The genus Coptotermes Wasmann 1896 is a large group of

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1 Solving the Hundred-Year Controversy of Coptotermes Taxonomy in Taiwan Hou-Feng Li, Nan-Yao Su, and Wen-Jer Wu The genus Coptotermes Wasmann 1896 is a large group of subterranean termites, including 70 described species (Constantino 2009, Vargo and Husseneder 2009), 28 of which are considered important structural pests (Edwards and Mill 1986, Su and Scheffrahn 2000). The global economic impact of all subterranean termites has been estimated at $22 billion annually (Su 2002), and Coptotermes has been reported as the most destructive of the group in the southern United States (Su 2003, Scheffrahn and Su 2005), Hawaii (Bess 1970, Woodrow et al. 2001), Brazil (Ferraz and Cancello 2001, Constantino 2002), Australia (Calaby and Gay 1956, Lo et al. 2006), southern China (Lin 1987, Wang et al. 2002), Taiwan (Oshima 1919, Li 2009), southern Japan (Mori 1987), Philippines and Mariana Islands (Su and Scheffrahn 1998, Yudin 2002, Acda 2004), Thailand (Sornnuwat 1996), Malaysia (Lee 2002), and India (Roonwal and Chhotani 1989). Coptotermes spp. have frequently been intercepted at ports (Gay 1967, Ping 1991), and some have become established in non-native areas (Gay 1969, Scheffrahn et al. 1990, 2004; Jenkins et al. 2007). They are continually dispersed by humanaided land transportation (Jenkins et al. 2002, Austin et al. 2008). The tendency of Coptotermes spp. to colonize cargo, boats, and ships may contribute to their wide distribution, especially in C. formosanus Shiraki and C. gestroi (Wasmann) (Scheffrahn and Su 2005). Despite its economic importance, taxonomy of the genus is still problematic. The inherent difficulties in Coptotermes taxonomy include: Lack of distinguishing features. Soldiers and alates are the two main castes used for termite identification. Soldiers among Coptotermes spp. are superficially similar (Calaby and Gay 1956). Few characters of the soldier are available for species identification (Kirton 2005), and a microscopic examination is usually required for observing diagnostic characters, such as the number of setae around the fontanelle. Alates offer more differentiable characters for species identification, including body length, body color, antennal spot pattern, and dispersal flight season. However, alates are only present in mature colonies, and for a short period of time each year. High degree of morphological variation among intraspecific populations. By using quantitative comparison of morphological characters, individuals of the same species collected from colonies at different ages and habitats likely show significant differences. Without comparing adequate numbers of samples, a plethora of new species have been proposed over the same geographic areas. For example, at least 12 synonyms have been recorded for C. formosanus alone (Snyder 1949, Li 2000). Difficulty in matching soldiers and alates of the same species. Soldiers and alates of the same colony are rarely collected together. Soldiers associated with damaged wood can be collected year-round, but alates are usually collected during their dispersal flight. Hence, alates and soldiers of the same species might be described as two species, as occurred with C. gestroi and C. havilandi Holmgren (Kirton and Brown 2003). To date, only half of Coptotermes spp. are based on descriptions of both soldier and alate castes (Snyder 1949, Roonwal and Chhotani 1962, Li 2000). Since most Coptotermes pest species originated in Asia, entomologists in these areas encountered the taxonomic challenge early. The identification of Coptotermes species in Taiwan and southern Japan was a controversial issue between 1909 and Many influential Japanese entomologists and European termite experts were involved in this debate. One hundred years after the controversy, we review this historical record, clarify the controversial issues based on our current study, and offer solutions to avoid similar problems in the future. Background of Termite Study in Taiwan in the Early 1900s After the First Sino-Japanese War between the Qing Dynasty of China and the Meiji government of Japan, Taiwan became a Japanese territory in 1895 under the Treaty of Shimonoseki until the end of WWII in Since most of Japan has a temperate climate and termites are primarily subtropical and tropical pests, Japanese construction practices were not adapted for termite prevention. Thus, structures built in Taiwan during the early Japanese colonial period suffered from severe termite damage (Oshima 1919). Termite control became a priority of the Governor of Taiwan in the early 1900s. In 1907, both Tokuichi Shiraki ( ) (Fig. 1A) and Masamitsu Oshima ( ) (Fig. 1B) started to work in Taiwan at the 222 American Entomologist Winter 2010

