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SYSTEMATICS
First Records of Genus Bruggmanniella
(Diptera: Cecidomyiidae: Asphondyliini) from Palaearctic and
Oriental Regions, with Descriptions of Two New Species
That Induce Stem Galls on Lauraceae in Japan
MAKOTO TOKUDA
1, 2
AND JUNICHI YUKAWA
3, 4
Ann. Entomol. Soc. Am. 99(4): 629Ð637 (2006)
ABSTRACT Two new species of the genus Bruggmanniella,Bruggmanniella actinodaphnes and
Bruggmanniella cinnamomi (Diptera: Cecidomyiidae: Asphondyliini: Asphondyliina) are described
from Japan. The former species induces stem galls on Actinodaphne lancifolia (Siebold et Zucc.)
(Lauraceae) in the Palaearctic Region, and the latter species induces stem galls on Cinnamomum
japonicum (Lauraceae) in the Oriental Region. Bruggmanniella, previously known to occur only in the
Neotropical and southern Nearctic regions, is recorded for the Þrst time from the Palaearctic and
Oriental regions. The number of genera, their components, and morphological features of Asphon-
dyliina are compared between different zoogeographical regions.
KEY WORDS Asphondyliini, Bruggmanniella, Cecidomyiidae, gall midge, Lauraceae
THE TRIBE ASPHONDYLIINI IS a well circumscribed mono-
phyletic group sharing unique characteristics of the
adult postabdomen and is divided into two subtribes,
Asphondyliina and Schizomyiina (Gagne´1994, 2004).
The tribe is a taxonomically and phylogenetically
well studied group compared with other tribes of
Cecidomyiidae (Mo¨hn 1961, Gagne´1994), but many
species are still unnamed in various parts of the world.
In addition, Asphondyliini exhibit various unique eco-
logical 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), prolonged diapause (Maeda
et al. 1982, Takasu and Yukawa 1984, Tabuchi and
Amano 2003), and association with fungal symbionts
(Meyer 1987, Bissett and Borkent 1988, Yukawa and
Rohfritsch 2005). Therefore, further taxonomic, phy-
logenetic, and ecological studies of Asphondyliini are
required to elucidate the evolution of these traits, in
particular the processes of host range expansion, in-
cluding host plant shift and galled organ shift (Tokuda
and Yukawa 2005).
Many species of the subtribe Asphondyliina have
been known to induce various galls on broad-leaved
evergreen trees of Lauraceae in the Oriental and
eastern Palaearctic regions, such as India (Kieffer
1905), southwestern Japan (through which the
boundary of the Palaearctic and Oriental regions ex-
tends; Fig. 1) (Yukawa 1974, Yukawa and Masuda
1996), Taiwan (Yang et al. 1999), and Indonesia and
Thailand (Yukawa et al. 2005). Accordingly, we have
been paying special attention to Lauraceae trees in our
search for galling species of Asphondyliina at various
localities in Japan. In recent Þeld surveys, we have
collected several galls of Asphondyliina on Lauraceae
from Japan. In this article, we describe two new spe-
cies of the genus Bruggmanniella that induce stem galls
on trees of Lauraceae. We compare generic compo-
nents and morphological features of Asphondyliina,
including Bruggmanniella, between the Neotropical
and Palaearctic regions.
Materials and Methods
Collection of Galls and Gall Midges. Some of the
collected galls were dissected under a stereoscopic
microscope to obtain larval and pupal specimens.
When some of the dissected galls contained mature
larvae or pupae, 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 99.5%
acetone.
Preservation of Specimens Examined. All speci-
mens examined, including holotypes and paratypes,
1
Entomological Laboratory, Graduate School of Bioresources and
Bioenvironmental Sciences, Kyushu University, Fukuoka 812-8581,
Japan.
2
Corresponding author, address: Institute for Biological Resources
and Functions, National Institute of Advanced Industrial Science and
Technology, Tsukuba, Ibaraki 305-8566, Japan (e-mail: tokuda-
m@aist.go.jp).
3
Entomological Laboratory, Faculty of Agriculture, Kyushu Uni-
versity, Fukuoka 812-8581, Japan.
4
Current address: Matsuzaki 1-5-12, Higashi-ku, Fukuoka 813-0035,
Japan.
0013-8746/06/0629Ð0637$04.00/0 䉷2006 Entomological Society of America
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are preserved in the collection of the Entomological
Laboratory, Kyushu University, Fukuoka, Japan.
Morphological Studies and Terminology. For mi-
croscopic study, some of the ethanol-preserved spec-
imens were mounted on slides in Canada balsam by
using the techniques outlined in Gagne´(1989). Draw-
ings were made with the aid of a drawing tube. Some
important structures of the pupae were examined with
a scanning electron microscope (S-3000N, Hitachi,
Tokyo, Japan) by using the acetone-preserved spec-
imens.
