Tetraponera attenuata

Tetraponera attenuata is distributed widely in Asia, ranging from northeast India, through continental and insular SE Asia to southern China, Taiwan and Japan. Specimens occur in lowland rainforest, primary rainforest, primary dipterocarp forest, lower montane mixed dipterocarp forest, hilltop dipterocarp forest, second-growth forest and (male only) mangrove. There are a few colony series from dead twigs or dead branches, indicating that this species has generalist nesting habits (Ward, 2001).

Identification
Ward (2001) - This species can be recognised by the broad head, elongate petiole, and dense covering of suberect pubescence and standing pilosity on most of the body. Workers from northern populations of T. attenuata tend to have more widely separated frontal carinae, smaller eyes, a shorter petiole and a longer metabasitarsal sulcus. But the relevant indices (FCI, REL, REL2, PLI and PWI) overlap substantially, and the variation appears to be clinal in nature. Queens of T. attenuata show comparable geographical variation in the same features. In addition, the queens vary in the shape of the clypeus (anterolateral teeth variably developed, occasionally absent) and the size of the malar pit (varying from well developed to absent), but variation in these latter traits does not show a clear-cut geographical pattern.

Distribution
Ward (2001) - Tetraponera attenuata ranges from northeast India to southern China, south to Sumatra, Java, and Borneo. The only Philippines records come from Palawan. A single dealate queen has also been taken on Okinawa in Japan.

Distribution based on Regional Taxon Lists
Indo-Australian Region: Borneo, Indonesia, Malaysia, Philippines, Singapore. Oriental Region: Cambodia, India, Laos, Myanmar, Thailand, Vietnam. Palaearctic Region: China, Japan.

Nomenclature

 *  attenuata. Tetraponera attenuata Smith, F. 1877b: 71 (w.) BORNEO. Wheeler, W.M. 1919e: 144 (q.m.). Combination in Sima: André, 1892b: 53; in Tetraponera: Wheeler, W.M. 1919e: 65. Senior synonym of tenuissima: Forel, 1912n: 55; of birmana, thagatensis: Ward, 2001: 622.
 * tenuissima. Sima attenuata var. tenuissima Emery, 1900d: 675, fig. 5 (w.q.) INDONESIA (Sumatra). Junior synonym of attenuata: Forel, 1912n: 55.
 * birmana. Sima birmana Forel, 1902c: 245 (q.) MYANMAR. Bingham, 1903: 112 (w.). Combination in S. (Tetraponera): Emery, 1921f: 25; in Tetraponera: Chapman & Capco, 1951: 79. Junior synonym of attenuata: Ward, 2001: 622.
 * thagatensis. Sima attenuata var. thagatensis Forel, 1902c: 249 (w.) INDIA. Combination in S. (Tetraponera): Emery, 1921f: 27; in Tetraponera: Wheeler, W.M. 1924b: 5; Chapman & Capco, 1951: 82. Junior synonym of attenuata: Bingham, 1903: 112; revived from synonymy as subspecies of nigra: Forel, 1909d: 226; raised to species: Forel, 1912n: 55; returned to junior synonymy of attenuata: Ward, 2001: 622.

Worker
Ward (2001) - HW 0.97-1.48, HL 1.07-1.67, LHT 0.97-1.47, CI 0.84-0.94, FCI 0.11-0.15, REL 0.29-0.34, REL2 0.32-0.38, SI 0.61-0.76, SI3 1.84-2.11, FI 0.33-0.40, PLI 0.38-0.47, PWI 0.29-0.44, PDI 0.97-1.06, LHT/HW 0.93-1.08, CSC 10-26, MSC 6-44.