2 Fig. 1. (A) Tokuichi Shiraki ( ) (Photo provided by the Entomology Dept. of the National Taiwan University); (B) Masamitsu Oshima ( ) (Photo provided by the Entomological Society of Japan). Agricultural Experiment Station, Government of Formosa (predecessor of Taiwan Agricultural Research Institute) and the Bureau of Civil Engineering. Later, they became the leading figures in termite research on the island. Their major control target was known at the time colloquially as house termites, which we now know to be Coptotermes species. Controversy in the Identification of Coptotermes spp. in Taiwan Shiraki and Oshima cooperated on the studies of termite control and taxonomy during the first two years of their careers in Taiwan. Shiraki (1909) briefly described C. formosanus, in Japanese, based on Oshima s collection from many locations in Taiwan. The article included descriptions of the morphology of several castes, including the male and female alate, nymph, soldier, worker, and queen, for which no figure was offered. In addition, the type locality and type specimen of C. formosanus were not included. The species name was mentioned twice, but incorrectly spelled as Captotermes formosanus [sic] and Coptotermes forrmosanus [sic] (Shiraki 1909). In the same year, based on Shiraki s description, Oshima (1909) offered a more detailed examination of the morphological characteristics of C. formo- sanus. This included illustrations of the dealate, soldier, worker, fore and hind wings (Fig. 2), and photographs of nests. Twelve collection locations in Taiwan and the Penghu islands were also described (Fig. 3A). Oshima (1909) vividly and comprehensively described damage caused by C. formosanus, its tunneling behavior, and soldier behaviors such as head banging and secreting from the fontanelle. In the following year, Oshima (1910a) questioned the validity of several termite species named by Shiraki (1909), including C. formosanus. Oshima (1910a) emphasized that there was no personal animosity between Shiraki and himself, and they worked together and shared references, books, and termite samples with each other. He had no wish to offend Shiraki, but he believed that the correct identification of termite species was a key requirement for their control. Oshima (1910b, 1911) mentioned that the soldiers of C. formosanus superficially resembled those of C. gestroi described by Haviland in Oshima did not compare the alates of these two species, probably due to absence of alate description of C. gestroi (Wasmann 1896, Haviland 1898). Since Taiwan was geographically close to Southeast Asia, where C. gestroi occurred, and no significant morphological difference of the soldier caste was found, Oshima (1910b, 1911) proposed that C. formosanus was a junior synonym Fig. 3. Coptotermes spp. collection sites in early 1900s (A) and in the current study (B). White area, altitude >500 m; light gray area, subtropical lowland; dark gray area, tropical lowland. Fig. 2. Coptotermes formosanus drawn by Oshima in 1909 provided by the National Taiwan University Library. Dealate (A); soldier (B); worker (C); fore (D) and hind (E) wings. Vertical scale bar for A-C; horizontal scale bar for D-E. American Entomologist Volume 56, Number 4 of C. gestroi. Oshima also provided 12 collection locations of C. gestroi/ formosanus in Taiwan and the Penghu islands (Oshima 1911) (Fig. 3A). In addition to Shiraki and Oshima, several other Japanese researchers such as Munemoto Yano ( ) of Forestry Experimental Station, Tokyo, Shozaburo Watasé ( ) and his student, Sanji Hozawa ( ) of the Zoological Institute, Science College, Tokyo Imperial University also became involved in identification of Coptotermes spp. of Taiwan and southern Japan. Yano (1911) mentioned that Watasé sent Japanese specimens to a German termite taxonomist, Eric Wasmann ( ), for identification. Wasmann believed that these samples were C. gestroi, which he named in However, Yano (1911) still had some doubt about Wasmann s opinion. Yano mentioned that the description of C. gestroi by Wasmann was too vague to differentiate it from other Coptotermes spp. Hence, Yano further compared Japanese/Taiwanese Coptotermes samples with the description of C. gestroi of Haviland (1898). He 223

