Gall-feeding habits in Lepidoptera of Japan. IV : Tortricid moths reared from two cecidomyiid fruit-galls collected in late autumn

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TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
ut8tw Trans.Iepid.Soc.Japan 55 (4):280-284,September20e4

Gall-feedin hgabits in Lepidoptera of Japan. IV: [Ibrtrici dmoths reared from
two cecidomyiid fruit-gali closllected in late autumn
Kazuo YAMAzAKi' )and Shinj iSuGiuRA2)
i' Osaka City Institu otfe Publi cHealt hand Environrnent aSlciences,
8-34 Tojo-cho, Tennoji, Osaka, 543-O026 Japan
i) Laborator yof Fores tEcology ,Graduate School of Agricultur eK,yoto Uniyersity,
Sakyo-ku, Kyote. 606-8502 Japan
Abstract This study examined tortricjds reared from two cecidomyiid fruit-ga lclolslected in late
autumn. 1) The ftui tgall sof a cecidomyiid on the herb Achyranthe s,foponic a(Amaranthaceae)
were sampled on 16 October and 20 November, and reared in the laboratory .The galls were fed on
by the 1arva eef two tortricid species, Lobesia aelopa Meyrick and Adoxophyes dubia Yasuda and
the attack rate was les sthan 5%. In addition, since the 1arva eof the gal lmidge had already
matured on 16 October and most larvae had egressed from the gall sby 20 November, the tortricids
imposed littl neegative effect on the cecidomyiid larvae. 2) Adoxqphyes d"bia emerged from the
frui tgalls of a cecidemyiid, Asterulobia sp. on the liana 7}"achetospermum asiatic"m (Apo-
cynaceae), The larv aofA. dubia bored into and fed on gal 1tissu eand the gall was considerably
damaged. However, mest cecidomyiid larvae matured or had already exited the galls .In addition,
the attack rate was 8.3% (IV!12 )T.hese two tortricid species are faeultati cveecidophages that
primarily utilize normal plant tissues rather than plant galls .The tortrici d1arva emay feed on gall
tissues as an alternatiye food source due to a dearth of fresh leaves in late autumn.
Key words Frui tgalls f,acultati cveecidophages, Tortricid aLeo,besia aelopa, Adoxophyes dubia,
Introduction
Plant ga]l sare usually palatab lteo herbivore sand contain more nitrogen, sugar and less
defensi vcehemicals than normal plant tissues, although there are some exceptions (Mani,
1964; Abrahamson & Weis, 1987; Price et al,, 1987). Galls bear nutritjous undilferentiated
tissue seven during autumn and winter when plant growth ceases, Therefore ,several
herbivore sappear to utilize gall sas a food source especially during these seasons.
However, records of gal1-feedi nLgepidopter afrom lat eautumn to winter are scarce.
The [Ibrtricid iase a species-rich microlepidopteran family whose members haye various
host taxa, host ranges and voltinism (Scobl e19,92; Yasuda, 1998a): Many tortrici dspecies
exhibit concealment activities such as leaftierin agnd boring .Their host plant scomprise
divers eangiosperms and gymnosperms (phaneroga masnd) their host ranges also vary
depending on the species from monophagy to polyphagy .Many tortrici dlarvae emerge in
the spring and feed on newly emerged leaves ,but some species pass several generation pser
year ,Thus, tortrici dlarva eare likel yto have many chances to encounter, bore into and feed
on various plant gatls .Tb our knowledge, among the lepidoptera fnamiIie smembers of the
rlbrtricidae
are most frequentl yreported as cecidophages (e .g. Ybshiyasu ,1987; Clancy,
1993; Abe, 1995; Sugiura & Yamazaki, 2e02). The larva eof multivoltine tomicid species
occur in autumn as well as in the spring and feed on plant tissues. Therefbre, some
multivoltine tortricid larvae may use gall sas a fbod source from late autumn to winter. We
fbund two gall-feeding tortrici dspecies from two differe ngtall scollected in lat eautumn.
In this report, their cecidophagous (gall-fee dhiabnigts) are recorded,
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Gall-feedin Hgabits in Lepidoptera of Japan (4) 281
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Fig. 1,FAig i.mit gal lof a cecidomyiid on A.japonica, Scale bar: 5 mm.
2,FAig i,mit gall ofA,yteralobia sp. on T. asiaticum. Scale bar: 1O mm.
3, A male Lobesia aelopa adult eclosed from a cecidomyiid gal lon A. japonic aW.ing span:
10.3 mm.
