Journal of Asia-Pacific Entomology 23 (2020) 364–370

Contents lists available at ScienceDirect

Journal of Asia-Pacific Entomology

journal homepage: www.elsevier.com/locate/jape

Full length article

New data on lance flies (Diptera, Lonchaeidae) associated with figs T

(Moraceae, Ficus spp.) in Japan and Taiwan, with descriptions of two new
species of the genus Silba Macquart
⁎
Kôichi Arimotoa, , Iain MacGowanb, Zhi-Hui Sua,c
a
JT Biohistory Research Hall, Murasaki Town 1-1, Takatsuki City, Osaka 569-1125, Japan
b
National Museums of Scotland, Collection Centre, 242 West Granton Rd, Edinburgh EH5 1JA, Scotland, United Kingdom
c
Department of Biological Sciences, Graduate School of Science, Osaka University, Machikaneyama Town 1-1, Toyonaka City, Osaka 560-0043, Japan

A R T I C LE I N FO A B S T R A C T

Keywords: Larvae of the genus Silba Macquart, 1875 were obtained from the syconia of three different species of Ficus
Diptera growing on the southern islands of Japan and Taiwan. Examination of the emerged adults identified four species,
Ficus two of which are described as new to science: Silba erecta MacGowan and Arimoto sp. nov. and Silba thunbergii
Fig wasps MacGowan and Arimoto sp. nov. Of the four species observed, the larvae of S. erecta, Silba ishigaki MacGowan
Lonchaeidae
and Okamoto, 2013 and S. thunbergii feed on the tissue of the fig syconia. Silba inubiwa MacGowan and Okamoto,
Predator
2013 was observed as a predator of three fig wasp species. The biogeographic distribution and biological data
Taxonomy
are provided for each of these fig-associated species.

Introduction
The association of species in the genus Silba Macquart, 1875 with fig
syconia has been known for over a century. Silvestri (1917) provided
the first account when he detailed the life history of Silba adipata
McAlpine, 1956 under the name Lonchaea aristella Becker, 1902 based
on misidentification. The larvae of this Mediterranean species live in
the flesh of Ficus carica L. syconia. More recently, five Silba species
associated with figs have been described from India (MacGowan et al.,
2012), Japan (MacGowan and Okamoto, 2013), Thailand, and Cam-
bodia (MacGowan and Compton, 2018). These Silba species are as-
signed to two groups: in the first, the larvae feed on the flesh of fig
syconia, and in the second, the larvae apparently prey on fig wasps
(Okamoto et al., 2012; MacGowan and Compton, 2018). Given the
approximately 850 species of fig, which comprise the most diverse
group in the family Moracea (Christenhusz et al., 2017), there must still
be many undescribed Silba species using this microhabitat. Further
study of the taxonomy, behaviour, and feeding habits of these Silba
species should lead to an increased understanding of the biological
diversity and interactions associated with fig–fig wasp mutualism, as
well as the evolution of larval feeding strategies within the genus Silba.
Therefore, we conducted field work to collect new materials and bio-
logical information on Silba species in Japan and Taiwan. This paper
describes two new fig-associated Silba species, the larvae of which feed
on syconium flesh. Data are also provided on the biology and dis-
tribution of two previously described species.
Materials and methods
Field work was conducted on Shikoku and the Ryukyu Islands,
Japan and in Taiwan during 2017 and 2018 to collect Silba larvae
closely associated with the fig–fig wasp mutualistic system (Fig. 1).
Specimens examined were collected by Kôichi Arimoto (KA), Yûji
Matsumoto (YM), Ayako Sasaki (AS), Zhi-Hui Su (ZHS), and members
of the laboratory of Taxonomy and Ecology of Forest Plants, National
Chung Hsing University (TEFP; Po-An Chou, Hiu-Wa Lee, Shin-Hung
Pan, Yu-Chun Yi, and Prof. Hsy-Yu Tzeng). Five to ten syconia in stages
ranging from C (wasp developing) to E (ripe syconia) or F (fallen sy-
conia) were examined per male fig tree [terminology after Galil and
Eisikovitch (1968) and Palmieri et al. (2018)]. When Silba larvae were
found in syconia, they were removed and transferred into plastic
rearing vials. These were kept at room temperature (ca. 25 °C) for about
2 weeks until adults emerged. Samples of the emerged adults were
preserved in 80% ethanol for morphological examination; the re-
mainder were stored in 99.5% ethanol for future molecular analyses.
