (k.). Senior synonym of yakushimensis: Yoshimura, Hosoishi, et al. 2009: 195. See also: Wilson, 1957b: 381; Taylor, 1967a: 49.
  • yakushimensis. Ponera yakushimensis Tanaka, 1974b: 32, fig. 1 (w.q.) JAPAN. Junior synonym of scabra: Yoshimura, Hosoishi, et al. 2009: 195.

Type Material

Taxonomic Notes

Leong et al. (2019): First of all, we found that a specimen (ANTC40459 collected from KiuShiu, Shiroyama, Japan) which was mistakenly pinned with the type label of Ponera scabra (see: Antweb.org 2017), deposited in Naturhistorisches Museum, Basel. Regarding the type locality of P. scabra Wheeler, 1928, the species was described based on specimens collected from Mt. Maya, Hyogo Prefecture, Japan. Therefore, the specimen (ANTC40459) does not belong to type series. Secondly, this specimen was reported by Santschi in 1937, who identified it as a gynecoid worker of P. scabra (see: Santschi, 1937: 364). However, this specimen is not P. scabra and can be easily identified as Ponera kohmoku Terayama, 1996 by the combination of the following characters: large compound eye (ca. 30 facets), distinct metanotal groove and subcircular petiolar node in dorsal view; in contrast to P. scabra, which present small compound eye (ca. 5 facets), an indistinct metanotal groove and an arched petiolar node in dorsal view. In summary, we assume that the other specimens of P. kohmoku workers were also misidentified as P. scabra gynecoid by Santschi (1937). Thus, no valid record of gynecoid worker exist in the Ponera genus after the exclusion of Santschi’s record (1937).

Taylor 1967 Ponera fig 27-32

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.



Taylor (1967):

1. The 3 large apical mandibular teeth occupy about 1/3 of the masticatory border, and are followed by a series of 10-12 minute denticles.

2. Median clypeal denticle much less pronounced, at least in smaller specimens (see below).

3. Scapes relatively short; when laid back along head they almost exactly reach the median occipital border.

4. Head narrower (cephalic index 78-83 opposed to 83-85 in chapmani).

5. Mesometanotal suture much less distinctly incised, although indicated by a break insculpturation.

6. Posterior border of node, viewed from above, distinctly concave. Node relatively wide (petiolar node index 84-89 opposed to 79-80 in chapmani).

7. Sculpturation of mesosoma and node much more pronounced; striae of propodeal declivity and posterior face of node lacking (see Wilson's description).

The posterior face of node with fine transverse superficial "scaly" sculpturation, like that on propodeal declivity of Ponera alpha, but more distinctly developed. This sculpturation, which probably represents a vestigial striate-rugosity, is more distinct on the lower 1 /2 of face than above.

Additional description. The following notes are based on 7 syntype workers and 2 dealated syntype queens from the MCZ and AMNH collections. An additional unstudied syntype worker is in the USNM collection (M. R. Smith, in litt). Wheeler presumably returned some of his types to Silvestri and it is not known whether he designated a holotype. Accordingly, no lectotype selection has been made here.

Syntypes (figs. 31, 32). In addition to the features indicated in the diagnosis above the following details should be noted: HL 0.77-0.80 mm; HW 0.61-0.64 mm; SL 0.56-0.59 mm; CI 78-80; SI 90-93; PW 0.46-0.50 mm; PNL 0.26-0.28 mm; PH 0.49-0.53 mm; DPW 0.39-0.43 mm; PNI 84-88.

Eyes about 0.03 mm in diameter, with 3-5 indistinct facets. Median clypeal tooth at best vestigial, represented by a low flat tumosity ; a little more distinct in larger specimens. Antennal club as described for chapmani. General form of propodeum as in chapmani; petiolar node somewhat bulkier (cf. figs. 30, 32).