Fig. 4. A male Adoxoph),es d"bia adult eclosed fi/om an Asteratobia gal lon T. asiaticum. Wing
span: 16.3 mm.
Galls examined
We collected the fbllowing two gall sin late autumn in a rural area of central Japan: 1) imit
galls of a gall midge (Cecidomyii dDaiep,tera >on the herb Achyranthes J'aponica
(Amaranthacea ea)nd, 2) fmit gall sof the gal lmidge Asteralobia sp. (Cecidomyiidae,
Diptera )on the ]iana 77achetospenmum asiaticum (Apocynaceae).
1) Cecidomyiid gall son A. japonica
An unknown cecidomyiid species induces oval gall sin the utric]es of A. joponi cian
October (Fi g1,) ,The gall sare about 5.0 mm in diameter and each one bears 1-5 larval
cells, of which each contains one yellow rnidge larva .Matured larva eexit from the gall in
November and crawl into the soil to hibernate .Severa glall soften fbrm as aggregates on
the spikes (Fig .1). However, the life cycle between winter and the fbllowing autumn has
not been clarified, In addition, this cecidomyiid has not been identifi e(dYUkaw a&
Masuda, 1996) . However, once it staxonomy is established, it will be easy to identify
based on it scharacteristic ga]1 morphology and host plant.
2)Astevalobia T.
galls on asiaticum
The cecidomyiid Asteralobi asp. defbrms the fruit osf T. asiaticum into curved club-shaped
gall sin autumn (Fi g2,). [[Ih geall sare about 30 mm in length and each bears 10-25 1arval
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282 Kazuo YAMAzAKi and Shiaj iSuGTuRA
Table 1. No. of emerged tortricid moths reared from
cecidomyiid frui tgall son Achyianthe sJ'aponica.
Samplingdatcs

Tortricidlarvae 16Oct20 Nov.

Lobesia aetqpa 6 6
Adoxophyes dubia 31739226 21465
No. ofgalls sampled
No. of spikes sampled l15
cells of which each contains one midge larva .In lat eautumn, most larvae mature in their
cells or have already exited from the gall. The larvae overwinter in the soil and the midge
ecloses as an adult in the spring (YUkaw a& Masuda, l 996).
Study
sites and procedures
The gall swere collected alengside a hiking route (c a80 m a, s. 1. ,34017'N, 13509'E) fi;om
Kyoshi te Aigaeri, Misaki-cho, Osaka Prefecture, central Japan, There, warm-temperate
forest sare dominated by the evergreen oak euereu pshitlyraeoid easnd plantatio nosf the
Japanese cedar CTIyptomeri ay'aponic aand the hinoki cypress Chamaec>paris obtusa.
Achynanthesjaponica also abundantly grows aleng this route. Several T. asia'ticum patches
also occur there.
Cecidomyiid gall son A, japoni cwaere sampled on 16 October (173 9galls )and 20
November (1465 2002. ElevenAsteralobia T.
galls) , galls on asiaticum were collected on
26 November, 2003. At the time of sampling, the Asteralobi agall shad many minute holes
from which most midge larvae and parasitoid shad alread yexited,
The gall swere cut off from the host plants p,laced in plasti cbags, brought to the laboratory,
and lef tin plasti cbags under laborator yconditions. The feeding habits of gall-feeding
]epidopteran larvae were observed, and the eclosed adults were identifie dG,alls infeste dby
the larvae were dissecte dwith a cutter and thei rinside swere inspecte dto determine the
effects of the larva eon the gal1 fbrmers.
Results
1) Cecidomyii gdall sonA. J'crponica
A small number of two tortricid species, Lobesia aelqpa Meyrick (Fig 3.) and Adoxqphyes
dubia Yasuda, eclosed in December from gall sthat were sampled on both 16 October and
20 November (Tlab 1l)e ,The larva espan the gall wsith the utricles and fed indiscriminately
on the galls ,spikes and utricles. The attacked gall swere seriously damaged, but most
midge larvae exited from the gall sby 20 November.