Leaves of the trees examined were also kept as identification records.
The syconia from which larvae were removed were preserved in 99.5%
ethanol and leaves were preserved using the teabag method (Wilkie

⁎
Corresponding author.
E-mail addresses: antbeginning@gmail.com (K. Arimoto), imacgowan9@gmail.com (I. MacGowan), su.zhihui@brh.co.jp (Z.-H. Su).

https://doi.org/10.1016/j.aspen.2019.11.007
Received 30 March 2019; Received in revised form 13 November 2019; Accepted 15 November 2019
Available online 05 February 2020
1226-8615/ © 2020 Korean Society of Applied Entomology. Published by Elsevier B.V. All rights reserved.
K. Arimoto, et al.
Fig. 1. Map of the Ryukyu Islands, Japan and Taiwan, showing the collection
sites of the Silba species examined. Red circles: S. inubiwa; yellow triangle: S.
ishigaki; green rectangle: S. thunbergii; blue cross: S. erecta. (For interpretation of
the references to colour in this figure legend, the reader is referred to the web
version of this article.)
et al., 2013). The specimens of the syconia and leaves collected in this
study are deposited in the JT Biohistory Research Hall, Osaka, Japan
[BRH].
Collection dates are given as the day on which larvae were col-
lected. Voucher numbers of the syconia were added alphabetically after
the tree numbers when more than one syconium was collected from the
same tree (e.g. J17-707 K). Tree and syconia numbers are given in
rounded brackets.
Pinned and mounted adult Silba specimens were deposited in the
collections of the National Museum of Scotland, Edinburgh, UK [NMS],
National Museum of Nature and Science, Tokyo, Japan [NSMT], and
Taiwan Agricultural Research Institute, Taichung, Taiwan [TARI].
Specimens preserved in alcohol for future molecular analyses were
deposited in BRH. Depositories are given in square brackets. The
taxonomic nomenclature of flies follows MacGowan and Rotheray (in
press).
Systematics: New species
Genus Silba Macquart, 1851
Included species associated with fig-fig wasp mutualism. Eight
species: S. adipata McAlpine, 1956; S. erecta MacGowan & Arimoto sp.
nov.; S. inubiwa MacGowan and Okamoto, 2013; S. ischnopoda
MacGowan and Compton, 2018; S. ishigaki MacGowan and Okamoto,
2013; S. japonica MacGowan and Okamoto, 2013; S. lanshker Mac-
Gowan & Razak, 2012; S. thunbergii MacGowan & Arimoto sp. nov.
Remarks. Asian fig feeding Silba species such as S. erecta, S. isch-
nopda, S. ishigaki, S. japonica, S. lashker and S. thunbergii, are char-
acterized with much reduced aristal plumosity.
Silba erecta MacGowan and Arimoto sp. nov.
Etymology. The specific epithet refers to Ficus erecta from which
the type series was reared.
Type material. Holotype. Male, along Huandao Road, Lüdao
Township (Green Is.), Taitung County, Taiwan, 22.6626° N, 121.5022°
E, 104 m, 30 VIII 2018, KA, AS, ZHS & TEFP leg., ex. Ficus erecta
(TW18-125A) [TARI]. Paratype. 1 female with the same data as the
holotype [NMS].
Description. Male. Head. Eyes bare. Frons black, dulled by
365
Journal of Asia-Pacific Entomology 23 (2020) 364–370
microsculpture. Orbital plate shining black, bare apart from orbital
seta. Frontal and interfrontal setulae very short, no more than
0.1 × length of orbital seta, longer setulae on anterior margin above
lunule. Lunule black, slightly grey dusted as face and parafacials.
Anterior genal setulae forming single row of 6–7 along mouth margin.
