Solving the
Hundred-Year Controversy
of Coptotermes Taxonomy
in Taiwan
Hou-Feng Li, Nan-Yao Su, and Wen-Jer Wu
T
he 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
222
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 1912. 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 1945. 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 (1882-1970) (Fig. 1A) and Masamitsu
Oshima (1884-1965) (Fig. 1B) started to work in Taiwan at the
American Entomologist • Winter 2010
Fig. 1. (A) Tokuichi Shiraki
(1882-1970) (Photo provided by the Entomology
Dept. of the National Taiwan
University); (B) Masamitsu
Oshima (1884-1965)
(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 1898. 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 (1883-1970) of Forestry
Experimental Station, Tokyo, Shozaburo Watasé (1862-1929) and
his student, Sanji Hozawa (1885-1947) 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 (1859-1931), for identification.
Wasmann believed that these samples were C. gestroi, which he
named in 1896. 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
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 (18771954), through Chiyomatsu Ishikawa (1861-1935) of Tokyo Imperial
University and another German termite researcher, Karl Escherich
(1871-1951). 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 (1.66–1.72 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 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 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 2003. 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 2005–2009. 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
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 1.66–1.72 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 1912.
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
1942. 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 (1883-1915), 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
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 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 EU805735 (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 21st 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. This study was supported in part by a grant from USDAARS under the grant agreement No. 58-6435-2-276.
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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. 3205 College Ave,
Davie, FL33314. [email protected]; [email protected]. Wen-Jer Wu is a Professor
of Entomology, National Taiwan University. [email protected].
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