3 thought the two species were similar, but that their soldiers could be distinguished by the ratio of head width to head length. The Japanese/Taiwanese soldier samples had elongated heads (1.5 mm in length and 1.2 mm in width) while C. gestroi as described by Haviland (1898) had more circular heads (1.4 mm in length and 1.3 mm in width). Additionally, Yano sent the Japanese/Taiwanese Coptotermes samples to a Swedish termite taxonomist, Nils Holmgren ( ), through Chiyomatsu Ishikawa ( ) of Tokyo Imperial University and another German termite researcher, Karl Escherich ( ). Holmgren examined Yano s specimens and described a new species, Coptotermes formosae Holmgren (Holmgren 1911), and compared it with other Coptotermes spp. of Sri Lanka. Based on Holmgren s opinion and his own measurements, Yano (1911) concluded that the Coptotermes sp. collected in Japan and Taiwan was not C. gestroi. Yano (1911) also criticized Shiraki s description of C. formosanus (Shiraki 1909), in which no figures were offered, the format was informal, and the description was in Japanese. Yano thought it should be appropriate to adopt Holmgren s nomenclature, C. formosae (Holmgren 1911), instead of C. formosanus named by Shiraki (1909), because Holmgren was a recognized authority on termite taxonomy. Oshima (1912) was torn between the conflicting opinions of the two internationally recognized authorities on termite taxonomy, Wasmann and Holmgren. He agreed with a part of Yano s opinions (1911) that there was a difference between Japanese/Taiwanese samples and C. gestroi in the ratio of head width and head length of the solider. However, he thought it was insufficient to name a new species (herein C. formosanus) because the difference might be due to the intraspecific variation. This was the major reason that he doubted the validity of C. formosanus (Oshima 1910b, 1911). To solve this controversial issue, Oshima collected more samples and measured head length and width with a more accurate method based on a clear definition. He had seven soldiers from two locations in southern Japan and 11 soldiers from three locations in Taipei, Taiwan (Oshima 1912, Fig. 3A). He removed the soldier heads and placed them on a glass plate in a horizontal position, then measured the distance between the medial base of the labrum to the hindmost margin of the head capsule. The data was rounded to two decimal places. Oshima found the head length ( mm) of Japanese/ Taiwanese samples to be much longer than that of C. gestroi (1.4 mm) as described by Haviland (1898). No significant difference was found between Japanese and Taiwanese samples (Taipei only), and the variation of these samples in head length was only 0.03 mm. It is worthy of note that the head length of soldiers reported by Oshima in 1910b and 1911 was 1.5 mm, but it increased to mm when he made another measurement in Oshima attributed the difference to different measuring methods. Based on the new measurements, Oshima confidently believed that Coptotermes samples collected in Japan and Taiwan (Taipei materials) were not C. gestroi. Oshima (1912) thought that the alate of C. formosanus described by Shiraki (1909) was similar to C. formosae described by Holmgren (1911). Since C. formosanus was described first, C. formosae should be a junior synonymy based on the rules of zoological nomenclature (Oshima 1912). Oshima (1912) also severely criticized Yano s attempt (Yano 1911) to invalidate C. formosanus based on the publication format and language used for description instead of the nomenclature rules and the content of the description. Oshima listed the original description of C. formosae (Holmgren 1911) in The Third Official Report on Termites (Oshima 1912) to show that Holmgren 224 only offered five measurements of alates, which was less informative than the first description of C. formosanus (Shiraki 1909), and that Holmgren failed to provide illustrations or photographs. After The Third Official Report on Termites was published by Oshima in 1912, most termite researchers such as Holmgren (1913) and Hozawa (1915) accepted Oshima s opinion that C. formosanus is the only Coptotermes species in southern Japan and Taiwan, and C. formosae is a junior synonymy of C. formosanus. Coptotermes gestroi in Taiwan After 1912, C. formosanus was believed to be the only Coptotermes species in Taiwan. Coptotermes gestroi in Taiwan had not been mentioned