Additional workers. 2 workers from Hikosan, Kyushu (K. Yasumatsu) have the following dimensions : HL 0.78 mm, 0.84 mm; HW 0.64 mm, 0.70 mm; SL 0.57 mm, 0.61 mm; CI 82, 83; SI 89, 87; PW 0.49 mm, 0.55 mm; PNL 0.25 mm, 0.30 mm; PH 0.51 mm, 0.55 Mm; DPW 0.44 mm, 0.47 mm; PNI 89, 85. These specimens agree well with the syntypes but have relatively high CI and low SI values. The larger specimen is probably slightly gynecoid; its eyes are about 0.05 mm long with 7 rather distinct facets, and the median clypeal denticle is well developed, as in queen. The palpal formula (smaller specimen dissected): Maxillary 2 : Labial 2.

Leong et al. (2019) - (n=3): HL 0.74–78; HW 0.61–0.64; SL 0.53–0.54; A06L 0.04; A07L 0.05; A08L 0.07; A09L 0.08; A10L 0.10; PrW 0.48–0.52; WL 0.97–1.05; PeH 0.48–0.53; PeNL 0.27–0.29; PeW 0.42–0.47; ATL 0.57–0.60; ATW 0.64–0.68; CI 78–85, SI 84–92, PeI 87–92, LPeI 53–56, DPeI 151–166, ATI 88–93.

In full-face view, head rectangular and distinctly longer than broad (CI: 78–85), with slightly convex posterior margin, convex lateral margins and angularly rounded posterolateral corners. Eye small; composed of a total of 5 to 7 indistinct facets. Anterior clypeal margin with strong and acute medial tooth. Masticatory margin of mandible with a series of about 15 indistinct denticles, and three large teeth on the apical part. Antennal scape, when laid backward, almost reaching posterolateral corner; average ratio of the length of antennomeres 7/6:8/6:9/6:10/6 = 1.25: 1.69: 2.93: 3.40 (n=3).

Mesosoma. Mesosomal dorsum in lateral view slightly convex. Pronotum in dorsal view with broadly convex anterior margin, and well convex lateral margins. Metanotal groove indistinct. Lateral mesopleural suture distinctly incised. Propodeal dorsum in dorsal view slightly narrow with straight lateral margins. Propodeal corner in lateral view angular; propodeal dorsum and declivity forming approximatively a 120 degree angle.

Metasoma. Petiolar node in dorsal view thick and arched, with broadly convex anterior margin, and concave posterior margin. Petiolar node in lateral view trapezoidal and thick, with straight anterior and posterior margins; anterodorsal corner convex and higher than posterodrosal corner. Subpetiolar process large with large and circular fenestra, anteroventral corner blunt, almost straight ventral margin with a pair of well developed teeth. Third abdominal tergum distinctly broader than long (ATI: 88–93), with straight anterior margin, and slightly convex lateral margins.

Sculpture. Head densely punctate. Mandible sparsely punctate. Mesosoma densely punctate. Mesopleuron with densely striate and punctate. Metapleuron and propodeum striate. Propodeal declivity shining, but sparsely punctate. Lateral face of petiole densely punctate, posterior face smooth. The third and fourth abdominal segments densely punctate, other segments smooth and shining with few punctures.

Pubescence. Head, antennae, mesosoma, petiole, and gaster with evenly distributed short hairs. Mesopleuron, metapleuron, propodeum and lateral face of petiole with few short hairs. Dorsal and ventral faces of head, anterior margin of clypeus, sides of mandibles, dorsum of petiolar node, gastral sterna and posterior half of gastral terga with many long erect hairs. Subpetiolar process with a few long erect hairs.

Color. Body color black. Mandible, clypeus, antennae, legs, and apex of metasoma brown.


dealate Length 4 mm. Very similar to the worker. Head with even less convex sides and the petiole not so thick and with more concave posterior surface.