Adult morphological terminology, except for that of
thoracic plates, follows use in McAlpine (1981) and
that of thoracic plates follows use in Tokuda (2004)
and Tokuda et al. (2004b). Counts of setae and scales
were based on the setal and scale insertions because
many setae and scales become lost through the pro-
cesses of collection, preservation, and preparation.
Morphological terminology of the immature stages
follows use in Mo¨hn (1955, 1961), which was orig-
inally written in German and later translated into
English in Yukawa (1971), but terminology of the
pupal antennal horn follows that in Gagne´(1994).
In addition, the term “anterior dorsal papillae”(To-
kuda et al. 2004a,b) is applied to pupal abdominal
papillae that are situated in the anterior fourth of
the dorsal surface and distinctly anterior to the row
of “dorsal papillae.”
Taxonomy
Genus Bruggmanniella Tavares
Bruggmanniella Tavares, 1909: 19. Type species: Brugg-
manniella braziliensis Tavares 1909
Hemibruggmanniella Mo¨hn, 1961: 6. Type species
Bruggmanniella oblita Tavares 1920
The genus Bruggmanniella belongs to the subtribe
Asphondyliina and comprises six known species in the
world (Gagne´2004, Gagne´et al. 2004). Among them,
Þve species occur in the Neotropical Region and one
in the southern parts of the Nearctic Region. Because
the morphological features of Bruggmanniella were
summarized and discussed in Gagne´(1994) and
Gagne´et al. (2004), we refer in this article only to
morphological differences between Bruggmanniella
and allied genera in the Palaearctic Region as fol-
lows: Bruggmanniella is similar to Pseudasphondylia
Monzen, 1955 in most male morphological features
such as the presence of parameres and the two sep-
arate teeth of the gonostylus. However, Bruggman-
niella can be distinguished from Pseudasphondylia by
the presence of setae on the larval cervical papillae
and on all papillae of the ventral surface (Tokuda and
Yukawa 2005). Bruggmanniella, with its two separate
teeth on the gonostylus, is distinguishable from Pro-
bruggmanniella Mo¨hn, 1961, which has a single tooth.
It differs from Illiciomyia Tokuda, 2004 by having
several morphological features such as less constricted
male ßagellomeres, convolute male circumÞla, pres-
ence of an apical spur on the Þrst tarsomeres, and
absence of asetose lateral papillae on larval thoracic
segments (Tokuda 2004).
Bruggmanniella actinodaphnes
Tokuda & Yukawa sp. n.
(Figs. 2A and B, 3A and B; Tables 1 and 2)
Male. Eye bridge four to six facets long. Frontocly-
peal setal count as in Table 1. Palpus two-segmented;
Þrst palpal segment ⬇38
m long, ⬇1.3 times as long
as basal width; second ⬇2.0 times as long as Þrst.
Antenna with 12 ßagellomeres; scape and pedicel with
rather dense setae; Þrst and second ßagellomeres not
fused; Þrst ßagellomere ⬇215
m long, 4.3 times as
long as wide, 1.2 times as long as second, Þfth ßag-
ellomere ⬇160
m long, 3.3 times as long as wide.
Thoracic setal counts as in Table 1. All legs covered
with many blackish scales; length of respective seg-
ments as in Table 2; Þrst tarsomeres of all legs each
with a short apical spur; claw simple on all legs, bent
nearly at right angle; empodium well developed, as
long as claw; pulvillus much shorter than empodium.
Fig. 1. Collection records of B.actinodaphnes and
B.cinnamomi based on Yukawa (1976), Usuba (1977), Yama-
uchi et al. (1982), Yukawa (1988), Yukawa and Masuda
(1996), and the present results. The broken line illustrates
the boundary between Oriental and Palaearctic zoogeo-
graphical regions.
Table 1. B. actinodaphnes sp. n.: frontoclypeal and thoracic
setal counts (nⴝ7 for each sex)
Male Female
Mean SD Range Mean SD Range
Frontoclypeal setae 25.0 2.94 21Ð28 21.3 4.72 18Ð28
ADL setae
a
73.0 19.76 61Ð108 69.6 5.13 65Ð76
PDL setae
b
96.2 12.03 83Ð112 91.8 17.63 81Ð118
Mesopleural setae 56.0 8.00 45Ð65 55.0 12.37 46Ð76
Mesepimeral setae 57.0 4.98 50Ð63 60.4 7.16 52Ð71
a
Anterior dorsolateral setae.
b
Posterior dorsolateral setae.