Antennae black; postpedicel with distinct orange medial base, length to
width ratio 2.0:1; arista yellow at base, with short plumosity, at its
greatest extent 0.3 × width of postpedicel.
Thorax. Proepisternum and proepimeron each with 1 seta. Scutum
glittering black, covered with appressed setulae, which approximately
0.4 × length of orbital seta on head. Scutellum shining black on disc,
with 4 marginal setae, with 2 short setulae between lateral and apical
setae, with a single short setula between apical setae. Anepisternum
with 2 anterior and 3 posterior setae; setulae between these rows no
more than 0.5 × length of the setae. Katepisternum with 1 weak
anterior and 1 strong posterior setae near dorsal margin; anterior seta
half as long and strong as posterior seta; 2–3 setulae lying anterior to
these setae along the anterior margin. Wing length 3.0 mm; membrane
covered with long dark microtrichia; veins dark brown. Calypteres
white, with white fringe. Halteres black. Legs entirely black including
ventral row of spicules on fore and hind metatarsi.
Abdomen. Gentalia (Fig. 2). Epandrium higher than long, with semi-
circular excavation on anterior margin. Cerci relatively large, rectan-
gular in shape, bearing numerous short, stiff setulae along posterior
margin and apex. Surstyli not extending the full length of the epan-
drium; outer margin with a row of stiff setae (Fig. 2A). Internal view
(Fig. 2B), a small setula located at base of each cercus; prensisetae in
rather uniform single row of 5 running medially from posterior margin
of surstyli; two setulae at dorsal end of the row. Phallus small and
lightly chitinised; intromittent organ long, thin and curving basally;
pair of long antler-like processes each with two strong ventral projec-
tions and a serrated apex (Fig. 2C).
Female. Similar to male except for the usual sexual differences.
Aculeus with apical segment diamond-shaped, in dorsal view twice as
long as wide; basal third embedded in rods of aculeus; with 2 short
setulae on lateral side and long setula on ventral side at apical half.
Remarks. Silba erecta has only 1 strong seta on the katepisternum,
an unusual feature in the genus but shared by three Asian fig-feeding
species: S. ishigaki, S. lashker and S. ischnopda. In external features S.
erecta is most similar to Silba ishigaki, but the key feature which clearly
separates these species is in the shape of the phallus. The phallus of S.
ishigaki is long and thin, serrated for most of its length and with only a
single basal process (MacGowan and Okamoto, 2013, Fig. 7) whilst in S.
erecta it is far more complex (Fig. 2C).
Distribution (Fig. 1). TAIWAN: Taitung County (Green Is.).
Host plant. Ficus erecta Thunb. var. erecta.
Bionomics. Larvae of this species were found in the syconia of F.
erecta. The syconia contained empty fig wasp galls of the E phase, from
which the fig wasps had already emerged. As no fig wasp exoskeleton
fragments were found in the syconia, we believe that S. erecta larvae do
not prey on fig wasp larvae, but feed on syconium tissue. Adults
emerged on 18. IX. 2018.
Silba thunbergii MacGowan and Arimoto sp. nov.
Etymology. The specific epithet refers to Ficus thunebergii from
which the type series was reared.
Type material. Holotype. Male, along Prefectural Route 79,
Shinokawa, Setouchi Town (Amami-Ôshima Is.), Ôshima District,
Kagoshima Pref., Japan, 28.2231° N, 129.2943° E, 15 m, 13 IV 2018,
KA, YM, AS & ZHS leg., ex. Ficus thunebergii (J18-127D) [NSMT].
Paratypes. 1 male and 3 females with the same data as the holotype
[NMS].
Description. Male. Head. Eyes bare. Frons greyish black, dulled by
microsculpture. Orbital plate black, dulled by microsculpture, bare
apart from orbital seta. Frontal and interfrontal setulae very short, no
K. Arimoto, et al.
Fig. 2. Male terminalia of Silba erecta sp. nov. A: Epandrium and associated structures, right side, lateral view; B: surstylus, left side, internal view; C: phallus, right
side, lateral view.
more than 0.1 × length of orbital seta; longer setulae on anterior
margin above lunule. Lunule ground colour black, covered by silver-
grey pollinosity; face and parafacials lightly grey dusted. Anterior genal
setulae forming a single row of 5 along mouth margin; basal two setulae
longest and strongest on genae. Antennae entirely black; postpedicel
length to depth ratio 2.5:1; arista with short plumosity, at its greatest
extent 0.5 × width of postpedicel.