Taylor (1967) - Differentiated from queen of Ponera chapmani by larger size and the same mandibular, scape, nodal and sculptural characters which distinguish the worker. Compound eye distinctly smaller, its maximum diameter 0.25 to 0.26 X head width, opposed to 0.29 to 0.30 X HW in chapmani. Wing venation unknown (unfortunately; considering the peculiar venation of chapmani).

Syntypes (N = 2): HL 0.87 mm; HW 0.71 mm, 0.74 mm; SL 0.64 mm, 0.65 mm; CI 82, 85; SI 90, 88; PW 0.60 mm, 0.66 mm; single measurements (1st specimen) for PNL 0.30 mm and PH 0.60 mm; DPW 0.51 mm, 0.55 mm; PNI 85, 83; maximum diameter of compound eye 0.18 mm; 0.19 mm; ocular index 25, 26.

Differing from the workers in the usual characters, and from other Ponera queens by the characters of the diagnosis above. The palpal formula has not been checked.


  • n = 4, 2n = 8, karyotype = 2M+6SM (2M + 2A) (Japan) (Imai & Kubota, 1972; Crozier, 1975; Mariano et al., 2015) (Mariano et al., 2015 list 2n=7).
  • n = 3 (Japan) (Crozier, 1975; Imai et al., 1988a).
  • n = 4 (Japan) (Crozier, 1975; Imai et al., 1988a).
  • 2n = 7 (Japan) (Imai et al., 1988a).
  • 2n = 8 (Japan) (Imai et al., 1988a).

Determination Clarifications

Taylor (1967) - The records of Ponera scabra published in the important ecological studies of Hokkaido ants by Hayashida (1957, 1960) were in fact based on incorrectly determined specimens of Ponera japonica, which were generously made available for this study by Dr Hayashida. For further information see japonica.