630 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 99, no. 4
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Wing length 2.8Ð3.4 mm, ⬇2.2 times as wide, densely
clothed in dark grayish hairs; R
5
joining costa a little
beyond wing apex.
Tergites without anterior pair of trichoid sensilla;
Þrst through seventh tergites rectangular; Þrst through
sixth tergites with double row of posterior setae,
seventh tergite with mostly triple row of posterior
setae; anterior portion of eighth tergite pigmented,
without setae; some lateral setae present on Þrst to
seventh tergites; elsewhere covered with scales on
Þrst through seventh tergites. Sternites without ante-
rior pair of trichoid sensilla; Þrst through sixth sternites
rectangular, 0.25Ð0.34 times as long as wide; seventh
and eighth sternites a little elongated, 0.44 and 0.57
times as long as wide, respectively. Genitalia (Fig. 2A):
cerci setose, each usually rounded at apex, rarely com-
pletely fused; hypoproct incised deeply by V-shaped
emargination, each lobe with two apical setae; gono-
stylus suboval, distally with two sclerotized teeth, the
teeth 8.5
m apart from one another; gonocoxite
rather elongate, ventrally extending a little beyond
insertion of gonostylus; paramere relatively large, usu-
ally with distinct seta; aedeagus laterally sclerotized,
distally tapering.
Female. Distal ßagellomeres shortened and termi-
nal one subglobular; Þfth ßagellomere 150
m long,
⬇3.0 times as long as wide. Wing length ⬇3.65 mm,
⬇2.2 times as long as wide. Seventh sternite ⬇1.2 times
as long as wide; eighth sternite without setae. Ovipos-
itor protractile, slender, aciculate, basally with a bi-
lobed cerci-like structure. Needle part of ovipositor
⬇1.05 mm long, ⬇1.6 times as long as the length of
seventh sternite; minute apical lobe of ovipositor with-
out setae. Otherwise as in male.
Mature Larva. Body color pale green, length 2.6Ð
3.1 mm. Second antennal segment short, conical,
13
m long, 1.3 times as long as basal width; cervical
papillae each with seta. Number and position of spi-
racles normal; four dorsal papillae present on all tho-
racic and Þrst through seventh abdominal segments,
each with seta; eighth abdominal segment with two
dorsal papillae, each with seta; two pleural papillae
present on each side, each with seta; two terminal
papillae present, each with minute seta. Sternal spat-
ula (Fig. 2B) 190Ð230
m long, anteriorly with two
triangular lobes; two inner and two outer lateral pa-
pillae present on all thoracic segments, each with seta;
sternal papillae each with seta on all thoracic seg-
ments; inner pleural papillae probably absent; anterior
and posterior ventral papillae not apparent; anal pa-
pillae not apparent. Each abdominal segment, except
terminal one, ventrally with many transverse rows of
minute spines and with many small triangular spines
ventro- and dorsolaterally.
Pupa. Pupal skin not pigmented except for apical
horn. Body length ⬇3.6 mm. Apical horn broad, dor-
soventrally ßattened, with three Þngerlike projec-
tions, the mesalmost longest, lateralmost shortest,
length from the base of antennal sheath to the tip of
apical horn 300Ð370
m (Fig. 3A); apical papillae
without setae; upper and lower frontal horns absent;
usually a pair of lower facial papillae and two pairs of
lateral facial papillae present, each without setae; pro-
thoracic horn 150Ð180
m long; stigmatal tubercles
present on second to sixth abdominal segments, each
35Ð50
m long; those on seventh and eighth abdominal
segments rudimentary; Þrst abdominal segment
densely covered with minute spines; second to eighth
abdominal segments densely covered with minute
spines on ventral surface and on middle third of dorsal
surface; second to eighth abdominal segments with
Þve to seven transverse rows of rather long spines on
Table 2. B. actinodaphnes sp. n.: measurements of legs (micrometers, nⴝ7 for each sex)
Male Female
Mean SD Range Mean SD Range
Foreleg
Femur 1,272 53.8 1,197Ð1,344 1,118 91.2 985Ð1,241
Tibia 1,154 94.1 1,083Ð1,348 1,016 113.5 837Ð1,154
Tarsomere I 174 13.7 154Ð192 178 30.4 141Ð210
Tarsomere II 970 157.8 763Ð1123 889 156.8 641Ð1,053
Tarsomere III 441 25.6 415Ð475 366 46.4 314Ð410
Tarsomere IV 320 40.6 281Ð375 249 22.9 224Ð269
Tarsomere V 207 12.8 194Ð224 200 39.9 158Ð254
Mid-leg
Femur 1,212 131.3 941Ð1324 1,163 114.9 961Ð1,269
Tibia 1,018 75.2 904Ð1109 956 104.7 773Ð1,063
Tarsomere I 172 18.6 151Ð206 178 17.0 148Ð193
Tarsomere II 933 78.4 885Ð1071 719 91.7 609Ð883
Tarsomere III 445 44.5 410Ð522 331 28.8 303Ð381
Tarsomere IV 312 30.3 280Ð359 269 39.9 210Ð323
Tarsomere V 208 20.8 183Ð227 172 18.1 149Ð196
Hindleg
Femur 1,279 54.5 1,203Ð1,350 1,247 65.3 1,173Ð1,362
Tibia 1,063 89.3 931Ð1198 1,018 118.5 904Ð1,184
Tarsomere I 184 18.4 167Ð222 173 24.8 143Ð212
Tarsomere II 935 163.4 721Ð1118 713 66.7 652Ð831
Tarsomere III 446 9.7 437Ð458 334 27.7 297Ð368
Tarsomere IV 332 23.5 303Ð354 255 28.5 207Ð280
Tarsomere V 224 18.5 198Ð242 183 37.4 150Ð256
July 2006 TOKUDA AND YUKAWA:Bruggmanniella,NEW TO PALAEARCTIC/ORIENTAL REGIONS 631
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Fig. 3. (A) Pupal head and frontal area of B.actinodaphnes sp. n. (B) Pupal abdominal segments (dorsal view) of B.