Thorax. Proepimeron and proepisternum each with 1 seta. Scutum
sub-shining black, covered with short appressed setulae, which no more
than 0.3 × length of orbital seta. Scutellum on margin with 4 short
setulae on right side and 5 short setulae on left side between lateral and
apical setae, with a pair of very short setulae between apical setae.
Anepisternum with a row of 3 anterior and 4 posterior setae, a scat-
tering of setulae between these rows no more than 0.5 × length of the
setae. Katepisternum with 2 setae near dorsal margin; a few setulae
lying anterior to these setae otherwise the sclerite almost bare. Wings
clear, veins dark brown, wing length 3.5 mm. Calypteres very slightly
greyish with a relatively dense fringe of white setulae. Halteres black.
366
Journal of Asia-Pacific Entomology 23 (2020) 364–370
Legs entirely black including ventral row of spicules on fore and hind
basal tarsomeres.
Abdomen. Genitalia (Fig. 3). In lateral view epandrium slightly
higher than long, anteriorly with a semi-circular excavation, numerous
long setulae located ventrally and posteriorly. Cerci rather small, rec-
tangular in shape, bearing numerous short, stiff setulae (Fig. 3A) In
ventral aspect large rounded surstyli project beyond shell of the epan-
drium, each with a marginal row of setulae. Medial margin of surstyli
with 5–6 lateral creases. Internal view, a pair of setulae present at base
of cerci (Fig. 3B); prensisetae in a single row of 6 near posterior margin
of surstyli, gradually becoming longer ventrally; the ventral 2 pre-
nsisetae long and thin, almost 3 × as long as the basal prensiseta.
Pregonites bifurcated, finger–like, with an apical seta. Postgonites large
and rectangular; medial margin with a row of 5–6 small setulae. Phallus
relatively small and lightly chitinised, with a pair of small, pointed
apical processes and basally a pair of large almost rectangular processes
(Fig. 3C); intromittent organ slender and slightly curved.
Female. Similar to male apart from the usual sexual differences.
K. Arimoto, et al.
Fig. 3. Male terminalia of Silba thunbergii sp. nov. A: Epandrium and associated structures, right side, lateral view; B: ditto, ventral view; C: phallus, right side, latral
view.
Aculeus with apical segment spear shaped, dorsally with a pair of basal
setulae 0.5 × length of apical segment, ventrally with a similar pair
setulae near apex; a further much shorter pair setulae located at mid-
point on ventral margin.
Remarks. In several features this species, with 2 setae on the ka-
tepisternum, is a more typical member of the genus Silba but differs in
the greatly reduced plumosity on the arista. The male terminalia are
also relatively typical of the genus. As is the case in other Silba species
the key specific character is the distinctive structure of the phallus
(Fig. 3C).
Distribution (Fig. 1). JAPAN: Ryukyu Islands (Amami-Ôshima Is.).
Host plant. Ficus thunbergii Maxim.
Bionomics. Eggs of this species were laid between the ostiolar
bracts of the syconia of Ficus thunbergii (Fig. 4A: arrows). On hatching,
larvae moved inside the syconia (Fig. 4B, C). It was not possible to
define the feeding habits of S. thunbergii larvae clearly, as two types of
larvae with different feeding habits were found in the same syconium
(Fig. 4C). Many larvae preyed on adult fig wasps inside their galls
(Fig. 4C: white arrow), whereas others fed on syconium tissue (Fig. 4C:
black arrow) in C to D phase. We believe that the larvae preying on the
367
Journal of Asia-Pacific Entomology 23 (2020) 364–370
fig wasps were those of S. inubiwa, whereas the larvae feeding on the
syconium tissue belonged to S. thunbergii (see also bionomics of S. in-
ubiwa).