Relatively large and slender species, with broad head (CI >0.82); clypeus short, its anteromedial margin varying from broadly convex to straight; anteromedial margin of clypeus joining anterolateral margin in a smooth curve or (less commonly) at a slight angle; frontal carinae moderately well separated, the distance between them exceeding basal scape width; eyes relatively small (see REL and REL2 values), scape length about twice eye length; lateral margins of head rounding broadly into posterior margin; profemur slender; pronotum usually lacking prominent lateral margins; mesopropodeal impression wellmarked, long and open, flanked by low metanotal tubercles that are usually not prominent in profile; propodeum about as high as wide, dorsal face rounding gradually and continously into the declivitous face; legs long relative to head size, LHT/HL 0.84-0.96; petiole somewhat variable in shape but always rather long and slender (PLI <0.48), often markedly so, and with a well differentiated anterior peduncle and node; petiole more than twice as long as broad; postpetiole 1.1-1.4x longer than broad; metabasitarsal sulcus lying in a darkened patch of cuticle subtended by a raised ridge, occupying about 0.15-0.30 the length of the basitarsus. Integument with numerous fine punctures, the interspaces (sub)lucid and smooth or weakly reticulate; punctures on dorsum of head and mesosoma mostly 0.005-0.015 mm in diameter (pronotal punctures sometimes larger) and separated by one to several diameters; punctures on side of mesosoma finer, less conspicuous; lower malar area with coarse punctures intermixed with irregular longitudinal rugulae. Standing pilosity common (see CSC and MSC values), the hairs up to 0.30-0.35 mm long, and tending to grade into shorter suberect and decumbent pubescence which forms a dense cover on most of the body, including the mesosoma dorsum and petiole. Black to dark brownish-black, scape and first several funicular segments often lighter (medium to yellowish-brown).

Type Material
Ward (2001) - Syntype (unique?), worker, Sarawak (labeled 'Sar.'), Malaysia.