actinodaphnes. (C) Pupal head and frontal area of B.cinnamomi sp. n. (D) Pupal abdominal segments (dorsal view) of B.
cinnamomi.
Fig. 2. (A) Male genitalia (dorsal view) of B.actinodaphnes sp. n. (B) Larval sternal spatula of B.actinodaphnes. (C) Male
genitalia (dorsal view) of B.cinnamomi. (D) Larval sternal spatula of B.cinnamomi. Scale bar ⫽0.1 mm.
632 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 99, no. 4
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anterior third of dorsal surface (Fig. 3B), each spine
8Ð20
m long; six dorsal papillae present on Þrst to
seventh abdominal segments, outermost pair each
with seta, remaining four usually without setae, rarely
each with minute seta; two anterior dorsal papillae
present on second to eighth abdominal segments, each
with a seta; a pleural papilla present on each side,
usually each with two minute setae, rarely with one
minute seta.
Host Plant. Actinodaphne lancifolia (Sieb. and
Zucc.) Meissn. (Lauraceae).
Gall. Subconical swelling on the stem, rounded api-
cally; usually many galls occurring together; mono-
thalamous (Fig. 4A; see also Gall No. C-250 in Yukawa
and Masuda 1996).
Biological Notes. The life history of this species has
not yet been intensively studied. It seems to be uni-
voltine, but some individuals may require 2 yr for one
generation (Yukawa and Masuda 1996). Stem galls
collected in December contained mature larvae and
most adults emerged from the galls on the host plant
between early April and early May in the following
year.
Holotype. Male (on slide, Cecid. No. B4001; Type
No. 3228, kept in the Entomological Laboratory,
Kyushu University, Fukuoka, Japan). JAPAN: Galls
were collected on 27-IV-1984 by J. Yukawa from Shi-
royama, Kagoshima, Kyushu. The holotype emerged
on 4-V-1984.
Paratypes. JAPAN: 5 么么,6乆乆, and 6 pupae (on
slides, Cecid. Nos. B4002Ð12, 405Ð56) same data as
holotype; 1 么,1乆, and 1 pupa (on slides, Cecid. Nos.
B4013Ð14, B4057), Iso, Kagoshima, Kyushu, galls col-
lected 1-V-1969, A. Mori, adults emerged 1-3-V-1969,
reared by J. Yukawa; two larvae (on slides, Cecid. Nos.
B4076Ð77), Ishiki, Kagoshima, Kyushu, 13-XII-1969,
J. Yukawa; 10 larvae (on slides, Cecid. Nos. B4078Ð
87), Kamiyama, Tokushima, Shikoku, 22-X-2001, M.
Yukinari.
Other Material Examined. JAPAN: 2 么么,3乆乆, and
8 pupae (in ethanol), same data as holotype; many
larvae (in ethanol), Ishiki, Kagoshima, Kyushu, 13-
XII-1969, J. Yukawa; 1 larva (in ethanol) and 1 larva (in
acetone), Mount Mukabaki, Miyazaki, Kyushu, 18-XI-
2000, M. Tokuda & J. Yukawa; 3 乆乆 and 2 pupae (in
ethanol), Izuhara, Tsushima Island, Nagasaki, galls col-
lected 27-IV-2001, M. Shoubu; one pupa (in acetone),
Izuhara, Tsushima Island, Nagasaki, galls collected 28-
IV-2001, M. Shoubu, adults emerged 5-10-V-2001,
reared by M. Tokuda; 4 larvae (in ethanol) and 4 larvae
(in acetone), Kamiyama, Tokushima, Shikoku, 22-XI-
2001, M. Yukinari.
