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SYSTEMATICS
Dimocarpomyia, a New Oriental Genus of the Tribe Asphondyliini
(Diptera: Cecidomyiidae) Inducing Leaf Galls on
Longan (Sapindaceae)
MAKOTO TOKUDA,
1,2
JUNICHI YUKAWA,
3
AND WIWAT SUASA-ARD
4
Ann. Entomol. Soc. Am. 101(2): 301Ð306 (2008)
ABSTRACT A new genus Dimocarpomyia belonging to the subtribe Asphondyliina (Diptera: Ce-
cidomyiidae: Asphondyliini) is erected for Dimocarpomyia folicola Tokuda & Yukawa, a new species
to science. Morphological features of the male, female, full-grown larva, and pupa are described. The
gall midge induces obovoid galls on the lower surface of leaves of longan, Dimocarpus longan
(Sapindaceae), in Thailand.
KEY WORDS Asphondyliina, Dimocarpomyia folicola,Dimocarpus longan, gall midge, Thailand
Various gall midges (Diptera: Cecidomyiidae) are
known to infest tropical fruit trees belonging to such
genera as Annona, Artocarpus, Citrus, Coffea, Ficus, and
Mangifera (Barnes 1948). In particular, mango, Man-
gifera indica L. (Anacardiaceae), is attacked by at least
17 gall midge species in tropical Asia (Prasad 1969;
Gagne´2004a,b), and some of them are recognized as
serious pests (Prasad 1971, Harris and Schreiner 1992,
Srivastava 1997, Uechi et al. 2002).
Longan, Dimocarpus longan Lour. (⫽Euphoria lon-
gan Steud., Euphoria longana Lam., and Nephelium
longana Cambess.) (Sapindaceae), is one of the im-
portant tropical fruit trees cultivated widely in South-
east Asia (e.g., Hotta et al. 1989). Up to the present,
taxonomy of gall midges associated with longan has
not been studied, even though gall midges inducing
galls with various shapes were often recognized as
pests in longan cultivations. Recently, we collected a
cecidomyiid inducing leaf galls on longan in Thailand
(Fig. 1), and we examined morphological features of
the adults, full-grown larvae, and pupae. As a result, we
conclude that the species belongs to the tribe Asphon-
dyliini and is new to science.
Asphondyliini exhibit various unique ecological and
life history traits, such as host alternation (Harris 1975;
Orphanides 1975; Yukawa et al. 2003; Uechi et al. 2004,
2005), polyphagy (Gagne´and Woods 1988, Tokuda et
al. 2005), extended diapause (Maeda et al. 1982,
Takasu and Yukawa 1984, Tabuchi and Amano 2003),
gall dimorphism (Tokuda and Yukawa 2005, Mishima
and Yukawa 2007), and association with fungal sym-
bionts (Meyer 1987, Bissett and Borkent 1988, Yukawa
and Rohfritsch 2005). For these reasons, ecological
and phylogenetic studies of Asphondyliini are of in-
terest in elucidating the evolution of these traits, and
in particular the processes of host range expansion,
host plant shift, and shift of gall-bearing organ (Toku-
da and Yukawa 2005, 2007). In addition, many species
belonging to this tribe have been recognized as serious
pests of various crops (Highland 1964, Tewari and
Moorthy 1986, Tewari et al. 1987, Dora and Kamala
1988, Gagne´and Orphanides 1992, Kumawat and Kan-
wat 1995, Yukawa et al. 2003). So, taxonomic and
applied studies of Asphondyliini are of fundamental
importance to establish control measures against these
pest species.
Because the gall midge obtained from leaf galls on
longan in Thailand has several remarkable character-
istics that have not been observed in known genera of
Asphondyliini, a new genus is erected for the species,
and we describe its morphological features.
Materials and Methods
Collection of Galls and Gall Midges. Leaf galls on
longan were collected in 2004 from orchards in
Chiangmai, northern Thailand. Some of the galls were
dissected under a stereoscopic microscope to obtain
larval and pupal specimens. The rest of the collected
galls were maintained in plastic bags (350 by 250 mm)
to obtain adults and pupal exuviae. All specimens
collected in this study were preserved in 75% ethanol
or in 99.5% acetone. In addition, we examined ethanol-
preserved specimens of the gall midge that had been
collected by C. Unahawutti in 1983 and 1985. All spec-
imens used in this study are kept in the collection of
1
Laboratory of Pest Management Systems, National Agricultural
Center for Kyushu Okinawa Region, Kumamoto 861-1192, Japan.
