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ThegenusCoptotermesWasmann1896 isa largegroupofsubterranean termites, including ≈70 described species (Constantino 2009, Vargo and Husseneder 2009), 28 of

whichareconsideredimportantstructuralpests(EdwardsandMill1986, Su and Scheffrahn 2000). The global economic impact of all subterraneantermiteshasbeenestimatedat$22billionannually(Su2002),andCoptotermeshasbeenreportedasthemostdestructiveofthe group in the southern United States (Su 2003, Scheffrahn and Su 2005), Hawaii (Bess 1970, Woodrow et al. 2001), Brazil (Ferraz and Cancello2001,Constantino2002),Australia(CalabyandGay1956,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(Sornnuwat1996),Malaysia(Lee2002),andIndia(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 human-aided land transportation (Jenkins et al. 2002, Austin et al. 2008). ThetendencyofCoptotermes spp. to colonize cargo, boats, and ships maycontributetotheirwidedistribution,especiallyinC. formosanus ShirakiandC. gestroi (Wasmann) (Scheffrahn and Su 2005).

Despiteitseconomicimportance,taxonomyofthegenusisstillproblematic. The inherent difficulties in Coptotermes taxonomyinclude:

Lack of distinguishing features.Soldiersandalatesarethetwomain castes used for termite identification. Soldiers among Cop-totermes 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 observingdiagnosticcharacters,suchasthenumberofsetaearoundthe fontanelle. Alates offer more differentiable characters for species identification, including body length, body color, antennal spot pat-tern, 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,individualsofthesamespeciescollectedfromcoloniesat 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,atleast12synonymshavebeenrecordedforC. formosanusalone (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. Soldiersassociatedwithdamagedwoodcanbecollectedyear-round,but alates are usually collected during their dispersal flight. Hence, alatesandsoldiersofthesamespeciesmightbedescribedastwospecies,asoccurredwithC. gestroiandC. havilandiHolmgren(Kirtonand Brown 2003). To date, only half of Coptotermes spp. are based on descriptionsofbothsoldierandalatecastes(Snyder1949,Roonwaland Chhotani 1962, Li 2000).

SincemostCoptotermespestspeciesoriginatedinAsia,entomolo-gists in these areas encountered the taxonomic challenge early. The identification of CoptotermesspeciesinTaiwanandsouthernJapanwas a controversial issue between 1909 and 1912. Many influential JapaneseentomologistsandEuropeantermiteexpertswereinvolvedin this debate. One hundred years after the controversy, we review thishistoricalrecord,clarifythecontroversialissuesbasedonourcurrent study, and offer solutions to avoid similar problems in the future.

Background of Termite Study in Taiwan in the Early 1900sAfter the First Sino-Japanese War between the Qing Dynasty of

ChinaandtheMeijigovernmentofJapan,TaiwanbecameaJapaneseterritoryin1895undertheTreatyofShimonosekiuntiltheendofWWII in 1945. Since most of Japan has a temperate climate and termitesareprimarilysubtropicalandtropicalpests,Japanesecon-struction practices were not adapted for termite prevention. Thus, structuresbuiltinTaiwanduringtheearlyJapanesecolonialperiodsuffered 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 (1882-1970) (Fig. 1A) and Masamitsu Oshima (1884-1965) (Fig. 1B) started to work in Taiwan at the

Solving the Hundred-Year Controversy of Coptotermes Taxonomy

in Taiwan

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

Solving the Hundred-Year Controversy of Coptotermes Taxonomy

in Taiwan

222

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Agricultural Experiment Station, Government of Formosa (prede-cessorofTaiwanAgriculturalResearchInstitute)andtheBureauofCivil 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

ShirakiandOshimacooperatedonthestudiesoftermitecontroland taxonomy during the first two years of their careers in Taiwan. Shiraki (1909) briefly described C. formosanus,inJapanese,basedon Oshima’s collection from many locations in Taiwan. The article includeddescriptionsofthemorphologyofseveralcastes,includingthe male and female alate, nymph, soldier, worker, and queen, for which no figure was offered. In addition, the type locality and type specimenofC. formosanus were not included. The species name was mentionedtwice,butincorrectlyspelledasCaptotermes formosanus[sic]andCoptotermes forrmosanus [sic] (Shiraki 1909). In the same year, based on Shiraki’s description, Oshima (1909) offered a more detailedexaminationofthemorphologicalcharacteristicsofC. 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)vividlyandcomprehensivelydescribeddamagecausedbyC. formosanus,itstunnelingbehavior,andsoldierbehaviorssuch as head banging and secreting from the fontanelle.