Distribution. This gall midge was previously col-
lected from Honshu and Kyushu, Japan (Fig. 1)
(Yukawa 1976, Usuba 1977, Yukawa 1988) and is re-
corded for the Þrst time from Shikoku, Japan, in this
article. It has not yet been found in the southwest
islands of Japan.
Remarks. This species is distinguishable from the
other congeners in the Neotropical and Nearctic re-
gions by the two-pointed anterior lobes of the larval
sternal spatula (Fig. 2B), whereas the spatula has
three- or four-pointed lobes in the Neotropical and
Nearctic species (Gagne´1994, 2004). The pupal apical
horn (Fig. 3A) is also unique by having three apical
Þngerlike projections.
A congener, which is morphologically very similar
to B. actinodaphnes was collected several times in
Japan so far (Yukawa 1976; Usuba 1977; Yukawa 1979,
1988; unpublished data) but has been left unnamed
because of the inadequate number of specimens for
description. It induces stem galls (Yukawa and
Masuda 1996) on Actinodaphne acuminata (Bl.)
Meissn. (⫽A. longifolia).
Bruggmanniella cinnamomi
Tokuda & Yukawa sp. n.
(Figs. 2C and D, 3C and D; Tables 3 and 4)
Male. Eye bridge six to eight facets long. Fronto-
clypeal setal count as in Table 3. Palpus three-seg-
mented; Þrst palpal segment ⬇45
m, ⬇1.4 times as
long as basal width; second ⬇1.4 times as long as Þrst;
third ⬇1.1 times as long as second. First ßagellomere
200Ð250
m long, ⬇4.5 times as long as wide, ⬇1.2
times as long as second, Þfth ßagellomere 160 Ð185
m,
⬇3.4 times as long as wide. Wing length ⬇2.6 mm,
⬇2.3 times as long as wide. First through sixth sternites
0.38Ð0.48 times as long as wide, seventh sternite
⬇0.63 times as long as wide, and that of eighth sternites
Fig. 4. (A) Stem galls caused by B.actinodaphnes on A.lancifolia. (B) Stem galls caused by B.cinnamomi on
C.japonicum.
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⬇0.48 times as long as wide, respectively. Genitalia
(Fig. 2C): cerci setose, each usually triangular at apex,
rarely completely fused; hypoproct incised very shal-
lowly by U-shaped emargination, each lobe with one
or two apical setae. Otherwise as in B. actinodaphnes.
Female. First ßagellomere 210
m long, 5.6 times as
long as wide, 1.3 times as long as the second; Þfth
ßagellomere 140
m long, 3.7 times as long as wide.
Wing length ⬇3.1 mm, 2.4 times as long as wide.
Seventh sternite ⬇1.4 times as long as wide. Needle
part of ovipositor 0.88 mm, ⬇1.7 times as long as the
length of seventh sternite. Otherwise as in the female
of B. actinodaphnes.
Mature Larva. Body color pale yellow. Body length
2.7Ð3.5 mm. Second antennal segment 11
m long,
1.8 times as long as basal width. Sternal spatula
(Fig. 2D) 290Ð320
m long, anteriorly with two tri-
angular lobes, additional sclerotized portion present
laterally to each triangular lobe. Otherwise as in
B. actinodaphnes.
Pupa. Head and thoracic parts of pupal skin slightly
pigmented. Body length 3.5Ð4.3 mm. Apical horn dor-
soventrally ßattened, anterior margin narrowed, pos-
terolaterally with small lobe, outer margin of the small
lobe Þnely denticulate, length from the base of an-
tennal sheath to the tip of apical horn 410Ð470
m
(Fig. 3C); apical papilla with 80
m long seta; upper
and lower frontal horns absent; lower and lateral facial
papillae not visible; prothoracic horn 270Ð290
m
long; stigmatal tubercles on second to sixth abdominal
segments 25Ð32
m long; Þrst abdominal segment
densely covered with minute spines on ventral and
dorsal surface; second to eighth abdominal segments
densely covered with minute spines on ventral surface
and on posterior half of dorsal surface; second to
eighth abdominal segments with nine to 11 transverse
rows of rather short spines on anterior one-half of
dorsal surface (Fig. 3D), each spine 12Ð23
m long;
usually eight dorsal papillae present on Þrst to seventh
abdominal segments, most outer and second inner
pairs each with minute seta, sometimes most inner and
second inner papilla more or less fused; anterior dorsal
papillae absent; each segment with one pleural papilla,
with a seta. Otherwise as in B. actinodaphnnes.