2
Corresponding author: Insect Interaction Research Unit, National
Institute of Agrobiological Sciences, Tsukuba, Ibaraki 305-8634, Japan
(e-mail: tokudam@affrc.go.jp).
3
Entomological Laboratory, Faculty of Agriculture, Kyushu Uni-
versity, Fukuoka 812-8581, Japan.
4
National Biological Control Research Center, Central Regional
Center, P.O. Box 1, Kasetsart University Kamphaengsaen Campus,
Nakhon-Pathom 73140, Thailand.
0013-8746/08/0301Ð0306$04.00/0 䉷2008 Entomological Society of America
the Entomological Laboratory, Faculty of Agriculture,
Kyushu University, Japan.
Morphological Studies and Terminology. Some of
the ethanol-preserved specimens were mounted on
slides for microscopic study in Canada balsam by using
the techniques outlined in Gagne´(1989). Drawings
were made with the aid of a drawing tube. Some
important structures of pupae were observed with a
scanning electron microscope (S-3000N, Hitachi, To-
kyo, Japan) by using the specimens stored in acetone.
Adult morphological terminology, except for that of
thoracic plates, follows McAlpine (1981) and that of
thoracic plates follows use in Tokuda (2004) and Toku-
da et al. (2004b). Recognition of female abdominal
segments and intersegmental membranes follows So-
linas and Isidoro (1991). Counts of setae and scales are
based on their insertions, in addition to actual setae
and scales, because many setae and scales become lost
through the processes of collection, preservation, or
preparation. Morphological terminology of immature
stages follows use in Mo¨hn (1955, 1961) and Yukawa
(1971), but the terminology of pupal antennal horn
follows that in Gagne´(1994). In addition, the term
“anterior dorsal papillae”(Tokuda et al. 2004a,b) is
applied to pupal abdominal papillae that are situated
anteriorly on the dorsal surface and distinctly anterior to
the row of “dorsal papillae.”The taxonomic investigation
in this article is the responsibility of M.T. and J.Y.
Description
Genus Dimocarpomyia Tokuda and Yukawa, gen. n.
Type species: Dimocarpomyia folicola sp. n.
Male. Palpus four-segmented (Fig. 2); antenna with
12 ßagellomeres; Þrst and second ßagellomeres not
Fig. 1. Leaf galls induced on longan by Dimocarpomyia
folicola sp. n.
Figs. 2–7. Dimocarpomyia folicola sp. n. (2) Male palpus, (3) male Þfth ßagellomere, (4) male Þrst tarsomere of mid leg,
(5) male genitalia in dorsal view, (6) ovipositor in lateral view, and (7) larval sternal spatula and associated papillae.
302 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 2
fused; ßagellomeres gradually shortened distally; cir-
cumÞla not closely appressed to the ßagellomeres
(Fig. 3). First tarsomeres each with minute apical spur
(Fig. 4); claws simple on all legs. Genitalia (Fig. 5):
gonostylus distally with sclerotized and bidentate
tooth; gonocoxite massive, extending ventrally well
beyond insertion of gonostylus; paramere absent; ae-
deagus basally wide, distally tapering.
Female. Flagellomeres shortened toward end of
ßagellum, the terminal one subglobular. Ovipositor
(Figs. 6 and 8): dorsal cerci-like structure bilobed,
situated on distal end of intersegmental membrane
between eighth and ninth abdominal segments; dis-
tal fourth needle-like and slightly bent upward;
cerci tiny.
Full-Grown Larva. Cervical papillae each bearing
seta. Sternal spatula elongate, distally with two-
pointed lobes (Fig. 7). Anus situated dorsally.
Pupa. Pupal skin not pigmented except for apical
horns. Spiracles of abdomen subglobular.
Remarks. The presence of a solid tooth on the
gonostylus and the dorsally situated larval anus in-
dicate that Dimocarpomyia belongs to the subtribe
Asphondyliina. This genus can be easily distin-
guished from other genera of Asphondyliina by the
unique structure of the female ovipositor. In other
Asphondyliina, the bilobed cerci-like structure on
female ovipositor is situated at the most proximal
portion of the intersegmental membrane between
eighth and ninth abdominal segments (Fig. 9); how-
ever, in the new genus, it is situated at the distal end
of this intersegmental membrane (Fig. 8). In the
resting female, almost all of this intersegmental
membrane and the ninth abdominal segment lies
retracted beneath the sixth to eighth abdominal
segments. In addition to the ovipositoral structure,
the genus also possesses other unique characteris-
tics (see Discussion).