until Based on soldier morphology, the new record of C. gestroi in Taiwan was proposed again (Tsai and Chen 2003). The first author of the current study and local pest control operators collected Coptotermes spp. from every county in Taiwan, both in urban areas and natural environments, from With the mitochondrial gene sequence data, the presence of C. gestroi in southern Taiwan was confirmed (Li et al. 2009). In total, 220 Coptotermes samples including museum specimens were identified and used for mapping their distribution (Fig. 3B). Coptotermes spp. were mostly collected in lowland areas (< 500 m). Coptotermes formosanus was distributed throughout the island of Taiwan, while C. gestroi was only collected in the southern tropical zone. In a review of articles published in the early 1900s, some evidence shows that C. gestroi may have already been present in Taiwan. Oshima (1909, 1911) collected Coptotermes samples from 12 locations in Taiwan (Fig. 3A), seven of which were in the tropical zone where C. gestroi and C. formosanus currently coexist. The drawing of a dark brown dealate (Fig. 2A) by Oshima (1909) resembled C. gestroi (Fig. 4A) instead of the light brownish-yellow dealate of C. formosanus (Fig. 4C). The color of soldier heads of C. gestroi (Fig. 4B) and C. formosanus (Fig. 4D) is light yellow, which is similar to the color of the C. formosanus alate abdomen (Fig. 4C), but in contrast to the dark brown color of the C. gestroi alate abdomen (Fig. 4A). Based on the color difference between soldier and alate, Oshima s drawing (Fig. 2A and 2B) was likely based on C. gestroi. In addition, the alate body length was recorded as 6 mm (Oshima 1909), which is closer to C. gestroi than C. formosanus (usually >7 mm). The drawing of the soldier head capsule (Fig. 5A, Oshima 1909) was somewhat rounded and was also Fig. 4. Dealate (A) and soldier (B) of C. gestroi, and dealate (C) and soldier (D) of C. formosanus collected in southern and northern Taiwan, respectively, in the current study. American Entomologist Winter 2010

4 Fig. 5. Soldier head capsules of Coptotermes spp. A solider collected in Taiwan by Oshima in 1909 resembled C. gestroi (A) (Photo provided by the National Taiwan University Library); C. formosanus collected in Taipei or southern Japan by Oshima in 1912 (B) (Photo provided by the Entomology Dept. of the National Taiwan University); SEM pictures of C. gestroi (C) and C. formosanus (D) collected in southern and northern Taiwan, respectively, in the current study. similar to C. gestroi (Fig. 5C), and its head length was recorded as 1.5 mm. Three years after his publication (Oshima 1909) in 1912, Oshima collected Coptotermes samples only in northern Taiwan (Taipei city) (Fig. 3A) and in Japan, where only C. formosanus is currently found. The image of a soldier head capsule (Fig. 5B, Oshima 1912) was elongated and similar to C. formosanus (Fig. 5D). The head length of soldiers reported by Oshima in 1912 was mm. Soldiers collected in 1909 and 1911 might include both C. formosanus and C. gestroi; hence, the average of their head length was shorter than that of C. formosanus collected only from Taipei in Oshima (1912) emphasized twice there was only one Coptotermes sp. in Taiwan. However, no effort was made to prove that all the Coptotermes samples in Taiwan were the same species (Oshima 1909, 1910b, 1911). Based on this false assumption, Coptotermes samples collected at many places in Taiwan were lumped together, and the mean of morphological measurements of the two species may have been presented as a single species, C. formosanus. When Holmgren and Wasmann identified Japanese/Taiwanese Coptotermes samples as C. formosae and C. gestroi, respectively (Yano 1911), Oshima and Yano were confused, but they did not suspect that there might be two Coptotermes 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, which may be the source of the controversy in early 1900s. A set of specimens (Fig. 6A) labeled Coptotermes formosanus Shiraki was found in TARI, where Shiraki worked between 1907 and The autograph on the label (Fig. 6B) is very similar to Shiraki s handwriting in an unpublished manuscript entitled Insect Fauna of Taiwan (Fig. 6C) offered by Wen-Jer Wu of the National Taiwan University. Termite soldiers preserved in vials (Fig. 6D) with two setae on each side of the fontanelle (Fig. 6D, inset) were identified as C. formosanus (C. gestroi has one seta on each side; Scheffrahn et al. 1990), but no collection information