Host Plant. Cinnamomum japonicum Sieb. ex Nees
(Lauraceae).
Gall. Massive subglobular or elongated subglobular
swelling on the stem, diameter 10Ð40 mm, containing
many small larval chambers (Fig. 4B; see also Gall No.
C-264 in Yukawa and Masuda 1996).
Biological Notes. The life history of this species is
still unclear although this species is suspected to be
univoltine (Yamauchi et al. 1982, Yukawa and Masuda
1996). Stem galls that were collected in December had
already developed to full size but contained Þrst in-
stars. Adults emerged from the galls between mid-
March and early April the following year.
Holotype. Male (on slide; Cecid. No. C7101; Type
No. 3229, kept in the Entomological Laboratory,
Table 3. B. cinnamomi sp. n.: frontoclypeal and thoracic setal
counts (nⴝ6 males and 5 females)
Male Female
Mean SD Range Mean SD Range
Frontoclypeal setae 18.7 3.08 16Ð24 16.3 4.35 12Ð20
ADL setae
a
37.2 4.07 30Ð41 43.8 1.71 42Ð46
PDL setae
b
34.7 4.46 30Ð40 38.2 3.77 34Ð44
Mesopleural setae 33.0 2.83 31Ð35 32.7 4.16 28Ð36
Mesepimeral setae 36.2 4.92 30Ð42 29.7 2.08 28Ð32
a
Anterior dorsolateral setae.
b
Posterior dorsolateral setae.
Table 4. B. cinnamomi sp. n.: measurements of legs (micrometers, nⴝ6 males and 5 females)
Male Female
Mean SD Range Mean SD Range
Foreleg
Femur 1,071 64.9 986Ð1,134 1,136 110.1 1,014Ð1,312
Tibia 1,100 56.3 1,010Ð1,169 1,226 51.9 1,158Ð1,291
Tarsomere I 135 19.8 104Ð157 157 19.8 125Ð176
Tarsomere II 905 58.3 824Ð962 934 35.6 895Ð972
Tarsomere III 516 20.3 487Ð531 494 30.0 466Ð524
Tarsomere IV 376 23.8 344Ð397 338 11.6 322Ð349
Tarsomere V 211 12.2 200Ð230 227 15.9 209Ð243
Mid-leg
Femur 958 40.7 890Ð1020 993 74.4 899Ð1088
Tibia 920 6.3 911Ð928 955 43.7 908Ð1,002
Tarsomere I 127 18.7 107Ð157 160 19.4 134Ð188
Tarsomere II 669 61.8 584Ð771 611 40.3 560Ð649
Tarsomere III 433 16.0 407Ð453 400 38.0 364Ð465
Tarsomere IV 307 24.5 273Ð336 276 8.9 265Ð284
Tarsomere V 206 21.9 173Ð233 208 14.7 192Ð224
Hindleg
Femur 1,132 104.5 989Ð1,295 1,157 162.5 906Ð1,359
Tibia 1,053 52.7 978Ð1,106 1,071 105.7 893Ð1,177
Tarsomere I 127 22.5 105Ð162 147 28.7 99Ð176
Tarsomere II 731 69.6 641Ð830 693 60.8 620Ð754
Tarsomere III 486 31.9 458Ð532 436 85.2 380Ð586
Tarsomere IV 373 15.4 358Ð394 315 40.4 264Ð356
Tarsomere V 239 22.4 207Ð269 214 16.0 192Ð235
634 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 99, no. 4
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Kyushu University, Fukuoka, Japan). JAPAN: Naka-
gusuku, Okinawa, galls collected on 25-II-2001 by
J. Yukawa and S. Yamauchi. The holotype emerged
15-III-2001, reared by M. Tokuda.
Paratypes. JAPAN: 5 么么,5乆乆, eight pupae, and
three larvae (on slides, Cecid. No. C7102Ð11, C7151Ð
58, C7176Ð78), same data as holotype except emer-
gence dates (between 15- and 21-III-2001); 3 么and 6
pupae (on slides, Cecid. No. C7112Ð14, C7159Ð64),
Aha, Kunigami, Okinawa, 5-III-2002, M. Tokuda and
H. Kuratomi, adults emerged between 10- and 19-III-
2002.
Other Material Examined. JAPAN: many adults and
pupae (in acetone or ethanol), same data as holotype;
many adults and pupae (in acetone or ethanol), Aha,
Kunigami, Okinawa, galls collected 5-III-2002,
M. Tokuda & H. Kuratomi, adults emerged 10Ð19-III-
2002, reared by M. Tokuda.
Distribution. The southwest islands, Japan (from
Kikaijima Island to Ishigakijima Island, included in the
Oriental Region; Fig. 1) (Yukawa and Masuda 1996).
