SolvingSolving thethe Hundred-YearHundred-Year ControversyControversy ofof CoptotermesCoptotermes TaxonomyTaxonomy inin TaiwanTaiwan

Hou-Feng Li, Nan-Yao Su, and Wen-Jer Wu

he genus Wasmann 1896 is a large group of characters, individuals of the same species collected from colonies T(Constantino 2009, Vargo and Husseneder 2009), 28 of which aresubterranean considered importanttermites, includingstructural pests≈70 described (Edwards andspecies Mill at different ages and habitats likely show significant differences. example,Without comparing at least 12 synonymsadequate numbers have been of recordedsamples, afor plethora C. formosanus of new subterranean has been estimated at $22 billion annually (Su species have been proposed over the same geographic areas. For 2002),1986, Su and and Coptotermes Scheffrahn has 2000). been reportedThe global as theeconomic most destructive impact of ofall Difficulty in matching soldiers and alates of the same species. alone (Snyder 1949, Li 2000). Soldiers associated with damaged wood can be collected year-round, Cancellothe group 2001, in the Constantino southern United 2002), States Australia (Su 2003, (Calaby Scheffrahn and Gay 1956,and Su Soldiers and alates of the same colony are rarely collected together. 2005), Hawaii (Bess 1970, Woodrow et al. 2001), Brazil (Ferraz and alates and soldiers of the same species might be described as two butspecies, alates as areoccurred usually with collected C. gestroi during and C. their havilandi dispersal Holmgren flight. (KirtonHence, Lo et al. 2006), southern China (Lin 1987, Wang et al. 2002), Taiwan Coptotermes 2004),(Oshima Thailand 1919, Li (Sornnuwat 2009), southern 1996), Japan Malaysia (Mori (Lee 1987), 2002), Philippines and India descriptions of both soldier and alate castes (Snyder 1949, Roonwal and Mariana Islands (Su and CoptotermesScheffrahn 1998, Yudin 2002, Acda and Brown 2003). To date, only half of spp. are based on Since most Coptotermes pest species originated in Asia, entomolo- (Roonwal and Chhotani 1989). spp. have frequently been and Chhotani 1962, Li 2000). intercepted at ports (Gay 1967, Ping 1991), and some have become Coptotermes species in Taiwan and southern Japan established in non-native areas (Gay 1969, Scheffrahn et al. 1990, gists in these areas encountered the taxonomic challenge early. The The2004; tendency Jenkins etof al.Coptotermes 2007). They are continually dispersed by human- Japaneseidentification entomologists of and European experts were involved mayaided contribute land transportation to their wide (Jenkins distribution, et al. especially2002, Austin in C. et formosanus al. 2008). was a controversial issue between 1909 and 1912. Many influential Shiraki and C. gestroi spp. to colonize cargo, boats, and ships this historical record, clarify the controversial issues based on our Despite its economic importance, of the genus is still in this debate. One hundred years after the controversy, we review (Wasmann) (ScheffrahnCoptotermes and Su 2005). taxonomy include: current study, and offer solutions to avoid similar problems in the problematic.Lack of distinguishing The inherent features. difficulties Soldiers in and alates are the two Backgroundfuture. of Termite Study in Taiwan in the Early 1900s Cop- totermes China and the Meiji government of Japan, Taiwan became a Japanese main castes used for termite identification. Soldiers among territoryAfter thein 1895 First underSino-Japanese the Treaty War of betweenShimonoseki the Qing until Dynasty the end of spp. are superficially similar (Calaby and Gay 1956). Few observingcharacters diagnostic of the soldier characters, are available such as the for number species of identification setae around termites are primarily subtropical and tropical pests, Japanese con- (Kirton 2005), and a microscopic examination is usually required for WWII in 1945. Since most of Japan has a temperate climate and - structures built in Taiwan during the early