2) Asteralobi agall son T, asiaticum
A male A, dubia adult (Fi g4.) emerged from the gal lof the gall midge Asteralobi asp, in
early January of the fo11owin gyear .The larva ofA. dubia bored into and fed on gall tissue
and the attacked gal lwas considerably damaged, However, most larvae of the gall midge
matured or had already exited from the galls and the attack rate was only 9, 1% (AJL11),
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Gall-feedin Hgabits in Lepidopter aof Japan (4) 283
Discussion
1)Cecidomyiid A.
galls on ,J'aponica
On]y 9 and 8 tortrici dlarvae fed on 1739 (16 Oct.) and 1465 (20 Nov.) gal!s ,respectively
(Tabl 1e). Each larva usually fed on 2-5 gall sduring it slarval period ,Thus, the galls
attacked by the tortricids accounted fbr less than 5% of the gall ssampled. Furtherrnore, the
larvae of the gall midge had already matured by 16 October and most 1arvae had egressed
from the gall sby 20 November. These tortricid stherefbre appeared to impose little
negative etfect on cecidomyiid larvae.
Larvae of L. aelopa feed on various plant part s(leav efsi,ower sand fruit so)f divers eplant
species such as the goldenrod Soti(lag aoltissima (Asteracea eth)e, labiat eRerill ,afrutescens
(Laibiat athee) t,ea Camellia sinensis (Theacea aend) the grape Viti slabr"sc a(Vitaceae)
(Ba e& Komai, 1991; Nasu, 1993). A related species, L. vir"lenta Bae et Komai, is also
polyphagou sand occasionally infest sCeratovacuna nekoashi (Sasak i()Homoptera,
Aphididae) gall son Styrex J'aponicu (sStyracac e(aBea) e& Komai, 1991).
Adoxophyes dubia is also polyphagous and feeds on many plants such as [,},oni aovalifolia
var. elliptica (Ericacea ean)d Ribes fascicul a(tSaaxifragrac e(aYea)suda ,1998b) . These
tortrici d]arva eare therefbr eclassified as facultativ ececidophages (gall-fee dtehrats)
primaril yutilize normal plant tissues rather than p]ant galls (sens Muani, 1964),
2) Asteratobia gall son T. asiaticum
For the Asteralobia galls ,the attack rate by A. dubia larva was ]ow (9,1% )and the attack
was lat eenough for cecidomyiid larva eto mature and exit from the galls .Thus, the impact
on the gall midge by attack from A. dubia larva appears to be negligible. As mentioned
above, A. dubia is a polyphagous tortrici dand facultati vceecidophage.
3) Gall sas alternative food sources for late-autum ntortricid larvae
In late autumn, most plant scease their growth, senesce and withen Even evergreen trees
bear few fresh leaves in late autumn. On A, japoni canad T. asiaticum, there were no fresh
leaves present on the sampling dates .These tortricid larva etherefore might feed on gall sas
an alternative food source due to a dearth of fresh leaves and young plant tissues .Sugiura
& Yamazaki (2002 )reported that two tortrici dlarvae ,(lypsonoma bijlasci aKtuaznetzov
and G, hivanoi Kawabe, feed on the cecidomyiid bud gall sof willows in early spring, and
suggested that the ]arva ehatched befbre bud break and utilized galls as an alternati vfeood
source. The utilization of gall sas an aiternative fbod source due to a lack of fresh plant
tissue smay be widespread during cold seasons.
Acknowledgments
We thank Y Nasu of the Agricultural, Food and Environmental Sciences Research Center
of Osaka Prefectur ef,or the identificat iofon tortricid moths and valuable advice,
References
Abe, Y,, 1995, Relationships between the gall wasp, Trichagalma ,yerrat a(eAshmead )(Hymenoptera:
Cynipidae ),and two moth species, Andrioptect apulverula (Meyric k()Lepidopte rToaf:tricidae )and
Characoma rqfici.rra (Hampson )(Lepidopte Nroac:tujdae ).Appt. Ent. Zbol. 30: 83-89,
Abrahamson, W. G. & A. E. Weis, 1987. Nutritiona lecology of arthropod gall makers. Jn Slansky, F. Jr &J.
G, Rodriguez (Eds )Md,ritiona lEcology ofInsec Mtist,es ,SPide ransd, related Invertehrates :235-258.
Wiley& Sons,New York.