Galls caused by this gall midge are commonly seen on
Okinawa Island and have been collected once in 1978
from Mount Omoto, Ishigakijima Island (Yamauchi et
al. 1982). Current and old galls could not be found in
March 2000 and March 2001 on Ishigakijima and
Iriomotejima islands, although at least 50 large trees
and many saplings were surveyed on respective is-
lands. The galls have never been collected in Honshu,
Shikoku, and Kyushu, Japan, which are included in the
Palaearctic Region (Yukawa and Masuda 1996), even
though the host plant is naturally distributed in these
areas (Horikawa 1972).
Remarks. This species differs from the other con-
geners by the unique pupal apical horn (Fig. 3C) and
shape of the larval sternal spatula (Fig. 2D). On the
southwest islands of Japan, similar stem galls have
been found on Cinnamomum doederleinii Engeler
and Machilus thunbergii Sieb. and Zucc. (Lauraceae)
(Yamauchi et al. 1982; Yukawa and Masuda 1996).
However, the taxonomic position of the gall midges
that are responsible for these stem galls was not de-
termined because we had no adult and pupal speci-
mens of the gall midges to examine.
Discussion
In the tribe Asphondyliini, two subtribes, Asphon-
dyliina and Schizomyiina, have been clearly divided
based on the Neotropical and western Palaearctic taxa
by many synapomorphies (Mo¨hn 1961, Gagne´1994).
However, recent taxonomic studies of the tribe in the
eastern Palaearctic Region have demonstrated that
some genera could not be clearly classiÞed into either
Asphondyliina or Schizomyiina in the light of the
previous subtribal concept (Tokuda 2004, Tokuda et
al. 2004a). At present, only two characteristics, the
solid tooth or teeth of the gonostylus and the dorsally
situated larval anus, are regarded as synapomorphies
of the subtribe Asphondyliina (Tokuda 2004),
whereas Schizomyiina seems to be a paraphyletic
group of Asphondyliina, because no synapomorphies
have been designated for the subtribe (Gagne´2004,
Tokuda 2004, Tokuda et al. 2004a).
The genus Bruggmanniella was previously recorded
from the Neotropical and Nearctic regions, but this
article reveals that it occurs also in the Palaearctic and
Oriental regions. As a result, eight genera of Asphon-
dyliina have been recorded from the Palaearctic Re-
gion, compared with ten genera in the Neotropical
Region (Gagne´2004, Tokuda 2004). In contrast to the
Neotropical and Palaearctic regions, the Nearctic Re-
gion is a genus-poor area, where only two genera of
Asphondyliina, Asphondylia H. Loew, 1850 and Brugg-
manniella, occur (Gagne´1989, 2004). Asphondyliina
have been relatively well-studied taxonomically in the
Nearctic (Gagne´1989), Neotropical (Mo¨hn 1961,
Gagne´1994), and Palaearctic regions (Mo¨hn 1961;
Kovalev 1964; Yukawa 1971, 1974; Tokuda 2004; To-
kuda and Yukawa 2005), but only a few species have
been described from the other zoogeographical re-
gions: for example, Coutin (1980) and Kolesik (1995)
in the Australian; Kieffer (1913) in the Afrotropical;
and Kieffer (1905), Gagne´(1973), Yukawa (1981),
and Uechi and Yukawa (2004) in the Oriental regions.
So, further studies are needed to discuss generic com-
ponents of Asphondyliina in these regions.
Although there is no big difference in the numbers
of genera between the Neotropical (10 genera) and
Palaearctic regions (eight genera), the generic com-
ponents of Asphondyliina are much different between
the two regions. In the Neotropical region, all genera,
except for Bruggmanniella, are considered to be phy-
logenetically rather close to Asphondylia, which is the
largest, cosmopolitan, and radiating genus of Asphon-
dyliina (Gagne´2004, Yukawa et al. 2005), because
these genera share all or some synapomorphies, such
as, e.g., pigmented pupal skin, presence of upper and
lower frontal horns, absence of parameres on gono-
coxites (Mo¨hn 1961, Gagne´1994). In contrast to the
Neotropical genera, all Palaearctic genera, except for
Asphondylia and Houardiella Kieffer, 1912, do not pos-
sess these synapomorphies. Indeed, Palaearctic genera
have some morphological features that are common to
those of Schizomyiina and not found in Nearctic As-
phondyliina other than Bruggmanniella, such as
stubby ovipositors in Daphnephila Kieffer, 1905, shal-
low construction of male ßagellomeres and absence of
apical spur on the Þrst tarsomeres in Illiciomyia, and
absence of setae on larval cervical papillae and on
some papillae on the ventral surface in Pseudasphon-
dylia (Yukawa 1971, 1974; Tokuda and Yukawa 2002,
2005; Tokuda 2004). Therefore, these Palaearctic gen-
era probably diverged earlier than the Nearctic genera
from a common ancestor of Asphondylia and its related
genera. Such differences in generic components of
Asphondyliina possibly imply that the common an-
cestor of the subtribe Asphondyliina originated in the
Old World and that of the genus Asphondylia in the
Neotropical Region.