Japanese colonial period the fontanelle. Alates offer more differentiable characters for species struction practices were not adapted for termite prevention. Thus, identification, including body length, body color, antennal spot pat tern,High and degreedispersal of flightmorphological season. However, variation alates among are only intraspecific present in suffered from severe termite damage (Oshima 1919). Termite control populations.mature colonies, and for a short period of time each year. became a priority of the Governor of Taiwan in the early 1900s. In 1907, both Tokuichi Shiraki (1882-1970) (Fig. 1A) and Masamitsu 222 By using quantitative comparison of morphological Oshima (1884-1965) (Fig. 1B) startedAmerican to Entomologist work in Taiwan • Winter at 2010 the sanus Fig. 1. (A) Tokuichi Shiraki (1882-1970) (Photo pro- . This included illustrations of the dealate, soldier, worker, fore vided by the Entomology and hind Oshima wings (1909) (Fig. vividly2), and and photographs comprehensively of nests. described Twelve collection damage Dept. of the National Taiwan locations in Taiwan and the Penghu islands were also described (Fig. University); (B) Masamitsu caused by C. formosanus, its tunneling behavior, and soldier behaviors Oshima (1884-1965) 3A). (Photo provided by the Entomological Society such as head banging and secreting from the fontanelle. of Japan). of several termite species named by Shiraki (1909), including C. formosanusIn the following year, Oshima (1910a) questioned the validity - animosity between Shiraki and himself, and they worked together cessor of Taiwan Agricultural Research Institute) and the Bureau of . Oshima (1910a) emphasized that there was no personal Agricultural Experiment Station, Government of Formosa (prede and shared references, books, and termite samples with each other. Civil Engineering. Later, they became the leading figures in termite He had no wish to offend Shiraki, but he believed that the correctC. research on the island. Their major control target was known at Coptotermes formosanusidentification of termite species was a key C.requirement gestroi described for their by the time colloquially as “house termites,” which we now know to be control. Oshima (1910b, 1911) mentioned that the soldiers of species. Controversy in the Identification ofCoptotermes spp. in species, probably superficially due to resembledabsence of those alate ofdescription of C. gestroi Taiwan Haviland in 1898. Oshima did not compare the alates of these two Shiraki and Oshima cooperated on the studies of termite control close to Southeast Asia, where C. gestroi (Wasmann 1896, Haviland 1898). Since Taiwan was geographically C. formosanus, in Japanese, based (1910b, 1911) proposed that C. formosanus occurred, was aand junior no significant synonym and taxonomy during the first two years of their careers in Taiwan. morphological difference of the soldier caste was found, Oshima includedShiraki (1909) descriptions briefly of described the morphology of several castes, including on Oshima’s collection from many locations in Taiwan. The article specimenthe male and of C. femaleformosanus alate, nymph, soldier, worker, and queen, for whichmentioned no figure twice, was but offered. incorrectly In spelledaddition, as the Captotermes type locality formosanus and type [sic] and Coptotermes forrmosanus were not [ included.sic The species name was detailed examination of the morphological] (Shiraki characteristics 1909). In of the C. formo same- year, based on Shiraki’s description, Oshima (1909) offered a more

Fig. 3. Coptotermes spp. collection sites in early 1900s (A) and in the cur- rent study (B). White area, altitude >500 m; light gray area, subtropical lowland; dark gray area, tropical lowland.

of C. gestroi C. ge- stroi/ formosanus . Oshima also provided 12 collection locations of In addition to in Shiraki Taiwan andand theOshima, Penghu several islands other (Oshima Japanese 1911) (Fig. 3A).