Dimocarpomyia folicola Tokuda & Yukawa sp. n.
Male. Eye bridge Þve to six facets long at vertex;
frontoclypeal setal count as in Table 1. First palpal
segment subglobular, 25
m long, 1.4 times as long as
basal width; second 1.5 times as long as Þrst; third as
long as second; fourth 1.2 times as long as third. Scape
and pedicel with sparse setae; Þrst ßagellomere 140
m long (including short distal stem), 3.2 times as long
as wide, 1.1 times as long as second; Þfth ßagellomere
120
m long, 3.0 times as long as wide.
Thoracic setal and scale counts as in Table 1. Tarsal
claws bent at right angle on all legs; empodia slightly
shorter than claws; pulvilli ca. one-Þfth length of
claws. Wing 1.8Ð2.1 mm long, densely covered with
dark grayish hairs; R
5
joining costa at wing apex.
Abdominal tergites with anterior pair of trichoid
sensilla; Þrst through seventh tergites rectangular,
with single row of posterior setae; lateral setae absent;
anterior portion of eighth tergite sclerotized, without
setae. Abdominal sternites covered with setae and
scales, without anterior pair of trichoid sensilla. Gen-
italia (Fig. 5): cerci triangular, apically setose, sepa-
rated by deep V-shaped emargination; hypoproct shal-
lowly incised, each lobe usually with short apical seta;
gonostylus longer than wide, at apex with pointed
tooth on outer edge, and broad rounded Þnely den-
ticulate lobe on inner side.
Female. Second palpal segment 1.2 times as long as
Þrst; third 1.3 times as long as second. First ßagellomere
155
m long, 4.5 times as long as wide, 1.4 times as long
as second; Þfth ßagellomere 95
m long, 3.2 times as long
as wide. Wing 2.1 mm long. Seventh abdominal sternite
480
m long, ovipositor rather short, tapered, length of
the needle-like portion 200
m (Fig. 8).
Full-Grown Larva. Body length 2.0Ð2.2 mm. Body
color yellow. Four dorsal papillae present on thoracic
segments, each bearing seta; two dorsal papillae present
on Þrst to seventh abdominal segments, each bearing
seta; dorsal papillae on eighth abdominal segment not
apparent; two pleural papillae present on each side of
thoracic and Þrst through seventh abdominal segments,
each bearing seta; a pair of pleural papillae visible on
eighth abdominal segment; terminal papillae not appar-
ent. Sternal spatula 180Ð200
m long (Fig. 7). Generally
two inner and two outer lateral papillae present on each
thoracic segment, each bearing seta; sternal papillae each
bearing seta; inner pleural papillae not apparent; anterior
Figs. 8–9. Ovipositors of (8) Dimocarpomyia folicola sp.
n. and (9) Asphondylia baca (one of the specimens used in
the redescription of A.baca by Yukawa 1971) showing po-
sition of cerci-like structure (Cls) on dorsal surface.
Table 1. Dimocarpomyia folicola sp. n.: frontoclypeal and
thoracic setal and scale counts
Setae/scales Male (n⫽8)
Female
(n⫽1)
Count
Mean SD Range
Frontoclypeal setae 8.3 1.91 6Ð10 9
ADL setae
a
16.4 3.34 12Ð22 22
PDL setae
b
17.4 4.14 14Ð25 24
Mesopleural scales 22.6 4.69 16Ð30 27
Mesepimeral setae 13.8 2.12 10Ð17 14
a
Anterior dorsolateral setae.
b
Posterior dorsolateral setae.
March 2008 TOKUDA ET AL.: Dimocarpomyia,NEW ORIENTAL GENUS OF ASPHONDYLIINI 303
ventral papillae without seta; posterior ventral papillae
not apparent; anal papillae not apparent. Each abdominal
segment, except terminal one, ventrally with several
transverse rows of minute spines, dorsally with many
triangular spines.