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284 Kazuo　YAM．　AzAKI 　and 　Shinj　iSuGJuRA
Bae，　Y．−S，＆ F．　Komai，199L　A　revision 　of　the　Japanes　especies 　ef　the　genus　Lobesia　Guen6e（Lepidopter，a
　　Tortricid）a，　ewith 　descripti　oofn　a　new 　subgenus ．7ンδσα 42：115−14L
Clancy，　K．　M ．，1993、　 Adaptation　sof　ga［lin　gsaw 且ies　to　natural　enemies ．　 In　Wagner ，　M ．　R ＆ K ，　F．　Raffa
　　（Eds），　Sawfl　NLtfe　Histo刑y　Adaptati．ons 　to　woody 　Plants：295−330、　Academic 　Press，　New 　York．
Mani ，　M 　S．，1964．　Ecology　ofPlant 　Ga正ls．434　pp．　Dr　W ．　Junk，　The　Hague．
Nasu，　Y．，1993．　New 　host−plant　sof　Lobesia（Lobesia）αゆραMeyrick　Proc．　Kansai　Plant　Prot，50c．（35）：
　　45−46 （in　Japanes）e．
Price，　P．　W ．，　Fernandes，　G 　W ．＆ G ．　L 　Waring ，1987．　Adaptive　nature 　of　insec　gtall．s翫 vか．　Ent．16：15−24．
Scoble，　M ．」．，1992．　The　Lepidop　te　ra：Form，　Function　and 　Diversit．y404　pp．　Oxfbrd　Uuiv．　Press，　Oxfbrd．
Sugiura，　S．＆ K ．　Yamazaki ，2002，　 Tortricid　moths 　reared 　f士o皿 cecidomyiid 　bud　gall　son 　willows ．　 Trans．
　　lepid．　Soc．」吻απ 53：116−118．
Yasuda ，　T．，1998α．　Biology　of　tortricid　moths ．　In　Yasuda ，　T．，　Hirowatari，　T．＆ M ．　Ishii（Eds），　Biology　of　the
　　Microlepidopte：r2a09−230．（In　Japanes）e．
　　　 ，1998わ．The　Japanese　species 　of　the　genus　Adoxophyes 　Me 〕加ck （Lepidoptera，　To貫ricidae ）．　Trans．
　　lep　id．　So（〕．ノapan 　49：159−173．
Yoshiyasu，　Y，，1986．　Notes　on　two　lepidoptero　usspecies　attacking 　the　galls　of　Schlechtendal　icahinensis
　　（BeU ）（Aphididae）on　Rんus ノαv硼 ica　L．乃δGα37：97−100 （in　Japaneヨe　with 　English　summary ）．
Yukawa ，」．＆H．　Masuda，1996．　Insect　and 　Mite　Gatls　o．プJapan　in　Colors．826　pp．　Zenkoku −n6son −ky6iku−
　　ky6kai，　Tokyo．（ln　JapaIles）．e
摘 要
日本の 鱗翅 囗に おける ゴール食の 習性 4．一晩秋に 2種の タマ バ エ による・果実 ゴール から得
ー 一 ・
られたハ マ キ ガ （山崎 夫 杉浦真治）
本論文で は 大阪府岬町孝子におい て 晩秋に採集 した 2種の ゴールか ら得られたハ マ キガ科に関し
，
て報告する．
1）イノ コ ズ チ ミフ ク レフ シ （タマ バ エ 科の 一種によ りイノ コ ズチ の 胞果に形成された ゴ ール）をサ
ン プリン グし室内飼育する と，ホソバ チビ ヒ メハ マ キLobesia　aelopa 　Meyrick とウス コ カクモ ンハ マ
キAd・xophyes 　dzabia　Yasudaの幼虫がゴ ール と胞果 を無差別 に摂食してい るの を確認するこ とがで き
た．ゴ ール の 2種のハ マ キガに よる攻撃率は 5％ 未満と低 く．寄主の タマ バ エ 幼虫はすで に ゴールか
ら越冬の ために脱出済みか成熟して 脱出直前で あっ た．その た め，これらの ハ マ キガ幼虫の タマ バ エ
に対する影響 はほとんどない と考えられた．
一
2）テ イカカズラ ミサキフクレフシ（タマ バ エ 科の 種Asteralobi　sap，によりテイカ カズラの実に形成
され たゴール）に は，ウス コ カ クモ ンハ マ キの 幼虫が潜入 してい た．攻撃されたゴ ール はか なり劣化
して い たが，タマ バ エ 幼虫はすで に脱出済みかその直前であっ た．また，ハ マ キガの ゴールへ の攻撃
率は8，3％ （N＝12）であっ たため，タマ バ エ に対する影響は ほ とん どない と考えられた．
これ ら2種のハ マ キガ幼虫は ゴール 以外の通常の植物組織をお もに摂食する機会的えい 食者である．
晩秋で 新鮮 な葉がない ため に ゴ ール 組織を代替食物として使用してい たと考えられ る．
，
（Accepted　May 　29，2004）
Publishe　dby　the　Lepidopterological　Society　of　Japan，
5−20 Motoyokoyama 2，Hachioj，iTokyo，192−0063Japan
， 　 　 　 　
一
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