researchers such as Munemoto Yano (1883-1970) of Forestry ScienceExperimental College, Station, Tokyo Tokyo,Imperial Shozaburo University Watasé also became (1862-1929) involved and in his student, SanjiCoptotermes Hozawa (1885-1947) of the Zoological Institute,

identification of spp. of Taiwan and southern Japan. Yano Wasmann(1911) mentioned believed that that Watasé these sent samples Japanese were specimens C. gestroi to, whicha German he termite taxonomist, Eric Wasmann (1859-1931), for identification. C. gestroi Fig. 2. Coptotermes formosanus drawn by Oshima in 1909 provided by named in 1896. However, Yano (1911) still had some doubtCoptotermes about the National Taiwan University Library. Dealate (A); soldier (B); worker Wasmann’s opinion. Yano mentioned that the descriptionCoptotermes of (C); fore (D) and hind (E) wings. Vertical scale bar for A-C; horizontal by Wasmann was too vague to differentiate it from other scale bar for D-E. samples with the description of C. gestroi spp. Hence, Yano further compared Japanese/Taiwanese American Entomologist • Volume 56, Number 4 of Haviland (1898).223 He thought the two species were similar, but that their soldiers could - C. formosanus (Shiraki 1909), and that only offered five measurements of alates, which was less informativeThe be distinguished by the ratio of headC. gestroi width toas headdescribed length. by The Haviland Japa Thirdthan the Official first Reportdescription on Termites of was published by Oshima in 1912, nese/Taiwanese soldier samples had elongated heads (1.5 mm in Holmgren failed to provide illustrations or photographs. After length and 1.2 mm in width) while Coptotermes (1915) accepted Oshima’s opinion that C. formosanus is the only (1898) had more circular heads (1.4 mm in length and 1.3 mm in Coptotermesmost termite species researchers in southern such asJapan Holmgren and Taiwan, (1913) and and C. formosae Hozawa 1954),width). through Additionally, Chiyomatsu Yano sent Ishikawa the Japanese/Taiwanese (1861-1935) of Tokyo Imperial is a junior synonymy of C. formosanus Universitysamples to and a Swedish another termite German taxonomist, termite researcher, Nils Holmgren Karl Escherich (1877- Coptotermes gestroi in Taiwan . a new species, Coptotermes formosae Holmgren (Holmgren 1911), After 1912, C. formosanus was believed to be the only Coptotermes and(1871-1951). compared Holmgren it with other examined Coptotermes Yano’s specimens and described Coptotermes gestroi in Taiwan had not been men- C. concluded that the Coptotermes spp. of Sri Lanka. Based gestroispecies in Taiwan. wason Holmgren’s not C. gestroi opinion and his own measurements, Yano (1911) authortioned untilof the 2003. current Based study on and soldier local morphology, pest control theoperators new record collected of of C. formosanus sp. collected in Japan and Taiwan Coptotermes in Taiwan was proposed again (Tsai and Chen 2003). The first . Yano (1911) also criticized Shiraki’s description thought it should be(Shiraki appropriate 1909), to in adopt which Holmgren’s no figures nomenclature, were offered, spp. from every county inC. Taiwan, gestroi inboth southern in urban Taiwan areas C.the formosae format was (Holmgren informal, 1911), and the instead description of C. formosanus was in Japanese. named Yano by and natural environments, from 2005–2009. WithCoptotermes the mitochondrial samples gene sequence data, the presence of was confirmed (Li et al. 2009).Coptotermes In total, 220 Shiraki (1909), because Holmgren was a recognized authority on including museum specimensCoptotermes were identified formosanus and used was for distributed mapping termite taxonomy. theirthroughout distribution the island (Fig. of 3B). Taiwan, while C.spp. gestroi were was mostly only collected Oshima (1912) was torn between the conflicting opinions of the in lowland areas (< 500 m). two internationally recognized authorities on termite taxonomy, In a review of articles published in the early 1900s, some evidence samplesWasmann and and C. Holmgren.gestroi in the He ratio agreed of headwith widtha part andof Yano’s head lengthopinions of