In the following year, Oshima (1910a) questioned the validity ofseveral termitespeciesnamedbyShiraki (1909), includingC. formosanus. Oshima (1910a) emphasized that there was no personal animositybetweenShirakiandhimself,andtheyworkedtogetherand 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. gestroidescribedbyHaviland in 1898. Oshima did not compare the alates of these two species,probablyduetoabsenceofalatedescriptionofC. gestroi(Wasmann 1896, Haviland 1898). Since Taiwan was geographically closetoSoutheastAsia,whereC. gestroi occurred, and no significant morphological difference of the soldier caste was found, Oshima (1910b,1911)proposedthatC. formosanuswasajuniorsynonym

Fig. 1. (A) Tokuichi Shiraki (1882-1970) (Photo pro-vided by the Entomology Dept. of the National Taiwan University); (B) Masamitsu Oshima (1884-1965) (Photo provided by the Entomological Society of Japan).

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.

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.

ofC. gestroi. Oshima also provided 12 collection locations of C. ge-stroi/ formosanus in Taiwan and the Penghu islands (Oshima 1911) (Fig. 3A).

In addition to Shiraki and Oshima, several other Japaneseresearchers such as Munemoto Yano (1883-1970) of Forestry Experimental Station, Tokyo, Shozaburo Watasé (1862-1929) and his student, Sanji Hozawa (1885-1947) of the Zoological Institute, ScienceCollege,TokyoImperialUniversityalsobecameinvolvedinidentification of Coptotermes spp. of Taiwan and southern Japan. Yano (1911) mentioned that Watasé sent Japanese specimens to a German termite taxonomist, Eric Wasmann (1859-1931), for identification. Wasmannbelieved that thesesampleswereC. gestroi,whichhenamed in 1896. However, Yano (1911) still had some doubt about Wasmann’s opinion. Yano mentioned that the description of C. gestroiby Wasmann was too vague to differentiate it from other Coptotermes spp. Hence, Yano further compared Japanese/Taiwanese CoptotermessampleswiththedescriptionofC. gestroi of Haviland (1898). He

American Entomologist•Winter2010224

thoughtthetwospeciesweresimilar,butthattheirsoldierscouldbe distinguished by the ratio of head width to head length. The Japa-nese/Taiwanese soldier samples had elongated heads (1.5 mm in length and 1.2 mm in width) while C. gestroiasdescribedbyHaviland(1898) had more circular heads (1.4 mm in length and 1.3 mm in width). Additionally, Yano sent the Japanese/Taiwanese Coptotermessamples to a Swedish termite taxonomist, Nils Holmgren (1877-1954),throughChiyomatsuIshikawa(1861-1935)ofTokyoImperialUniversityandanotherGermantermiteresearcher,KarlEscherich(1871-1951). Holmgren examined Yano’s specimens and described anewspecies,Coptotermes formosaeHolmgren(Holmgren1911),andcompareditwithotherCoptotermes spp. of Sri Lanka. Based on Holmgren’s opinion and his own measurements, Yano (1911) concludedthattheCoptotermes sp. collected in Japan and Taiwan wasnotC. gestroi. Yano (1911) also criticized Shiraki’s description ofC. formosanus (Shiraki 1909), in which no figures were offered, the format was informal, and the description was in Japanese. Yano thoughtitshouldbeappropriatetoadoptHolmgren’snomenclature,C. formosae(Holmgren1911),insteadofC. formosanusnamedbyShiraki (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 samplesandC. gestroiintheratioofheadwidthandheadlengthofthe solider. However, he thought it was insufficient to name a new species(hereinC. formosanus) because the difference might be due to the intraspecific variation. This was the major reason that he doubted thevalidityofC. formosanus (Oshima 1910b, 1911).