Further data on previously described species associated with figs
Silba inubiwa MacGowan and Okamoto, 2013
Silba inubiwa MacGowan and Okamoto, 2013: 197 (original de-
scription; type locality: Kii-Ôshima Island, Wakayama Prefecture,
Honshu, Japan).
Material examined. Ex. Ficus erecta. JAPAN. Shikoku: 2 males,
Ôshima, Sukumo City, Kôchi Pref., 32.9157° N, 132.6986° E, 19 m, 7 IX
2017, KA leg. (J17-695E) [BRH]; 1 female, same place, 32.9155° N,
132.6971° E, 69 m, 7 IX 2017, KA leg. (J17-707 K) [NMS]. Ryukyu.
Kagoshima Pref.: Yakushima Is.: 2 female, Anbô, Yakushima Town,
Kumage District, 30.3164° N, 130.6353° E, 163 m, 19 VII 2018, KA leg.
(J18-614B) [1 female: NMS; 1 female: BRH]; Amami-Ôshima Is.: 1
female, Agina, Setouchi Town, Ôshima District, 28.1610° N, 129.3228°
E, 133 m, 13 IV 2018, KA, YM, AS & ZHS leg. (J18-120A) [NMS]; 1
K. Arimoto, et al.
Fig. 4. Silba spp. and Wiebeia fig wasps from the syconium of Ficus thunbergii, Amami-Ôshima Island, the Ryukyu Islands, Japan (J18-127B). A: Fly eggs between
ostiolar bracts of the syconium; B: hatched or unhatched fly eggs (arrows: unhatched egg); C: fly larvae feeding on fig wasp or syconium tissue (white arrow:
predation of an adult female fig wasp inside a fig gall; black arrow: feeding on syconium tissue); D: adult male fig wasps predated by fly larvae (arrows: abdomen
from which internal organs were lost).
female, Nishikomi, Setouchi Town, Ôshima District, 28.2382° N,
129.1604° E, 54 m, 13 IV 2018, same collectors as the former (J18-133)
[NSMT]; 2 males, Amami City, 28.3003° N, 129.4724° E, 159 m, 14 IV
2018, same collectors as the former (J18-151A) [NMS]. Okinawa
Pref.: Okinawa-jima Is.: 1 male, 1 female, near Mt. Terukubi,
Kunigami Village, Kunigami District, 26.7614° N, 128.2574° E, 358 m,
2 VII 2018, KA leg. (OK18-1E) [NMS]; Tokashiki-jima Is.: 2 females,
Tokashiki, Tokashiki Village, Shimajiri District, 26.2044° N, 127.3549°
E, 169 m, 5 V 2018, KA leg. (J18-441I) [1 female: NMS; 1 female: BRH];
Kume-jima Is.: 1 male, Zenda, Kume-jima Town, Shimajiri District,
26.3206° N, 126.8039° E, 46 m, 25 IV 2018, KA leg. (J18-216C) [NMS].
TAIWAN. Taoyuan City: 3 males, 4 females, along Northern Cross-
County Highway, Hualing Village, Fuxing District, 24.6515° N,
121.4177° E, 1,289 m, 27 VIII 2018, KA, AS, ZHS & TEFP leg. (TW18-
29A) [2 males, 2 females: NMS; 1 male, 2 female: BRH]; 1 male, 3
females, same tree as the former (TW18-29B) [1 male: NMS; 3 females:
BRH]; 2 males, same tree as the former (TW18-29C) [1 male: NMS; 1
male: BRH]; 2 males, 2 females, same tree as the former (TW18-29D) [2
males: NMS; 2 females: BRH]; 3 females, same tree as the former
(TW18-29F) [1 female: NMS; 2 females: BRH]; 1 male, along Northern
Cross-County Highway, Shuanyuan, Fuxing District, 24.6446° N,
121.4442° E, 1,094 m, 27 VIII 2018, KA, AS, ZHS & TEFP leg. (TW18-
32A) [1 male: NMS;]; 4 male, 3 females, same tree as the former
(TW18-32D) [1 male, 2 females: NMS; 3 males, 1 female: BRH]; 1 male,
4 females, along Northern Cross-County Highway, Shuanyuan, Fuxing
District, 24.6448° N, 121.4443° E, 1,101 m, 27 VIII 2018, KA, AS, ZHS
& TEFP leg. (TW18-33A) [1 male, 1 female: NMS; 3 females: BRH].