showsin the southern that C. gestroi tropical zone. - (1911) that there was a difference between Japanese/Taiwanese hima (1909, 1911) collected Coptotermes samples from 12 locations species (herein C. formosanus may have already been present in Taiwan. Os the solider. However, he thought it was insufficient to name a new C. gestroi and C. formosanus the validity of C. formosanus ) because the difference might be due to in Taiwan (Fig. 3A), seven of which were in the tropicalC. zonegestroi where the Tointraspecific solve this controversialvariation. This issue, was the Oshima major collected reason that more he doubtedsamples 4A) instead of the light brownish-yellow currently coexist. dealate The of C. drawing formosanus of a dark and measured head length and (Oshima width 1910b,with a more 1911). accurate method brown dealate (Fig. 2A) by Oshima C.(1909) gestroi resembled C. formo (Fig.- sanus (Fig.C. in southern Japan and 11 soldiers from three locations in Taipei, formosanus4C). The color of soldier heads of (Fig. 4B) and based on a clear definition. He had seven soldiers from two locations color of(Fig. the 4D) C. gestroi is light yellow, which is similar to the color of the alate abdomen (Fig. 4C), but in contrast to the dark brown theTaiwan distance (Oshima between 1912, the Fig. medial 3A). Hebase removed of the labrum the soldier to the heads hindmost and 2B) was likely based on alate C. gestroi abdomen (Fig. 4A). Based on the color placed them on a glass plate in a horizontal position, then measured difference between soldier and alate, Oshima’s drawing (Fig.C. gestroi2A and than C. formosanus . In addition, the alate body length Taiwanesemargin of the samples head tocapsule. be much The longer data than was thatrounded of C. gestroi to two decimal was recorded as ≈6 mm (Oshima 1909), which is closer to places. Oshima found the head length (1.66–1.72 mm) of Japanese/ (usually >7 mm). The drawing of the soldier head between Japanese and Taiwanese samples (Taipei only), and the(1.4 varia mm)- capsule (Fig. 5A, Oshima 1909) was somewhat rounded and was also as described by Haviland (1898). No significant difference was found note that the head length of soldiers reported by Oshima in 1910b and tion of these samples in head length was only 0.03 mm. It is worthy of

1911 was 1.5 mm, but it increased to 1.66–1.72 mm when he made another measurement in 1912. OshimaCoptotermes attributed samples the collected difference in toJapan different and Taiwan measuring (Taipei methods.materials) Based were on not the C. newgestroi measurements, OshimaOshima confidently (1912) thought believed that that the alate of C. formosanus described by Shiraki (1909) was similar to C. formosae described by. Holmgren C. formosanus C. formosae should

(1911). Since was described first, be a junior synonymy basedC. on formosanus the rules of based zoological on the nomenclature publication (Oshimaformat and 1912). language Oshima used (1912) for also description severely criticizedinstead of Yano’s the nomenattempt- (Yano 1911) to invalidate Fig. 4. Dealate (A) and soldier (B) of C. gestroi, and dealate (C) and original description of C. formosae (Holmgren 1911) in The Third clature rules and the content of the description. Oshima listed the soldier (D) of C. formosanus collected in southern and northern Taiwan, Official Report on Termites (Oshima 1912) to show that Holmgren respectively, in the current study.

224 American Entomologist • Winter 2010 Fig. 5. Soldier head Coptotermes formosanus capsules of Coptotermes spp. A solider collected in A set of specimens (Fig. 6A) labeled “ Taiwan by Oshima in 1909 resembled C. gestroi (A) Shiraki”handwriting was found in an inunpublished TARI, where manuscript Shiraki worked entitled between 1907 Fauna and (Photo provided by the Na- of1942. Taiwan The autograph on the label (Fig. 6B) is very similar to Shiraki’s tional Taiwan University Li- brary); C. formosanus col- lected in Taipei or southern (Fig. 6C) offered by Wen-Jer Wu of the National Taiwan Japan by Oshima in 1912 asUniversity. C. formosanus Termite (C. gestroisoldiers preserved in