References based on Global Ant Biodiversity Informatics

  • Azuma M. 1953. On the myrmecological fauna of Mt. Rokko, Hyogo Prefecture. Warera 2:1-7.
  • CSIRO Collection
  • 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. 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., 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 Park, K.S. 1991. Studies on the distribution of ants (Formicidae) in Korea (7). Ant fauna in Mt. Kyeryongsan. Korean Journal of Applied Entomology 30: 80-85.
  • Collingwood C. A. 1976. Ants (Hymenoptera: Formicidae) from North Korea. Annales Historico-Naturales Musei Nationalis Hungarici 68:
  • Eto S., and K. Ogata. 1983. Ants of Hirado Island, Kyushu. Bulletin of the Nagasaki Prefecture Biological Group 25: 7-11.
  • 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., M. Enomoto, N. Nishimata, and K. Nishimuta. 2014. Ants of the Tokara Islands, northern Ryukyus, Japan. Nature of Kagoshima 40: 111–121.
  • 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.
  • Hosoichi S., M. Yoshimura, Y. Kuboki, and K. Ogata. 2007. Ants from Yakushima Island, Kagoshima Prefecture. Ari 30: 47-54.
  • 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.
  • 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.
  • Imai, H. T.; Kubota, M. 1972. Karyological studies of Japanese ants (Hymenoptera, Formicidae) III. Karyotypes of nine species in Ponerinae, Formicinae and Myrmicinae. Chromosoma (Berlin) 37:194. [1972-05-12] PDF 126055
  • Ito. F., Kondoh. M., Kubota. S., Masuko. K., Morishita. M., Murata. K., Ogata. K., Sato. T., Takamine. H., Yamaoka. H. and Kondoh. M. 1986. A list of ants collected at Akiyoshi-dai (Yamaguchi-ken) by the members of the Myrmecologists Society (Japan) in 1985. ARI Reports of the Myrmecologists Society (Japan) 14: 5-6
  • Janda M., G. D. Alpert, M. L. Borowiec, E. P. Economo, P. Klimes, E. Sarnat, and S. O. Shattuck. 2011. Cheklist of ants described and recorded from New Guinea and associated islands. Available on http://www.newguineants.org/. Accessed on 24th Feb. 2011.
  • 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., S.J. Park, and J.H. Kim. 1996. Ants from Naejangsan national park (Hymenoptera: Formicidae). Korean J. Soil. Zoology &(2): 120-133.
  • Kim B.J.; Kim, J.H.; Kim K 1998. Systematic study of Ponerinae (Hymenoptera: Formicidae) from Korea. Korean Journal of Entomology 28:145-154.
  • Kim C.H., B.M. Choi, and J.R. Bang. 1992. Studies on the distribution of ants (Formicidae) in Korea (8)-Ant fauna in 10 islands, Chollanam-do. Korean J. Appl. Entomol. 31(4): 345-359.
  • 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.
  • 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., C. M. Lee, J. H. Chun, J. H. Sung, and S. K. Kim. 2011. Ants in Hongneung forest. Korea Forest Research Institute, 92 pages.
  • Leong C. M., B. Guénard, S. F. Shiao, & C. C. Lin. 2019. Taxonomic revision of the genus Ponera Latreille, 1804 (Hymenoptera: Formicidae) of Taiwan and Japan, with a key to East Asian species. Zootaxa 4594: 1-86.
  • Lyu D. 2008. Taxonomic study on the Poneromorph subfamilies group (Hymenoptera: Formicidae) in Korea. Korean J. Appl. Entomol. 47(4): 315-331.
  • 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 (second report; ants of mountain zone). Ari 17: 8.
  • Masuko K. 1986. A list of ants collected at Akiyoshi-dai (Yamaguchi-ken) by the members of the Myrmecologists Society (Japan) in 1985. Ari 14:5.
  • Masuko, K. 2010. Nest density and distribution of subterranean ants in an evergreen broadleaf forest in Japan with special reference to Amblyopone silvestrii. Entomological Science 13:193
  • 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.
  • Minato M., T. Kameyama, F. Ito, and T. Itino. 1996. A preliminary report of ant fauna in Gagawa Prefecture. Ari 20: 9-13.
  • 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
  • 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 55(2): 127-221.
  • Santschi F. 1937. Fourmis du Japon et de Formose. Bulletin et Annales de la Société Entomologique de Belgique. 77: 361-388.
  • Sato T., N. Tsurusaki, K. Hamaguchi, and K. Kinomura. 2010. Ant fauna of Tottori prefecture, Honshu, Japan. Bulletin of the Tottori Prefectural Museum 47: 27-44.
  • 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.
  • Tanaka H. O., T. F. Haraguchi, I. Tayasu, and F. Hyodo. 2019. Stable and radio-isotopic signatures reveal how the feeding habits of ants respond to natural secondary succession in a cool-temperate forest. Insectes Sociaux 66(1): 37-46.
  • Tanaka M. 1974. A new species of the ant genus Ponera from Yaku Island (Hymenoptera, Formicidae). Entomol. Rev. Jpn. 27: 32-36.
  • Taylor R. W. 1967. A monographic revision of the ant genus Ponera Latreille (Hymenoptera: Formicidae). Pacific Insects Monograph 13: 1-112.
  • 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., 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 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. 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
  • Touyama Y. 2000. Estimating species richness: an application of the time unit sampling method to a myrmecofaunal survey. Jpn. J. Enviro. Entomol. Zool. 11: 51-60.
  • 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.
  • Xu Z. 1998. A report of fourty-one ant species newly recorded in China from Xishuangbanna District of Yunnan Province (Hymenoptera: Formicidae). Zhongguo Xue Shu Qi Kan Wen Zhai 4: 1119-1121.
  • 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.
  • Yoshimura M., S. Hosoishi, Y. Kuboki, K. Onoyama, and K. Ogata. 2009. New synonym and new Japanese record of the ant genus Ponera (Hymenoptera: Formicidae). Entomological Science 12: 194-201.
  • 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.