Yilan County: 7 males, 9 females, along Northern Cross-County
Highway, Sidauban, Datong Township, 24.6309° N, 121.4826° E,
1,224 m, 27 VIII 2018, KA, AS, ZHS & TEFP leg. (TW18-39A) [1 male, 3
females: NMS; 2 males, 6 females: BRH]; 2 males, 2 females, same data
as the former, (TW18-40A) [1 male, 1 female: NMS; 1 male, 1 female:
BRH]; 1 female, same tree as the former (TW18-40B) [BRH]; 4 males, 3
females, same tree as the former, (TW18-40C) [2 males: NMS; 2 males,
368
Journal of Asia-Pacific Entomology 23 (2020) 364–370
3 females: BRH]. Chiayi County: 2 males, 1 female, along Dadungshan
Walkway, Meishan Township, 23.5022° N, 120.7137° E, 1,678 m, 4 IX
2018, KA, AS, ZHS & TEFP leg. (TW18-277A) [1 male, 1 female: NMS; 1
male: BRH]. Ex. Ficus thunbergii. JAPAN. Kagoshima Pref.:
Yakushima Is.: 2 males, 3 females, Miyanoura, Yakushima Town,
Kumage District, 30.4127° N, 130.5659° E, 30 m, 20 VII 2018, KA leg.
(J17-647 J) [2 males, 2 females: NMS; 1 female: BRH].
Distribution. JAPAN: Honshu (Wakayama Pref.), Shikoku (Kôchi
Pref.), Ryukyu Islands (Yakushima Is., Amami-Ôshima Is., Okinawa-
jima Is., Tokashiki-jima Is., Kume-jima Is., Ishigaki-jima Is.). TAIWAN:
Taoyuan City, Yilan County, Chiayi County.
New records from the all islands, except for Honshu and Ishigaki-
jima Island (Fig. 1). This species is widely distributed from mainland
Japan to Taiwan and from 30 to 1700 m in altitude.
Host plant. Ficus erecta Thunb. var. erecta (MacGowan and
Okamoto, 2013) and Ficus thunbergii Maxim. (new record).
Bionomics. Females were often observed flying around, alighting
on, and walking on syconia of F. erecta (Fig. 5A). On finding the en-
trance to the syconia (ostiole) in the C phase (Fig. 5B), oviposition took
place in the ostiolar bracts (Fig. 5C). On hatching, the larvae moved to
the interior of the syconia (Fig. 5D: black arrows). The oviposition
behaviours observed were generally similar to those noted for Silba
adipata (Katsoyannos, 1983). It took approximately 10 days for adults
to emerge after pupation.
The larvae of S. inubiwa prey on fig wasps (Okamoto et al., 2012;
MacGowan and Okamoto, 2013), as do those of Silba ischnopoda
MacGowan and Compton, 2018 (MacGowan and Compton, 2018).
Okamoto et al. (2012) reported that larvae of Silba sp. (=Silba inubiwa)
fed on adult female Blastophaga nipponica Grandi, 1922 on syconia of F.
erecta. The Silba larvae in the syconia of F. thunbergii were observed
preying on both adult female and adult male fig wasps (Fig. 4C: white
arrow). The larvae fed mainly on the internal organs within the abdo-
mens of male wasps, entering from the ventral aspect (Fig. 4D: arrows).