vials (Fig. 6D) with two (B) (Photo provided by setae on each side of the fontanelle (Fig. 6D, inset) were identified the Entomology Dept. of the National Taiwan has one seta on each side; Scheffrahn et University); SEM pictures al. 1990), but no collection information of these soldiers was found.th, 41st of C. gestroi (C) and C. Three alate specimens were labeled, but damaged to some degree. formosanus (D) collected in southern and northern The best-preserved sample (Fig. 6E) was collected on June 10 Taiwan, respectively, in the year of Meiji era (1908) from a location called “Old Farm,” which is current study. Nitobeunknown (1883-1915), to us, and the Shiraki’s collector assistant, was not on recorded. April 23 Therd, 41 otherst year two of severely damaged samples (Fig. 6F and 6G) were collected by Inao similar to C. gestroi - Meiji era (1908) from Pingtung county, Gangkou research station collected Coptotermes (Fig. samples 5C), and only its headin northern length Taiwanwas recorded (Taipei as city) 1.5 of TFRI (Fig. 3A). A comparison ofC. formosanusthese three alates with the origiC. mm. Three years after his publicationC. (Oshimaformosanus 1909) in 1912, Oshima formosanusnal description specimens (Shiraki preserved 1909) and in our TARI recent once collectionsbelonged to (Li Shiraki et al. 2009)and might has confirmedhave been themused forto be describing C. formosanus. Even though in 1909, these the elongated(Fig. 3A) and and in similar Japan, towhere C. formosanus only is currently found. The image of a soldier head capsule (Fig. 5B, Oshima 1912) was C. collected in 1909 and 1911 might include (Fig. both 5D). C. The formosanus head length and ofC. formosanuscollector of these specimens does not fit the original description gestroisoldiers; hence, reported the averageby Oshima of their in 1912 head was length 1.66–1.72 was shorter mm. than Soldiers that (Shirakino part of 1909). Oshima’s Shiraki collection (1909) was mentioned found in thethat six the insect new collectionsspecies, of C. formosanus , was described based on Oshima’s collection. However, Coptotermes collected only from Taipei in 1912. Cop- totermesOshima samples (1912) inemphasized Taiwan were twice the there same was species only one (Oshima 1909, sp. in Taiwan. However, Based on no this effort false assumption,was made to Coptotermesprove that all samples the collected at many places in Taiwan were lumped together, and the mean1910b, of 1911). morphological measurements of the two species may have been presented as a single species, C. formosanus Coptotermes samples as C. formosae and C. gestroi . When Holmgren and Wasmann identified Japanese/Taiwanese two Coptotermes , respectively (Yano 1911), Oshima and Yano were confused, but they did not suspect that there might be spp. in Taiwan. Shiraki (1909), Oshima (1909, 1910b, 1911, 1912), and Yano (1911) drew their conclusions based on termite samples collected in different places in Taiwan and Japan, Typewhich Specimen may be the of source C. formosanus of the controversy in early 1900s. of C. formosanus and C. gestroi in the early 1900s, museum specimens preservedDuring ourin six investigation major insect into collections the controversy in Taiwan on the were identification examined to search for those used by Shiraki and Oshima:

Wufeng, Taichung, Taiwan, ROC (TARI) 1. Taiwan Agricultural Research Institute, Insect and Mite Collection,

2. Taiwan Forestry Research Institute, Insect Collection, Taipei, (NMNS)Taiwan, ROC (TFRI) 3. National Museum of Natural Science, Taichung, Taiwan, ROC Fig. 6. Shiraki’s C. formosanus collection preserved in the Taiwan Agri- cultural Research Institute. Entire collection with an identification label; Collection, Taipei, Taiwan, ROC (NTU) workers, soldiers in vials, and three pinned alates (A); the autography on 4. National Taiwan University, Department of Entomology, Insect the label (B) is similar to Shiraki’s handwriting in an unpublished manu- script entitled Insect Fauna of Taiwan (C); soldiers (D) preserved in vials Insect Collection, Taichung, Taiwan, ROC (NCHU) with two setae (inset) on one side