In the Neotropical and Nearctic regions, Bruggman-
niella is associated with various plant families such as
Anacardiaceae, Celastraceae, Malpighiaceae, Mora-
ceae, and Sapotaceae (Gagne´2004). In these regions,
July 2006 TOKUDA AND YUKAWA:Bruggmanniella,NEW TO PALAEARCTIC/ORIENTAL REGIONS 635
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Bruggmanniella perseae Gagne´, 2004 is the only species
that is associated with Lauraceae, inducing fruit galls
on avocado, Persea americana Mill. (Gagne´et al. 2004).
Morphologically, B. perseae seems closer to the other
Neotropical congeners than to the Palaearctic and
Oriental species that induce stem galls on Lauraceae.
For example, the larval sternal spatula has four ante-
rior lobes in B. perseae and some congeners in the
Neotropical Region (Gagne´et al. 2004), but only two
lobes in the Palaearctic species. In addition, the male
cerci are fused basally in B. perseae and Neotropical
congeners but usually discrete two lobes in the Palae-
arctic species. These differences indicate that Brugg-
manniella species possibly have become gallers on
Lauraceae independently in the Palaearctic and Neo-
tropical regions. However, there are still many unde-
scribed species associated with Machilus, which is an
allied genus of Persea, in the Palaearctic and Oriental
regions (Yukawa et al. 2005). Further morphological
and phylogenetic studies are needed to determine
relationships among the Neotropical and Palaearctic
species of Bruggmanniella. According to Chanberbali
et al. (2001), the “Persea group,”which includes
Machilus, Persea, and related genera, diverged from
Laureae including Actinodaphne and Cinnamomeae
including Cinnamomum in the Eocene (⬇44 MYA),
and then Laureae and Cinnamomeae diverged from
one another, also in the Eocene (⬇40 MYA). At
present, no information is available to assess the time
when the Bruggmanniella-Lauraceae association be-
gan. Molecular phylogenetic analysis among the
Palaearctic and Neotropical Bruggmanniella species
could possibly be used to assess the time of divergence
for the respective species, as has been used in the case
of Ilex-associated species of the genus Asteralobia
Kovalev, 1964 (Diptera: Cecidomyiidae: Asphondyli-
ini) (Tokuda et al. 2004b).
In the Palaearctic and Oriental regions, the genus
Daphnephila, belonging to the Asphondyliina, also is
associated with Lauraceae and produces leaf and stem
galls on Machilus (Yukawa 1974, Yang et al. 1999).
Although the “Persea group”sensu Chanderbali et al.
(2001) is distributed also in the Neotropical Region,
Daphnephila has never been reported there (Mo¨hn
1961; Gagne´1994, 2004). As mentioned by Yukawa et
al. (2005), this suggests that Daphnephila has be-
come associated with Machilus after the Asian Persea
group had diverged from the American group in the
EoceneÐOligocene boundary (⬇32 MYA; Chander-
bali et al. 2001). Moreover, Pseudasphondylia neolit-
seae Yukawa, 1974, which also belongs to the sub-
tribe Asphondyliina, is known to induce leaf galls on
Neolitsea sericea (Bl.) Koidz. (Lauraceae) in Japan
(Yukawa 1974, Tokuda and Yukawa 2005). That so
many genera and species of Asphondyliini are associ-
ated with Lauraceae in the Palaearctic and Oriental
regions and that only B. perseae is in the Neotropical
Region is remarkable.
The broad distribution of Bruggmanniella across
the southern Nearctic, Neotropical, eastern Palaearc-
tic, and Oriental regions possibly suggests that the
genus had formerly spread elsewhere in the world, as
in the present radiation of the genus Asphondylia
(Gagne´2004, Yukawa et al. 2005).
Acknowledgments
We express our thanks to R. J. Gagne´and K. M. Harris for
critical reading of a draft. We are grateful to H. Kuratomi,
A. Mori, S. Yamauchi, and M. Yukinari for offering gall midge
specimens. M.T. thanks O. Tadauchi, S. Kamitani, and
D. Yamaguchi for support. This study was partly supported by
the Research Fellowships of the Japanese Society for the
Promotion of Sciences for Young Scientists (to M.T.).
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Received 20 July 2005; accepted 6 December 2005.
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