Pupa. Body length 2.1Ð2.2 mm including apical
horns. Apical horn short, triangular (Fig. 10); apical
papilla situated on a small protuberance and bearing
seta; upper and lower frontal horns absent; a pair of
lower facial papillae without setae; lateral facial pa-
pillae not apparent; prothoracic spiracle 150Ð200
m;
spiracles present on each of Þrst eight abdominal seg-
ments, these abdominal segments ventrally with dense
minute spines; second to eighth abdominal segments
dorsally with 15Ð25 transverse rows of short spines,
each spine 10
m long. Four dorsal papillae present on
Þrst seven abdominal segments, usually each bearing
seta; a pair of anterior dorsal papillae present, each
bearing minute seta; two pleural papillae visible each
side, each bearing minute seta.
Host Plant. D. longan (Sapindaceae).
Gall. Obovoid swelling mostly on the lower surface
of host leaf, 3.0 mm in diameter (Fig. 1); single cham-
bered; surface brownish yellow; apically with a hole
through which the adult emerges.
Biological Notes. The life history of this species has
not yet been intensively studied. In Chiangmai, full-
grown larvae and pupae were found in galls on 13 and
14 January 2004. Adults emerged from the galls be-
tween 18 and 23 January 2004.
Holotype. Male (on slide, Cecid. No. TY0101; Type
no. 3248, kept in the Entomological Laboratory, Ky-
ushu University, Fukuoka, Japan). THAILAND: Galls
were collected on 13-I-2004 by M. Tokuda and W.
Suasa-ard from Tambol Mae, Faek Mai, San Sai Dis-
trict, Chiangmai (18⬚50⬘N, 99⬚02⬘E). The holotype
emerged on 19-I-2004.
Paratypes. Seven males and one female (on slides,
Cecid. Nos. TY0102Ð0109), same data as holotype ex-
cept for the emergence date (18 Ð23-I-2004); six pupae
(on slides, Cecid. Nos. TY0110Ð0115), same data as
holotype; four full-grown larvae (on slides, Cecid.
Nos. 0116Ð0119), THAILAND: Galls collected and
dissected on 14-I-2004 by M. Tokuda and W. Suasa-ard
from Maejo University orchard, Tambol Nong Han,
San Sai District, Chiangmai (18⬚51⬘N, 99⬚03⬘E)
(Cecid. No. TY0116Ð19).
Other Specimens Examined. Two males, eight fe-
males (on slides) and many adults (in ethanol), 1-I-
1983, C. Unahawutti, THAILAND: Chiangmai, four
full-grown larvae (on slides), many adults, pupae, and
full-grown larvae (in ethanol), 1985, C. Unahawutti,
THAILAND: Chiangmai.
Discussion
The tribe Asphondyliini is a well-circumscribed
monophyletic group sharing unique characteristics of
the adult postabdomen (Gagne´1994), and the tribe is
divided into two subtribes, Asphondyliina and Schizo-
myiina (Gagne´2004a). At present, two characteristics,
the solid tooth or teeth of the gonostylus and the
dorsally situated larval anus, are regarded as synapo-
morphies of the subtribe Asphondyliina (Tokuda
2004; Tokuda and Yukawa 2005, 2006, 2007), whereas
Schizomyiina are considered to be a paraphyletic
group of Asphondyliina because of the lack of syna-
pomorphies (Gagne´2004a; Tokuda 2004; Tokuda et al.
2004a, 2005). This relationship was also supported by
phylogenetic analysis for eastern Palaearctic and Ori-
ental Asphondyliini by using 14 morphological char-
acters (Tokuda and Yukawa 2007).
The genus Dimocarpomyia is regarded as a member
of Asphondyliina, based on the two synapomorphies,
but it has several remarkable features. Male genitalia
of Dimocarpomyia exhibit more or less intermediate
characteristics between Asphondyliina and Schizo-
myiina: the ventrally well-developed gonocoxite ex-
tending beyond the insertion of gonostylus is a com-
mon feature of Schizomyiina (Mo¨hn 1961, Tokuda et
al. 2004a, Tokuda et al. 2005), and it is only known in
Daphnephila, an Oriental and eastern Palaearctic ge-
nus, in the Asphondyliina (Yukawa 1974, Tokuda and
Yukawa 2007); the bidentate claw on the gonostylus
with pointed tooth on outer edge and rounded lobe on
inner side is a unique feature of the genus; and the
shape of the aedeagus and absence of parameres are
typically exhibited by Asphondyliina. The ovipositor
of Dimocarpomyia is unusual in the subtribe Asphon-
dyliina. In other genera of Asphondyliina, the bilobed
cerci-like structure on the female ovipositor is situated
at the most proximal portion of the intersegmental
membrane between eighth and ninth abdominal seg-
ments (Fig. 9); however, in the new genus, it is situ-
ated at the distal end of the intersegmental membrane
(Fig. 8). Moreover, cercal lobes that are situated at the
apex of the ovipositor are relatively large in Dimocar-
pomyia (Figs. 6 and 8) compared with other genera of
Asphondyliina, in which they are usually very tiny and
indistinct.