The fig wasp species associated with F. erecta in Taiwan was
K. Arimoto, et al.
Fig. 6. Puparium of Silba ishigaki MacGowan and Okamoto, 2013 (TW18-262A). A: Normal empty puparium from which an adult fly emerged; B: empty puparium
due to predation by the larva of a staphylinid beetle (arrow: hole opened by the larva to enter the puparium).
previously thought to be B. nipponica (Chen and Chou, 1997). However,
subsequent analysis of sequence data suggested that the Taiwanese
population of B. nipponica and two other fig wasp species (Blastophaga
taiwanensis Chen and Chou, 1997 and Blastophaga tannoensis Chen and
Chou, 1997) represent a single panmictic population and are distinct
from the Japanese population of B. nipponica (Wachi et al., 2016). These
Taiwanese species have not yet been revised taxonomically. The Tai-
wanese fig wasps from F. erecta must differ from B. nipponica and be
assigned to B. taiwanensis based on the nomenclature. Otherwise, the fig
wasp species associated with F. thunbergii is considered to be an un-
described species of Wiebesia Bouček, 1988 (Yokoyama and Iwatsuki,
1998). In conclusion, the larvae of S. inubiwa are currently believed to
prey on three fig wasp species: B. nipponica, B. taiwanensis, and Wiebesia
sp. (also see the bionomics of Silba thunbergii).
Silba ishigaki MacGowan and Okamoto, 2013
Silba ishigaki MacGowan and Okamoto, 2013: 198 (original de-
scription; type locality: Ishigaki-jima Island, Ryukyu Islands, Japan).
Material examined. 4 males, 5 females, along Neishan Road, Fanlu
Township, Chiayi County, Taiwan, 23.3640° N, 120.6115° E, 546 m, 3
IX 2018, KA, AS, ZHS & TEFP leg., ex. Ficus variegata (TW18-262A) [3
males, 3 females: NMS; 1 male, 2 females: BRH]; 1 male, 1 female, same
tree as the former (TW18-262B) [NMS]; 3 males, same tree as the
369
Journal of Asia-Pacific Entomology 23 (2020) 364–370
Fig. 5. Silba inubiwa MacGowan and
Okamoto, 2013, Amami-Ôshima Island, the
Ryukyu Islands, Japan. A: Female adult
walking on syconium; B: female adult lo-
cating the entrance of the syconium (os-
tiole) in late C phase (J18-133); C: ovipo-
sition at the ostiolar bracts (J18-133); D:
hatched larvae on ostiolar bracts (J18-133)
(black arrows: larvae; white arrow: empty
egg).
former (TW18-262C) [NMS].
Distribution. JAPAN: Ryukyu Islands (Ishigaki-jima Is.). TAIWAN:
Chiayi County.
New record from Taiwan (Fig. 1).
Host plant. Ficus variegata Bl. and Ficus benguetensis Merr.
Bionomics. Larvae were collected from fallen syconia of F. var-
iegata, which contained no fig wasps. Adults emerged during the period
14–17. IX. 2018. Beetle larvae of the family Staphylinidae attacked
several pupae, making entry holes to predate the pupae (Fig. 6). They
left the puparium after consuming the contents (Fig. 6B).
MacGowan and Okamoto (2013) found larvae of S. ishigaki in sy-
conia of F. variegata, which contained no fig wasps, in the B (receptacle)
phase and suggested that the larvae fed on the tissue of fresh syconia.
We assumed that syconia damaged by larvae feeding in the B phase had
fallen to the ground without ripening, and that the larvae kept feeding
on the syconia tissue regardless of status of the syconia.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
K. Arimoto, et al.
Acknowledgements
We thank the researchers mentioned in the Materials and Methods,
Reo Itô (Ôita City, Ôita, Japan), Taisuke Kanao (Okinawa Institute of
Science and Technology Graduate University, Kunigami District,
Okinawa, Japan), Naoki Ogawa (University of California, Riverside,
California, U.S.A.), Takaomi Sugimoto (Naha City, Okinawa, Japan)
and Takahiro Yoshida (Ehime University, Matsuyama City, Ehime,
Japan) for their help with the field work. Hsy-Yu Tzeng also helped to
obtain permission to conduct research in Taiwan. The research activ-
ities on Yakushima and Okinawa Islands were approved by the
Yakushima District Forest Administration Office (Yakushima Town,
Kumage District, Kagoshima Pref.), Kunigami Village Office (Kunigami
Village, Kunigami District, Okinawa Pref.), and Hokubu Agriculture,
Forestry and Fisheries Promotion Centre, Okinawa Prefectural
Government (Nago City, Okinawa Pref.). This is a contribution from
BRH.