References based on Global Ant Biodiversity Informatics

 * André E. 1892. Voyage de M. Chaper à Bornéo. Catalogue des fourmis et description des espèces nouvelles. Mém. Soc. Zool. Fr. 5: 46-55.
 * CSIRO Collection
 * 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. 1970. Formicidae (Hymenopter: Aculeata) of Nepal. Himalaya Khumbu Himal, 3: 371-388.
 * Crawley W.C. 1924. Ants from Sumatra, with biological notes by Edward Jacobson. Annals and Magazine of Natural History (9)13: 380-409
 * Davidson D. W., S. C. Cook, R. R. Snelling and T. H. Chua. 2003. Explaining the Abundance of Ants in Lowland Tropical Rainforest Canopies. Science 300: 969-972.
 * Dlussky G. M.; Radchenko A. G. 1990. The ants (Hymenoptera, Formicidae) of Vietnam. Subfamily Pseudomyrmicinae. Subfamily Myrmicinae (tribes Calyptomyrmecini, Meranoplini, Cataulacini). Pp. 119-125 in: Akimov, I. A.; Emelianov, I. G.; Zerova, M. D. et al. (eds.) 1990. News of faunistics and systematics. [In Russian.]. Kiev: Naukova Dumka, 184 pp.
 * Eguchi K., and S. Yamane. 2003. Species diversity of ants (Hymenoptera, Formicidae) in a lowland rainforest, northwestern Borneo. New Entomol. 52(1,2): 49-59.
 * Eguchi K.; Bui T. V.; Yamane S. 2011. Generic synopsis of the Formicidae of Vietnam (Insecta: Hymenoptera), part I  Myrmicinae and Pseudomyrmecinae. Zootaxa 2878: 1-61.
 * Emery C. Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei. Annali del Museo Civico di Storia Naturale 40: 661-722.
 * Emery, C. "Formiche raccolte da Elio Modigliani in Sumatra, Engano e Mentawei." Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova) (2) 20, no. 40 (1900): 661-722.
 * Floren A., W. Wetzel, and M. Staab. 2013. The contribution of canopy species to overall ant diversity (Hymenoptera: Formicidae) in temperate and tropical ecosystems.  Myrmecological News 19: 65-74.
 * 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. 1901. Formiciden des Naturhistorischen Museums zu Hamburg. Neue Calyptomyrmex-, Dacryon-, Podomyrma- und Echinopla-Arten. Mitt. Naturhist. Mus. Hambg. 18: 43-82.
 * Forel A. 1902. Myrmicinae nouveaux de l'Inde et de Ceylan. Rev. Suisse Zool. 10: 165-249.
 * Forel A. 1903. Les Formicides de l'Empire des Indes et de Ceylan. Part X. J. Bombay Nat. Hist. Soc. 14: 679-715.
 * Forel A. 1909. Ameisen aus Java und Krakatau beobachtet und gesammelt von Herrn Edward Jacobson. Notes Leyden Mus. 31: 221-232.
 * Forel A. 1912. Einige neue und interessante Ameisenformen aus Sumatra etc. Zool. Jahrb. Suppl. 15: 51-78.
 * Forel A. 1913. H. Sauter's Formosa-Ausbeute: Formicidae II. Arch. Naturgesch. (A)79(6): 183-202
 * Forel A. 1913. Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien ausgeführt im Auftrage der Kgl. Preuss. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen. II. Ameisen aus Sumatra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren 1911-1912. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 36:1-148.
 * Gumawardene, N.R., J.D. Majer and J.P. Edirisinghe. 2008. Diversity and richness of ant species in a lowland wet forest reserve in Sri Lanka. Asian Myrmecology 2:71-83
 * Gunawardene N. R., J. D. Majer, and J. P. Edirisinghe. 2008. Diversity and richness of ant species in a lowland wet forest reserve in Sri Lanka. Asian Myrmecology 2: 71-83.
 * Gunawardene N. R., J. D. Majer, and J. P. Edirisinghe. 2012. Correlates of ant 5Hymenoptera: Formicidae) and tree species diversity in Sri Lanka. Myrmecological News 17: 81-90.
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Hashimoto Y., M. Mohamed, and H. Sakata. 1999. The ants (Hymenoptera: Formicidae) of the Tabin Wildlife Reserve, Sabah. Tabin Scientific Expedition 69-74.
 * Hashimoto Y., and M. Mohamed. 2004. A preliminary survey of ant fauna at Crocker Range Park. In: Maryati Mohamed, Zulhazman Hamzah, T. Tachi & J. Nais (eds.). Crocker Range scientific expedition 2002. Kota Kinabalu: Universiti Malaysia Sabah, pp. 51-71.
 * Itino T., and S. Yamane. The vertical distribution of ants on canopy trees in a Bornean lowland rainf forest. Tropics 4(2/3): 277-281.
 * Jaitrong W., B. Guenard, E. P. Economo, N. Buddhakala, and S. Yamane. 2016. A checklist of known ant species of Laos (Hymenoptera: Formicidae). Asian Myrmecology 8: 1-32. DOI: 10.20362/am.008019
 * Jaitrong W., and T. Ting-Nga. 2005. Ant fauna of Peninsular Botanical Garden (Khao Chong), Trang Province, Southern Thailand (Hymenoptera: Formicidae). The Thailand Natural History Museum Journal 1(2): 137-147.
 * Jaitrong W.; Nabhitabhata, J. 2005. A list of known ant species of Thailand. The Thailand Natural History Museum Journal 1(1): 9-54.