of these soldiers was found. Three alate specimens were labeled, but damaged to some degree. The best-preserved sample (Fig. 6E) was collected on June 10th, 41st year of Meiji era (1908) from a location called Old Farm, which is unknown to us, and the collector was not recorded. The other two severely damaged samples (Fig. 6F and 6G) were collected by Inao Nitobe ( ), Shiraki s assistant, on April 23rd, 41st year of Meiji era (1908) from Pingtung county, Gangkou research station of TFRI (Fig. 3A). A comparison of these three alates with the original description (Shiraki 1909) and our recent collections (Li et al. 2009) has confirmed them to be C. formosanus. Even though these C. formosanus specimens preserved in TARI once belonged to Shiraki and might have been used for describing C. formosanus in 1909, the collector of these specimens does not fit the original description (Shiraki 1909). Shiraki (1909) mentioned that the new species, C. formosanus, was described based on Oshima s collection. However, no part of Oshima s collection was found in the six insect collections Type Specimen of C. formosanus During our investigation into the controversy on the identification of C. formosanus and C. gestroi in the early 1900s, museum specimens preserved in six major insect collections in Taiwan were examined to search for those used by Shiraki and Oshima: 1. Taiwan Agricultural Research Institute, Insect and Mite Collection, Wufeng, Taichung, Taiwan, ROC (TARI) 2. Taiwan Forestry Research Institute, Insect Collection, Taipei, Taiwan, ROC (TFRI) 3. National Museum of Natural Science, Taichung, Taiwan, ROC (NMNS) 4. National Taiwan University, Department of Entomology, Insect Collection, Taipei, Taiwan, ROC (NTU) 5. National Chung-Hsing University, Department of Entomology, Insect Collection, Taichung, Taiwan, ROC (NCHU) 6. National Pingtung University of Science and Technology, Department of Plant Medicine, Insect Collection, Neipu, Pingtung, Taiwan, ROC (NPUST) American Entomologist Volume 56, Number 4 Fig. 6. Shiraki s C. formosanus collection preserved in the Taiwan Agricultural Research Institute. Entire collection with an identification label; workers, soldiers in vials, and three pinned alates (A); the autography on the label (B) is similar to Shiraki s handwriting in an unpublished manuscript entitled Insect Fauna of Taiwan (C); soldiers (D) preserved in vials with two setae (inset) on one side of the fontanelle; the most well-preserved 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 on April 23rd, 41st year of Meiji era (1908) at Gangkou. Photos of labels and insects not to scale. 225

5 of Taiwan. In order to prevent any further confusion on identifying C. formosanus, we designated neotypes herein. NEOTYPE: male alate collected by H.-F. Li at TAIWAN, Taoyuan Co., Taoyuan City: N, E; 20-VI-2006 (TW49) will be deposited in NMNS. The neotype perfectly fit the re-description by Hozawa (1915). NEOPARATYPES: female and male alates, and soldiers collected from the same colony as the neotype will be deposited in NMNS, NTU, TFRI, and University of Florida Termite Collection, Fort Lauderdale Research and Education Center. Mitochondrial genes including COII, 12S rrna, and 16S rrna of workers collected from the same colony with the neotype have been partially sequenced and submitted to the GenBank database with accession numbers: EU805758, EU805712, and EU (Li et al. 2009). This review of the historical controversy is intended to present the challenge for Coptotermes taxonomy, and to prompt international cooperation on a revision of Coptotermes. The genus Coptotermes is in serious need of revision, especially the Chinese species (Crosland 1995, Ruelle 1996, Eggleton 1999). In the 21 st century, the inherent difficulties in Coptotermes taxonomy remain, but new tools such as statistical methods and molecular techniques present new possibilities for solving the problem (Kirton 2005, Vargo and Husseneder 2009). Unprecedented levels of international communication, including sharing gene sequences through GenBank and releasing morphological images through Morphbank, ease information exchange. International entomologists must work together just as Holmgren, Oshima, Shiraki, Wasmann, and Yano did a hundred years ago. Without a revision of Coptotermes, regular identification for quarantine and control purposes cannot be achieved. As Oshima (1910) wrote, the correct identification of termite species is a key requirement for their control. 