Table Of ContentTThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
matwh 7)JOtoCa 45(2):105-112,June1994
Factors determining laryal, pupal and adult polyphenisms in the tropical
butterflyC,Zztopsilia (F.)(LepidopterPaie,ridae)'
pomona
Minoru IsHII
'Osaka
Entomological Laboratory, Colleg eof Agrieulture, Universit yof Prefecture,
Sakai,Osaka 593,Japan
Abstract To know the determinant sof adu]t wing forms ,f .crocale and f .Po7nona ,and larva!
and pupal coloration of the Sabahan population of the Lemon Migrant, CatoPsil Pioamona, the
larvae inisolation in (10-hour)
were rcared or crowds under a short photoperiod or natural
photoperiods (abou t13-heur) at room temperatures (22-3℃4) .The larval photoperiocl
affected the adult forms : several intermediate forms emerged under the short day, while all
the inclividual wsere of f .crocate under natural photoperiods. A phase variation was observed
in the larva lstage : conspicuous black latera sltripes appeared in the larva ereared in crowds,
while the larval body was almost green in the solitary rearing. The high densit yalso caused
the sLight shortening of larval stage and the decreas ein pupal size. The pupal coloration was
proved to be deterrnined by the ground color of substratum for pupation : yellowish green and
brown backgrounds induce dgreenish ancl brownish pupae, respectively.
Key words Butterfly ,CaloPsili aPomona, Malaysia, deterininant ,wing form, phase variation,
pupal coloration, photoperiod.
TrQpical butterfli aerse subjected to nearly constant conditions of daylengt hand tempera-
ture all the year round. It is ,however, not that we can not fin dany `tseasonalities" in the
lif eof tropical butterfii esF.or example, Owen (197 1p)ointed out that in tropical Africa a
considerable number of satyrids and some nymphalids have two "seasonal forms'' i, ,e,, the
wet- and dry-seaso nforms, which replace each other as the season changes. I also
collected differen tseasonal forms of such butterfli aess Catopsili aPomona, Eurema hecabe,
Melanitis ledo, etc. during the short research period in Sabah, Malaysia in Jul yand August
of 1981 (Tabl 1e) .Among them the Lemon Migrant, C. Pomona has the two adult forms,
f .crocaie and f. Pomona detemined by the larval photoperiod in the population of southern
Japan(Yata Tanaka, 1979).
and
Here arises an intriguing question, what factors are responsible for the contrel of the
seasonal replacement of adult forms in the tropics , In order to elucidate the determination
of adult seasonal forms in tropical butterfii ersea,ring experiments were made with C.
Pomona (Fabriciu isn) S,abah, Malaysia in 1981. In addition, the determinants of larval
melanization and pupal coloration will also be deal tin this paper.
Materials and Methods
Eggs of the Lemon Migrant, CatoPsil ipaomona were obtained from females of f. crocale
captured around the Forest Research Center in the suberbs of Sandakan, Sabah in late July
of 1981. The hatched larvae were divide dinto 2 groups and reared under a 10-hour or
natural photoperiods (abo u1t3 hours includin gmorning and evening twilights) in the
' IL'hi wsork was supported bir a Grant-in-ai fdor Scientifi cResearch (Oversea sScientifi cSurvey}
from the Ministry of Education, Science and Culture J,apan (Nes .56041038 and 57043034).
NNIII-IE-leEcltreoncitcronic LMbirabrryary Service
TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
106 Minoru IsHii
`'seasonai
Table 1. Nurnbers of each forms" of 3 butterfl yspecies collected in Sabah in
Jul yand August, 1981, WSFi wet season form incrudin fg .crocage in C. Pomona,
DSF : dry season form including f. Pomona in C. Pomona, IMF : intermediat feorm.
Species VLiSF iMF DSF Total
Pieridae
Ctztopsi tPioamona 1136 3o llle 2546
Eurema hecabe
Satyridae
Melanitis lecla 3 3 1 7
laborator yof the center in August. Each group was subdivided into 2 groups and reared
in isolation (an individua lper a 200-ml transparent plastic cup) or in crowds (5 individuals
per the same plastic cup). Larvae were daily previded fresh Ieaves of Cassi asp. as food,
The photoperiod was controlled manually : larvae of the short-day rearing were put on a
desk illuminate bdy fluoresce nltamps and/or the Iigh tthrough the window in photophases,
and put in light-tig bhotxes in scotophases. In the present study, the temperature was not
controlled so that it fiuctuate bdetween 22 and 34"C , Duration sof larva land pupal periods,
larva lcoloration, and pupal body lengt hwere recorded,
In the course of experiment, I found a variation in pupal coloration of this pierid: pupae
formed on the Cassi aleaf were green, while some en the wall or li dof plastic cup bore
brown or intermediat ceolors (se Feigs 6-7). Thus in order to know the factor determining
the pupal coloration, 10 fina linstar larvae reared under natural photoperiods were put into
each of the two types of clear 200-ml plasti ccups, those wrapped in the thin yellowish-
green paper and those in the thin brown paper, at the wandering stage, and they were put
on the desk by the window.