 * Karavaiev V. 1933. Ameisen aus dem Indo-Australischen Gebiet, VII. (Schluss). Konowia 12: 260-271.
 * Karavaiev V. 1935. Neue Ameisen aus dem Indo-Australischen Gebiet, nebst Revision einiger Formen. Treubia 15: 57-118.
 * Katayama M., K. Kishimoto-Yamada, H. O. Tanaka, T. Endo, Y. Hashimoto, Sk. Yamane, and T. Itioka. 2015. Negative correlation between ant and spider abundances in the canopy of a Bornean tropical rain forest. Biotropica (in press).
 * Li Q., Y. Chen, S. Wang, Y. Zheng, Y. Zhu, and S. Wang. 2009. Diversity of ants in subtropical evergreen broadleaved forest in Pu'er City, Yunnan. Biodiversity Science 17(3): 233-239.
 * Li Z.h. 2006. List of Chinese Insects. Volume 4. Sun Yat-sen University Press
 * Lin S. Y., B. Di Giusto, A. Bain, and L. S. Chou. 2016. Variation of ant community structure on Ficus benguetensis. Taiwania 61(1): 49-57.
 * 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
 * Malsch A. K. F., K. Rosciszewski, and U. Maschwitz. 2003. The ant species richness and diversity of a primary lowland rain forest, the Pasoh Forest reserve, West Malaysia. in T. Okuda, N. Manokaran, Y. Matsumoto, K. Niiyama, S. C. Thomas, and P. S. Ashton, eds. Pasoh: Ecology and Natural History of a Southeast Asin Lowland Tropical Rain Forest, pp 347-374.
 * Mohamed M. 1995. A preliminary list of ants (Hymenoptera: Formicidae) of the Tawau Hills Park, Sabah. In: Ghazally Ismail  et al.  (eds.),  Tawau  Hills Park,  Sabah,  Pelanduk Pub.  Pp.  205-213.
 * Pfeiffer M.; Mezger, D.; Hosoishi, S.; Bakhtiar, E. Y.; Kohout, R. J. 2011. The Formicidae of Borneo (Insecta: Hymenoptera): a preliminary species list. Asian Myrmecology 4:9-58
 * Radchenko A. G. 1993. Ants from Vietnam in the collection of the Institute of Zoology, PAS, Warsaw. I. Pseudomyrmicinae, Dorylinae, Ponerinae. Annales Zoologici (Warsaw) 44: 75-82.
 * 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. Fourmis de Sumatra, récoltées par Mr. J. B. Corporaal. Tijdschrift voor Entomologie 71: 119-140.
 * 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.
 * Sukimin S., M. Mohamed, and H. Aris. 2010. Ant diversity of Maliau Basin Conservation Area, Sabah, Malaysia. Journal of Tropical Biology and Conservation 6:89-101.
 * Tanaka H. O., S. Yamane, and T. Itioka. 2012. Effects of a fern-dwelling ant species, Crematogaster difformis, on the ant assemblages of emergent trees in a Bornean tropical rainforest. Ann. Entomol. Soc. Am. 105(4): 592-598.
 * 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 M., S. Kubota, and K. Eguchi. 2014. Encyclopedia of Japanese ants. Asakura Shoten: Tokyo, 278 pp.
 * Terayama, M. 2009. A synopsis of the family Formicidae of Taiwan (Insecta; Hymenoptera). The Research Bulletin of Kanto Gakuen University 17: 81-266.
 * 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.
 * Ward P. S. 2001. Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions. Invertebrate Taxonomy 15: 589-665.
 * Ward, P. S. 2001. Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions. Invertebrate Taxonomy 15:589-665.
 * Wheeler W. M. 1919. The ants of Borneo. Bulletin of the Museum of Comparative Zoology 63:43-147.
 * 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, William Morton. 1924. Ants of Krakatau and Other Islands in the Sunda Strait. Treubia. 5(1-3):1-20.
 * Widodo E.S., M. Mohamed, and Y. Hashimoto. 2001. Canopy ant diversity assessment in the fragmented rainforest of Sabah, East Malaysia. Nature and Human activities 6: 13-23.
 * Xu Z. and Z.-Q. Chai. 2004. Systematic study on the ant genus Tetraponera F. Smith (Hymenoptera, Formicidae) of China. Acta Zootaxonomica Sinica 29(1): 63-76.
 * Yamane S. 2013. A Review of the ant fauna of the Krakatau Islands, Indonesia. Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist. Ser: A, 11: 1-66
 * Yamane S.; Nona, A. R. 1994. Ants from Lambir Hills National Park, Sarawak. Pp. 222-226 in: Inoue, T.; Hamid, A. A. (eds.) 1994. Plant reproductive systems and animal seasonal dynamics. Long-term study of dipterocarp forests in Sarawak. Kyoto: Center for Ecological Research, Kyoto University, vii + 255 pp.
 * Yamane Sk., and T. Itino. 1995. The vertical distribution of ants on canopy trees in a Bornean lowland rain forest. Tropics 4(2-3): 277-281.
 * 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.
 * Zryanin V. A. 2011. An eco-faunistic review of ants (Hymenoptera: Formicidae). In: Structure and functions of soil communities of a monsoon tropical forest (Cat Tien National Park, southern Vietnam) / A.V. Tiunov (Editor). – M.: KMK Scientific Press. 2011. 277 р.101-124.