of the fontanelle; the most well-pre- 5. National Chung-Hsing University, Department of Entomology,- served alate (E) was collected on June 10th, 41st year of Meiji era (1908) at “Old Farm”; two severely damaged alates (F and G) collected by Nitobe rd st 6. National Pingtung University of Science and Technology, Depart on April 23 , 41 year of Meiji era (1908) at Gangkou. Photos of labels and not to scale. ment of Plant Medicine, Insect Collection, Neipu, Pingtung, Taiwan, AmericanROC (NPUST) Entomologist • Volume 56, Number 4 225 Constantino, R. 2009. On-line termite database, Genus Coptotermes C. formosanus of Taiwan. In order to prevent any further confusion on identifying Wasmann, 1896. [http://vsites.unb.br/ib/zoo/docente/constant/ , we designated neotypes herein. Crosland, M. 1995. catal/catnew.html]. NEOTYPE: male alate collected by H.-F. Li at TAIWAN, Taoyuan Edwards, R., and A. E.Taxonomic Mill. 1986. splitters Termites in inChina. buildings, Isoptera their Newsletter biology and 5: Co., Taoyuan City: 24.99°N, 121.30°E; 20-VI-2006 (TW49) will be- 1. depositeddiers collected in NMNS. from the The same neotype colony perfectly as the neotype fit the re-description will be deposited by Eggleton, P. 1999. Termite species description rates and the state of termite Hozawa (1915). NEOPARATYPES: female and male alates, and sol control. Rentokil Ltd., East Grinstead, UK. Ferraz, M. V., and E. M. Cancello. 2001. Swarming behavior of the economi- taxonomy.cally most Insect.important Soc. termite, 46: 1-5. Coptotermes havilandi (Isoptera: Rhino- includingin NMNS, COII,NTU, 12STFRI, rRNA, and Universityand 16S rRNA of Florida of workers Termite collected Collection, from Fort Lauderdale Research and Education Center. Mitochondrial genes Gay, F. J. 1967. termitidae), in southeastern Brazil. Sociobiology 38: 683-694. and submitted to the GenBank database with accession numbers: the same colony with the neotype have been partially sequenced Gay, F. J. 1969. A world review of introduced species of termites.In Aust. Res. Organ. Commonw. Sci. Ind. Bull. 286: 1-88. This review of the historical controversy is intended to present Haviland, G. D.Species 1898. Observationsintroduced by on man, termites: pp. 459-494. with descriptions K. Krishna of and new F. EU805758,the challenge EU805712, for Coptotermes and EU805735 taxonomy, (Li and et to al. prompt 2009). international M. Weesner [eds.], Biology of termites I. Academic Press, New York. cooperation on a revision of Coptotermes Coptotermes is Holmgren, N. 1911. In species. J. Linn. Soc. Lond. 26: 358-422. in serious need of revision, especially the Chinese species (Crosland Ceylon-Termiten, pp. 185-212. K. Escherich [ed.], . Thest genus Holmgren, N. 1913. century, the inherent Termitenleben auf Ceylon. Jena, Germany. Coptotermes taxonomy remain, but new tools such as Termitenstudien. VI. Versuch einer systematischen 1995, Ruelle 1996, Eggleton 1999). In the 21 - Hozawa,Monographie S. 1915. der Termiten der orientalischen Region. K. Svenska difficultiesties for solving in the problem (Kirton 2005, Vargo and Husseneder Vetensk. Akad. Handl. 50: 1-276. Revision of the Japanese termites. J. Coll. Sci. Tokyo Imp. statistical methods and molecular techniques present new possibili- Jenkins, T. M., R. E. Dean, and B. T. Forschler. 2002. DNA technology, Univ. 35: 1-161. - 2009). Unprecedented levels of international communication, includ interstate commerce, and the likely origin of Formosan subterranean ing sharing gene sequences through GenBank and releasing mor International entomologists must work together just as Holmgren, Jenkins,termite T. (Isoptera:M., S. C. Jones, ) C.-Y. Lee, B. T.infestation Forschler, in Z.Atlanta, Chen, Georgia.G. Lopez- J. phological images through Morphbank, ease information exchange.- Martinez,Econ. Entomol. N. T. 95: Gallagher, 381-389. G. Brown, M. Neal, B. Thistleton, and S. out a revision of Coptotermes Kleinschmidt. 