References
Chen, C.H., Chou, L.Y., 1997. The Blastophagini of Taiwan (Hymenoptera: Agaonidae:
Agaoninae). J. Taiwan Museum 50, 113–154.
Christenhusz, M.J.M., Fay, M.F., Chase, M.W., 2017. Plants of the World: An Illustrated
Encyclopedia of Vascular Plants. University of Chicago Press, Chicago.
Galil, J., Eisikovitch, D., 1968. Flowering cycles and fruit types of Ficus sycomorus in
Islael. New Phytol. 67, 745–758. https://doi.org/10.1111/j.1469-8137.1968.
tb05497.x.
Katsoyannos, B.I., 1983. Field observation on the biology and behavior of the black fig fly
Sibla adipata McAlpine (Diptera, Lonchaeidae), and trapping experiments. J. Appl.
370
Journal of Asia-Pacific Entomology 23 (2020) 364–370
Entomol. 95, 471–476.
MacGowan, I., Compton, S.G., 2018. A new species of Silba (Diptera; Lonchaeidae) as-
sociated with figs. Zootaxa 4455 (1), 196–200. https://doi.org/10.11646/zootaxa.
4455.1.10.
MacGowan, I., Okamoto, T., 2013. New species of Lonchaeidae (Diptera: Schizophora)
from Japan and a re-evaluation of genus Setisquamalonchaea Morge. Entomol. Sci. 16,
196–202. https://doi.org/10.1111/ens.12003.
MacGowan, I., Razak, N., Rotheray, G.E., Ahmad, I., 2012. A new species of fig-feeding
Lonchaeidae (Diptera: Schizophora) from India and a checklist for the family in the
Indian sub-continent. Zootaxa 3242, 47–52. https://doi.org/10.11646/zootaxa.3242.
1.3.
MacGowan, I., Rotheray, G.E., in press. Chapter 67. Lonchaeidae (lance flies) pp.
1397–1406. In Kirk-Spriggs, A.H., Sinclair, B.J., (eds), 2019. Manual of Afrotropical
Diptera. Volume 3. Brachycera: Cyclorrhapha, excluding Calyptratae. Suricata 6.
South African National Biodiversity Institute, Pretoria.
Okamoto, T., MacGowan, I., Su, Z.-H., 2012. Predation on the pollinating fig wasps of
Ficus erecta by larvae of Silba sp. (Diptera: Lonchaeidae). Entomol. Sci. 15, 288–293.
https://doi.org/10.1111/j.1479-8298.2012.00522.x.
Palmieri, L., Augusto, R., Pereira, S., 2018. The role of non-fig-wasp insects on fig tree
biology, with a proposal of the F phase (Fallen figs). Acta Oecol. 90, 132–139.
https://doi.org/10.1016/j.actao.2017.10.006.
Silvestri, F., 1917. Lonchaea aristtella Beck. (Diptera, Lonchaeidae) dannosa alle in-
florescenze e fruitescenze del Caprifico e del Fico. Bolletino del laboratorio di
Zoologica generale e agraria della R.Scuola superiore d'agricoltura in Portici. 13,
123–146.
Wachi, N., Kusumi, J., Tzeng, H.Y., Su, Z.-H., 2016. Genome-wide sequence data suggest
the possibility of pollinator sharing by host shift in dioecious figs (Moracea, Ficus).
Mol. Ecol. 25, 5732–5746. https://doi.org/10.1111/mec.13876.
Wilkie, P., Poulsen, A.D., Harris, D., Forrest, L.L., 2013. The collection and storage of
plant material for DNA extraction: the teabag method. Gardens’ Bull. Singapore 65
(2), 231–234.
Yokoyama, J., Iwatsuki, K., 1998. A faunal survey of fig-wasps (Chalcidoidea:
Hymenoptera) distributed in Japan and their associations with figs (Ficus: Moraceae).
Entomol. Sci. 1 (1), 37–46.