7 Acknowledgements The authors thank Jing-Fu Tsai (National Chung-Hsing University) for assistance with specimen photography and identification of Shiraki s specimens. We also thank Paul Bardunias, Rudolf Scheffrahn, and Aaron Mullins (University of Florida) for reviewing the manuscript, and Yau-I Chu and Ai-Chi Lin (National Taiwan University) for providing literature references. We are grateful to Yen-Chiu Lan (Leader University), Shu-Pei Chen and Chi-Feng Lee (TARI), Jung-Tai Chao (TFRI), Mei-Ling Chan (NMNS), Man-Miao Yang (NCHU), and Tsui-Ying Chang (NPUST) for their assistance in the search and examination of Shiraki and Oshima s termite collection. We also thank Shinya Miyano (Entomological Society of Japan) and Meng-Chun Hsieh (National Taiwan University Library) for processing the copyright permission of Oshima s photograph and illustration, respectively. 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6 Oshima [ed.], The second official report on termites. Taiwan Sōtokufu, Taihoku, Japan. Oshima, M The taxonomy and distribution of termites in Taiwan, pp In M. Oshima [ed.], The third official report on termites. Taiwan Sōtokufu, Taihoku, Japan. Oshima, M Formosan termites and methods of preventing their damage. Philipp. J. Sci. 15: Ping, Z Termite problem in plant quarantine. Plant Quarantine 5: Roonwal, M. L., and O. B. Chhotani Indian species of termite genus Coptotermes. Government of India, Delhi, India. Roonwal, M. L., and O. B. Chhotani The fauna of India and the adjacent countries, Vol. 1. Zoological Survey of India, Calcutta, India. Ruelle, J. E Comments on taxonomic splitters in China. Isoptera Newsletter 6: 1. Scheffrahn, R. H., and N.-Y. Su Distribution of the termite genus Coptotermes (Isoptera: Rhinotermitidae) in Florida. Fla. Entomol. 88: Scheffrahn, R. H., N.-Y. Su, and B. Diehl Native, introduced, and structure-infesting termites of the Turks and Caicos Islands, B.W.I. (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae). Fla. Entomol. 73: Scheffrahn, R. H., J. Krecek, B. Maharajh, N.-Y. Su, J. A. Chase, J. R. Mangold, A. L. Szalanski, J. W. Austin, and J. Nixon Establishment of the African termite, Coptotermes sjostedti (Isoptera: Rhinotermitidae), on the Island of Guadeloupe, French West Indies. Ann. Entomol. Soc. Am. 97: Shiraki, T Japanese termites. Trans. Entomol. Soc. Japan 2: Snyder, T. E Catalog of the termites (Isoptera) of the world. Smithsonian Misc. Coll. 112: Sornnuwat, Y Studies on damage of constructions caused by subterranean termites and its control in Thailand. Wood Res. 83: Su, N.-Y Novel technologies for subterranean termite control. Sociobiology 40: Su, N.-Y Overview of the global distribution and control of the Formosan subterranean termite. Sociobiology 41: Su, N.-Y., and R. H. Scheffrahn Coptotermes vastator Light (Isoptera: Rhinotermitidae) in Guam. Proc. Hawaii. Entomol. Soc. 33: Su, N.-Y., and R. H. Scheffrahn Termites as pests of buildings, pp In T. Abe, D. E. Bignell and M. Higashi [eds.], Termites: evolution, sociality, symbioses, ecology. Kluwer Academic Publishers, Dordrecht, The Netherlands. Tsai, C.-C., and C.-S. Chen First record of Coptotermes gestroi (Isoptera: Rhinotermitidae) from Taiwan. Formosan Entomol. 23: Vargo, E. L., and C. Husseneder Biology of subterranean termites: Insights from molecular studies of Reticulitermes and Coptotermes. Annu. Rev. Entomol. 54: Wang, C., J. E. Powell, and Y. Liu A literature review of the biology and ecology of Coptotermes formosanus (Isoptera: Rhinotermitidae) in China. Sociobiology 40: Wasmann, E Neue Termitophilen und Termiten aus Indien. Viaggo di Leonardo Fea in Birmania E Regioni Vicine. Annali del Museo civico di storia naturale di Genova 16: Woodrow, R. J., J. K. Grace, and S. Y. Higa Occurrence of Coptotermes vastator (Isoptera: Rhinotermitidae) on the island of Oahu, Hawaii. Sociobiology 38: Yano, M On the scientific names of Japanese termites. Dobutsugaku Zasshi 23: Yudin, L Termites of Mariana Islands and Philippines, their damage and control. Sociobiology 40: Hou-Feng Li and Nan-Yao Su are a Post Doctoral Research Associate and a Professor of Entomology, respectively, in the Fort Lauderdale Research and Education Center, University of Florida. Their research interests include general termite biology and control of subterranean termite College Ave, Davie, FL houfeng@ufl.edu; nysu@ufl.edu. Wen-Jer Wu is a Professor of Entomology, National Taiwan University. wuwj@ntu.edu.tw. American Entomologist Volume 56, Number 4 227