2007. Oshima, Shiraki, Wasmann, and Yano did a hundred years ago. With exotic Coptotermes gestroi Phylogeography illuminates maternal origins of , regular identification for quarantine Kirton, L. G. 2005. The importance (Isoptera: of Rhinotermitidae).accurate termite taxonomyMol. Phylogenet. in the and control purposes cannot be achieved. As Oshima (1910) wrote, 7 Evol. 42: 612-621. In “the correct identification of termite species is a key requirement for broader perspective of termite management, pp. 1-7. Proceedings Acknowledgementstheir control.” Kirton,of the L. Fifth G., and International V. K. Brown. Conference 2003. on Urban Pest, 10-13 July, 2005, - The taxonomic status of pest spe- Suntec,cies of Coptotermes Singapore. in southeast Asia: resolving the paradox in the pest status of the termites Coptotermes gestroi, C. havilandi, and C. travians The authors thank Jing-Fu Tsai (National Chung-Hsing Univer sity) for assistance with specimen photography and identification- Lee, C.-Y. 2002. Subterranean termite pests and their control in the urban of Shiraki’s specimens. We also thank Paul Bardunias, Rudolf (Isoptera: Rhinotermitidae). Sociobiology 42: 43-63. Scheffrahn, and Aaron Mullins (University of Florida) for review Li, G. 2000. Coptotermes In environment in Malaysia. Sociobiology 40: 3-9. ing the manuscript, and Yau-I Chu and Ai-Chi Lin (National Taiwan Li, H.-F. 2009. , pp. 299-341. F. Huang, S. Zhu, Z. Ping, X. He, G. Li University) for providing literature references. We are grateful to Coptotermesand D. Gao [eds.], formosanus Fauna andSinica, Coptotermes Isoptera. Sciencegestroi (Isoptera: Press, Beijing, Rhinotermi China.- Yen-Chiu Lan (Leader University), Shu-Pei Chen and Chi-Feng Lee Phylogeography, interspecific competition, and control of (TARI), Jung-Tai Chao (TFRI), Mei-Ling Chan (NMNS), Man-Miao Yang Li, H.-F., W. Ye, N.-Y. Su, and N. Kanzaki. 2009. Copto- (NCHU), and Tsui-Ying Chang (NPUST) for their assistance in the termestidae) in gestroi Taiwan, and Ph.D. Coptotermes dissertation, formosanus University (Isoptera: of Florida, Rhinotermitidae) Gainesville. We also thank Shinya Miyano (Entomological Society of Japan) and Phylogeography of search and examination of Shiraki and Oshima’s termite collection. Meng-Chun Hsieh (National Taiwan University Library) for process- Lin, S.-Q. 1987. Coptotermes formosanus and its control ing the copyright permission of Oshima’s photograph and illustration, in Taiwan. Ann. Entomol.In Soc. Am. 102: 684-693. Present status of in China, pp. 31-36. M. Tamashiro and N.-Y. Su [eds.], Biology and respectively. This study was supported in part by a grant from USDA- Lo, controlN., R. H. of Eldridge, the Formosan and M. subterranean Lenz. 2006. termite. College of Trop.Copto Agr.- Human Resources, Univ. of Hawaii, Honolulu, HI. ARS under the grant agreement No. 58-6435-2-276. termes (Isoptera: Rhinotermitidae) species inferred from mitochondrial References Cited Phylogeny of Australian Acda, M. N. 2004. - Mori, H. 1987. - COII sequences. Bull. Entomol. Res. 96: 433-437. In Austin, J. W., G. J. Glenn, Economically and R. E.important Gold. 2008. termites (Isoptera) of the Philip- The Formosan subterranean termite in Japan: its distribu pines and their control. Sociobiology 43: 159-168. tion, damage, and current and potential control measures, pp. 23-26. Protecting urban infrastruc M. Tamashiro and N.-Y. Su [eds.], Biology and control of the Formosan ture from Formosan termite (Isoptera: Rhinotermitidae) attack: a case Bess, H. A. 1970. Oshima,subterranean M. 1909. termite. College of Tropical Agriculture,In Human Resources, Instudy for United States railroads. Sociobiology 51: 231-247. University of Hawaii, Honolulu, HI. Termites of Hawaii and the oceanic islands, pp. 449-476. Oshima, M. 1910a.Taiwanese termites, pp. 29-40. M. Oshima [ed.], The Calaby,K. J. Krishna H., and and F. J. F.Gay. M. Weesner1956. The [eds.], distribution Biology and of termites biology II.of Academicthe genus first official report on termites. Taiwan Sōtokufu, Taihoku, Japan. Press, New York. Coptotermes Oshima, M. 1910b. Taiwanese termites. Dobutsugaku Zasshi 22: 343- Constantino, R. 2002. The pest termites of South America: taxonomy, 346. (Isoptera) in Western Australia. Aust. J. Zool. 4: 19-39. Oshima, M. 1911. Taiwanese termites. Dobutsugaku Zasshi 22: In376- 382. distribution and status. J. Appl. Entomol. 126: 355-365. 226 The taxonomy ofAmerican termites Entomologist in Taiwan, pp.• Winter 1-11. 2010 M. Su, N.-Y. 2003. -