Degeneration of papillae is known in the full-grown
larvae of Asphondyliina that pupate in galls (Mo¨hn
1961, Tokuda et al. 2004a). In Dimocarpomyia, degen-
eration of various larval papillae, such as the posterior
Fig. 10. Pupal head of Dimocarpomyia folicola sp. n.
304 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 101, no. 2
ventral, terminal, and anal papillae, are observed. Full-
grown larvae of some Old World genera of Asphon-
dyliina such as Pseudasphondylia and Illiciomyia are
known to have asetose papillae on the ventral surface
of the thoracic segments (Yukawa 1971; Tokuda and
Yukawa 2002, 2005; Tokuda 2004). This feature is com-
mon to Schizomyiina and regarded as plesiomorphic.
Dimocarpomyia possesses setose papillae alone on the
ventral surface as seen in Daphnephila, Bruggman-
niella, and all genera of Neotropical and Nearctic
Asphondyliina (Mo¨hn 1961, Yukawa 1974, Gagne´
1994, Tokuda and Yukawa 2006).
The pupa of Dimocarpomyia seems to retain most of
the plesiomorphic characters in the Asphondyliina.
Usually, Asphondyliina have well-developed pupal
apical horns (Mo¨hn 1961), but Dimocarpomyia has
very small horns. The pupal skin is not pigmented in
Dimocarpomyia, which is also known in some Palae-
arctic genera such as Daphnephila, Pseudasphondylia,
Illiciomyia, and a worldwide genus Bruggmanniella
(Yukawa 1974; Tokuda and Yukawa 2002, 2005, 2006;
Gagne´et al. 2004; Tokuda 2004). In some genera of
Asphondyliina, such as Illiciomyia, Pseudasphondylia,
and Bruggmanniella, pupal abdominal spiracles on an-
terior segments are elongate and those on posterior
segments are degenerated (Yukawa 1971; Tokuda and
Yukawa 2002, 2005, 2006; Tokuda 2004). However,
such elongation and degeneration of abdominal
spiracles were not observed in Dimocarpomyia, Daph-
nephila, and Asphondylia.
Even though some remarkable genera of Asphon-
dyliini, e.g., Luzonomyia, Daphnephila, and Dimo-
carpomyia, occur in the Oriental Region, faunistic
studies of Asphondyliini are still insufÞcient in this
area (Tokuda and Yukawa 2007). According to un-
published information (M.T. and J.Y., unpublished
data in 2004), several gall midge species seem to be
associated with longan in Thailand. Further studies
of the gall midge fauna on longan will provide useful
information on systematics and phylogeny of As-
phondyliini. Ecological and life history studies also
are needed to develop control measures against gall
midges infesting longan.
Acknowledgments
We are grateful to M. Takagi, B. Napompeth, S. Kamitani,
D. Yamaguchi, N. Uechi, K. Akimoto, A. Matsunaga, and T.
Ganaha for help in collecting materials. Our thanks are also
extended to O. Tadauchi for allowing use the scanning elec-
tron microscope and to C. Unahawutti for offering specimens
of D.folicola and its information. M.T. thanks N. Minagawa,
H. Kanno, and other members of Laboratory of Pest Man-
agement Systems, National Agricultural Research Center for
Kyushu Okinawa Region, Japan, for support. We thank R. J.
Gagne´for critical reading of an early draft. This study was
partly supported by Grant-in-Aid 15208007 (to J.Y. through
team leader M. Takagi) from the Japanese Society for Pro-
motion of Sciences (JSPS) and the JSPS Research Fellowship
for Young Scientists (to M.T.).
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