Tosolvethiscontroversialissue,Oshimacollectedmoresamplesandmeasuredheadlengthandwidthwithamoreaccuratemethodbased on a clear definition. He had seven soldiers from two locations insouthern Japanand11soldiers fromthree locations inTaipei,Taiwan (Oshima 1912, Fig. 3A). He removed the soldier heads and placed them on a glass plate in a horizontal position, then measured thedistancebetweenthemedialbaseofthelabrumtothehindmostmargin of the head capsule. The data was rounded to two decimal places. Oshima found the head length (1.66–1.72 mm) of Japanese/TaiwanesesamplestobemuchlongerthanthatofC. gestroi (1.4 mm) as described by Haviland (1898). No significant difference was found betweenJapaneseandTaiwanesesamples(Taipeionly),andthevaria-tion of these samples in head length was only 0.03 mm. It is worthy of notethattheheadlengthofsoldiersreportedbyOshimain1910band1911 was 1.5 mm, but it increased to 1.66–1.72 mm when he made another measurement in 1912. Oshima attributed the difference to different measuring methods. Based on the new measurements, Oshima confidently believed that CoptotermessamplescollectedinJapanandTaiwan(Taipeimaterials)werenotC. gestroi.

Oshima(1912)thoughtthatthealateofC. formosanusdescribedbyShiraki(1909)wassimilartoC. formosaedescribedbyHolmgren(1911). Since C. formosanus was described first, C. formosaeshouldbe 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. formosanusbasedonthepublicationformatand languageused fordescription insteadof thenomen-clature rules and the content of the description. Oshima listed the originaldescriptionofC. formosae(Holmgren1911)inThe Third Official Report on Termites(Oshima1912)toshowthatHolmgren

only offered five measurements of alates, which was less informative than the first description of C. formosanus(Shiraki1909),andthatHolmgren failed to provide illustrations or photographs. After The Third Official Report on TermiteswaspublishedbyOshimain1912,most termite researchers such as Holmgren (1913) and Hozawa (1915)acceptedOshima’sopinionthatC. formosanus is theonlyCoptotermesspeciesinsouthernJapanandTaiwan,andC. formosaeisajuniorsynonymyof C. formosanus.

Coptotermes gestroi in TaiwanAfter1912,C. formosanuswasbelievedtobetheonlyCoptotermes

species in Taiwan. Coptotermes gestroiinTaiwanhadnotbeenmen-tioned until 2003. Based on soldier morphology, the new record of C. gestroi in Taiwan was proposed again (Tsai and Chen 2003). The first authorofthecurrentstudyandlocalpestcontroloperatorscollectedCoptotermes spp. from every county in Taiwan, both in urban areas and natural environments, from 2005–2009. With the mitochondrial gene sequence data, the presence of C. gestroiinsouthernTaiwanwas confirmed (Li et al. 2009). In total, 220 Coptotermessamplesincluding museum specimens were identified and used for mapping their distribution (Fig. 3B). Coptotermes spp. were mostly collected in lowland areas (< 500 m). Coptotermes formosanuswasdistributedthroughouttheislandofTaiwan,whileC. gestroiwasonlycollectedin the southern tropical zone.

Inareviewofarticlespublishedintheearly1900s,someevidenceshowsthatC. gestroi may have already been present in Taiwan. Os-hima(1909,1911)collectedCoptotermes samplesfrom12locationsin Taiwan (Fig. 3A), seven of which were in the tropical zone where C. gestroiandC. formosanus currently coexist. The drawing of a dark brown dealate (Fig. 2A) by Oshima (1909) resembled C. gestroi (Fig. 4A)insteadofthelightbrownish-yellowdealateofC. formosanus (Fig. 4C). The color of soldier heads of C. gestroi (Fig. 4B) and C. formo-sanus (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 coloroftheC. gestroi alate abdomen (Fig. 4A). Based on the color difference between soldier and alate, Oshima’s drawing (Fig. 2A and 2B)waslikelybasedonC. gestroi. In addition, the alate body length was recorded as ≈6 mm (Oshima 1909), which is closer to C. gestroithanC. 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.

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similartoC. gestroi (Fig. 5C), and its head length was recorded as 1.5 mm. Three years after his publication (Oshima 1909) in 1912, Oshima collectedCoptotermes samplesonlyinnorthernTaiwan(Taipeicity)(Fig. 3A) and in Japan, where only C. formosanus is currently found. The image of a soldier head capsule (Fig. 5B, Oshima 1912) was elongatedandsimilartoC. formosanus (Fig. 5D). The head length of soldiers reported by Oshima in 1912 was 1.66–1.72 mm. Soldiers collectedin1909and1911mightincludebothC. formosanusandC. gestroi;hence,theaverageoftheirheadlengthwasshorterthanthatofC. formosanus collected only from Taipei in 1912.