Oshima,Oshima M. [ed.],1912. The second official report on termites. Taiwan Sōtokufu, Su, N.-Y., and R. OverviewH. Scheffrahn. of the 1998. global Coptotermes distribution vastator and control Light of (Isoptera: the For Taihoku,In Japan. mosan subterranean termite. Sociobiology 41: 7-16. The taxonomy and distribution of termites in Taiwan, pp. Su, N.-Y., and R. H. Scheffrahn. 2000. Oshima,54-94. M. 1919.M. Oshima [ed.], The third official report on termites. Taiwan Rhinotermitidae)In in Guam. Proc. Hawaii. Entomol. Soc. 33: 13-18. Sōtokufu, Taihoku, Japan. Termites as pests of buildings, pp. Ping, Z. 1991. Formosan termites and methods of preventing their 437-453. T. Abe, D. E. Bignell and M. Higashi [eds.], Termites: evolution, damage. Philipp. J. Sci. 15: 319-383. Tsai,sociality, C.-C., and symbioses, C.-S. Chen. ecology. 2003. Kluwer AcademicCoptotermes Publishers, gestroi Dordrecht, (Isop- Roonwal, M. L., Termite and O. B. problem Chhotani. in plant 1962. quarantine. Indian species Plant of Quarantinetermite genus 5: The Netherlands. Coptotermes53-59. Vargo, E. L., and C. Husseneder. First2009. record Biology of of subterranean termites: Roonwal, M. L., and O. B. Chhotani. 1989. The fauna of India and the tera:Insights Rhinotermitidae) from molecular from studies Taiwan. of Reticulitermes Formosan andEntomol. Coptotermes 23: 157-161. . Government of India, Delhi, India. Ruelle, J. E. 1996. Wang, C., J. E. Powell, and Y. Liu. 2002. A literature review of the biology. Annu. adjacent countries, Vol. 1. Zoological Survey of India, Calcutta, India. Rev.and ecologyEntomol. of 54: Coptotermes 379-403. formosanus (Isoptera: Rhinotermitidae) in Scheffrahn, R. H., andComments N.-Y. Su. on 2005. “taxonomic Distribution splitters of inthe China”. termite Isoptera genus CoptotermesNewsletter 6: 1. Wasmann, E. 1896. China. Sociobiology 40: 343-363. Scheffrahn, R. H., (Isoptera: N.-Y. Su, Rhinotermitidae) and B. Diehl. 1990. in Florida. Native, Fla. introduced, Entomol. and88: Neue Termitophilen und Termiten aus Indien. Viaggo 201-203. Woodrow,di Leonardo R. J., J. Fea K. Grace,in Birmania and S. E Y. Regioni Higa. 2001. Vicine. Occurrence Annali del of Museo Coptotermes civico vastatordi storia naturale di Genova 16: 613-630. structure-infesting termites of the Turks and Caicos Islands, B.W.I. Scheffrahn,(Isoptera: R. Kalotermitidae, H., J. Krecek, B. Rhinotermitidae, Maharajh, N.-Y. Termitidae). Su, J. A. Chase, Fla. J. Entomol. R. Man- Yano, M. 1911. (Isoptera: Rhinotermitidae) on the island of Oahu, Dobutsugaku Hawaii. gold,73: 622-627. A. L. Szalanski, J. W. Austin, and J. Nixon. 2004. Establishment Sociobiology 38: 667-673. of the African termite, Coptotermes sjostedti (Isoptera: Rhinotermitidae), Yudin, L. 2002.On the scientific names of Japanese termites. Zasshi 23: 364-368. Termites of Mariana Islands and Philippines, their damage Shiraki,on the T. Island1909. of Guadeloupe, French West Indies. Ann. Entomol. Soc. Hou-Fengand control. Li and Sociobiology Nan-Yao Su 40: 71-74. Snyder,Am. 97:T. E. 872-876. 1949. - Japanese termites. Trans. Entomol. Soc. Japan 2: 229-242. are a Post—Doctoral Research Associate and- Sornnuwat, Y. 1996. Catalog Studies of on the damage termites of constructions (Isoptera) of thecaused world. by subter Smith- a Professor of Entomology, respectively, in the Fort Lauderdale Research and sonian Misc. Coll. 112: 1-490. Education Center, University of Florida. Their research interests include gen Su, N.-Y. 2002. - eral termite biology and control of subterranean termite. 3205 College Ave, ranean termites and its control in Thailand. Wood Res. 83: 59-139. Davie, FL33314. [email protected]; [email protected]. Wen-Jer Wu is a Professor Novel technologies for subterranean termite control. So of Entomology, National Taiwan University. [email protected]. ciobiology 40: 95-101.

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