Oshima (1912) emphasized twice there was only one Coptotermessp. in Taiwan. However, no effort was made to prove that all the Cop-totermessamplesinTaiwanwerethesamespecies(Oshima1909,1910b, 1911). Basedonthisfalseassumption,Coptotermes samplescollectedatmanyplacesinTaiwanwerelumpedtogether,andthemeanofmorphologicalmeasurementsofthetwospeciesmayhavebeenpresentedasasinglespecies,C. formosanus. When Holmgren and Wasmann identified Japanese/Taiwanese Coptotermes samplesasC. formosaeandC. 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.

Type Specimen of C. formosanusDuring our investigation into the controversy on the identification

ofC. formosanusandC. gestroiintheearly1900s,museumspecimenspreservedinsixmajorinsectcollectionsinTaiwanwereexaminedtosearchforthoseusedbyShirakiandOshima:

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, InsectCollection,Taichung,Taiwan,ROC(NCHU)

6. National Pingtung University of Science and Technology, Depart-ment of Plant Medicine, Insect Collection, Neipu, Pingtung, Taiwan, ROC (NPUST)

A set of specimens (Fig. 6A) labeled “Coptotermes formosanus Shiraki” was found in TARI, where Shiraki worked between 1907 and 1942. The autograph on the label (Fig. 6B) is very similar to Shiraki’s handwriting inanunpublishedmanuscriptentitled 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 asC. 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,41styear 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(1883-1915),Shiraki’sassistant,onApril23rd,41styearofMeiji era (1908) from Pingtung county, Gangkou research station of TFRI (Fig. 3A). A comparison of these three alates with the origi-nal description (Shiraki 1909) and our recent collections (Li et al. 2009) has confirmed them to be C. formosanus. Even though these C. formosanusspecimenspreservedinTARIoncebelongedtoShirakiandmighthavebeenusedfordescribingC. formosanus in1909,thecollector 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, nopartofOshima’scollectionwasfoundinthesixinsectcollections

Fig. 6. Shiraki’s C. formosanus collection preserved in the Taiwan Agri-cultural 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 manu-script entitled Insect Fauna of Taiwan (C); soldiers (D) preserved in vials with two setae (inset) on one side of the fontanelle; the most well-pre-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 on April 23rd, 41st year of Meiji era (1908) at Gangkou. Photos of labels and insects not to scale.

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 Na-tional Taiwan University Li-brary); C. formosanus col-lected 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.

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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: 24.99°N, 121.30°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 sol-dierscollectedfromthesamecolonyastheneotypewillbedepositedin NMNS, NTU, TFRI, and University of Florida Termite Collection, Fort Lauderdale Research and Education Center. Mitochondrial genes includingCOII,12SrRNA,and16SrRNAofworkerscollectedfromthe same colony with the neotype have been partially sequenced andsubmittedtotheGenBankdatabasewithaccessionnumbers:EU805758, EU805712, and EU805735 (Li et al. 2009).

ThisreviewofthehistoricalcontroversyisintendedtopresentthechallengeforCoptotermestaxonomy,andtopromptinternationalcooperationonarevisionofCoptotermes. The genus Coptotermesisinseriousneedofrevision,especiallytheChinesespecies(Crosland1995, Ruelle 1996, Eggleton 1999). In the 21stcentury,theinherentdifficulties in Coptotermestaxonomyremain,butnewtoolssuchasstatistical methods and molecular techniques present new possibili-tiesforsolvingtheproblem(Kirton2005,VargoandHusseneder2009). Unprecedented levels of international communication, includ-ing sharing gene sequences through GenBank and releasing mor-phological images through Morphbank, ease information exchange. InternationalentomologistsmustworktogetherjustasHolmgren,Oshima, Shiraki, Wasmann, and Yano did a hundred years ago. With-outarevisionofCoptotermes, 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

AcknowledgementsThe authors thank Jing-Fu Tsai (National Chung-Hsing Univer-

sity) 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 review-ing 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. WealsothankShinyaMiyano(EntomologicalSocietyofJapan)andMeng-ChunHsieh(NationalTaiwanUniversityLibrary)forprocess-ingthecopyrightpermissionofOshima’sphotographandillustration,respectively. This study was supported in part by a grant from USDA-ARS under the grant agreement No. 58-6435-2-276.

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Hou-Feng LiandNan-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 gen-eral termite biology and control of subterranean termite. 3205 College Ave, Davie, FL33314. houfeng@ufl.edu; nysu@ufl.edu. Wen-Jer Wu is a Professor of Entomology, National Taiwan University. wuwj@ntu.edu.tw.