Results discussion
and
Larval sttrge and Pmpai si2e
Larval photoperiods seemed to uninfiuence on the larval development in the present pierid :
the mean larval stage under a 10-hour photoperiod did not diffe rfrom that under natura]
daylengths in both solitaly and crowded conditiens (Tabl 2e, P>O.05 by t-test). On the
other hand, the rearing densit yappeared to have an effect on the larva lgrowth : under each
photoperiodic condition the mean larva lstage of the crowded group was slightly shorter
than that of the solitary one and the differenc ewesre significant (P<O.05 )T.he pupal size
appeared to reflect the duration of larval stage : the mean pupal sizes estimated as the
pupal body length were greater in the solitary groups than in the crowcled ones (Table2;
P<0,05) . The differen cseeemed more evident under the 10-hour photoperiod than under
natural photoperiods. The results dernonstrat tehat the high densit yduring larval period
causes both the shortening of larval stage and the decrease in adult size in this pierid.
Phase inlarvae
van'ation
It was interesti ntghat the larva lcoloration varied with the rearing density ,whereas
photoperiods seemingly had no effect on the determination of the larva lcoloration. In
Figure 1, fiv egrades of･the larva lcoloration used by Fukuda (1981 )an,d the percentage of
NII-Electronic Library Service
TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
Polyphenism in a Tropical Butterfly 107
Table 2. Mean larval stages and pupal sizes, and nuinbers of crocale, Pomona and intermedi-
ate forms emerged under differe ndtensit yand photoperiodic conditins in the Lemon
Migrant, CZztoPsii iPaomona.
No.
Conditions No. ofPupati on Larval stage Pupal size of adults emerged
(mean±S.D. days) (mean- S･.D. mm) f .crocaleInt.f. Pomona
solitary
leL-14D 1515 8.40+O.69a 3LOO+1.19a 910 4o oo
natural 8.60+O.51a 30.36+1.20a
crowded
10L-14D I312 8.15 ±, O.38b 27.23±1.29b 811 3o oo
natural 8.08;O.29b 29.05+L71b
Means followed by the same lette risn a column are not significantly differe natt 5% 1evel by
t-test.
NaturatDaytength 10-hourDaytength
MSo[itary
BO
------e---- ---------------
EIiEgcrowded
BI
-----}--t----------------e- ---"--------------"--t---------
B2
-----te-----t-- ---t----------
B3
..
IV IV
B4
BO
BI
B2
.-- -----------t
B3
v '"-"-"----- '.-'-･-'-･-"-"-'-･.'-"-･'･'"･'" v
B 4 BO BI B2 B3B4
--}t---------------------tt---- ----------------t---------
(%) 50 OO 50 (%)
Fig. 1. Five grades of larval coloration (BO-B4 u)sed by FuKuDA (1980 a)nd, percentages
of 4th (IV )and 5th (V) instar larvae of each grade in solitary and crowcled rearings
under a 10-hour and natural-(ca. ]3-hour) photoperiods in the Lemon Migrant,
CZztoPsig iPaomona Cse etext).
4th and 5th (fin ainl$)ta rlarvae of each grade under the 4 differe nctonditions are shown.
The "density effect" became remarkable after the 4th instar stage: although the larvae
were uniformly yellowish green til lthe 3rd Iarval instar ,about a half of the 4th instar
larvae and all the 5th instar ones of the crowded group had conspicuous black Iateral
stripes (Fig s2.-5). In most 4th instar larvae and about a half of the 5th insta rones of the
NII-Electronic Library Service
TThhee LLeepipdiopdteorpoltoegircaollSoocgleitycal Society oofJfap anJapan
I08 ]vlinorIusHu
4
s,..",,.i,,-i,I
;5i
Figs2-5 .Final {5th )instar larvae of CJLitopsi lPioamona,N2- 3L,arvatiof grade BO. 4-5.
Larva B3.
of grade
.'L't'"
''
't /tt ttttit
:, t /.t't'"
tt tt
e.
･.z. alj I . agg-
. 7- . ti G . '. B "u'･
Figs6-7. Variation of pupal colorationin Ctztof}s Piolimaona ,6. Agreenish pupa formed
on the underside of CZiss ilaeaf .7. Greenis h(G) and brownish (B )pupae obtained
from the present experiment (se etext).
NII-Electronic Library Service
TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
Polyphenism in a Tropica lButterfly 109
l
H
'Kg.-
,T
..r'-'
x
Xtt
S
8' -- o[
･ll
"'t;
iiil-,
f-
't- ttt.tl.,
,
9 11 K.,mp
Figs8-11. Intermediate-fonm adults of datopsil iPaomona obtained from the present
experiment. Dorsal and ventral views of a male <8-9a)nd a female (10-11),
solitary group, contrastively, the body coloration was alrnost green. The differenc eisn the
proportions of larvae belongin gto each grade were significant between solitary and
crowded groups and between 4th and 5th insta rlarvae despite the photoperiodic conditions
(Tab]e3).
The phenomenon in which such characters as morphoTogy, coloration, physiology, behavier
etc. of an insect species change remarkably in response to the population densit yhas been
known as the phase variation, and in the insects such as Iocust sand armyworms, the phase
variation is deeply related to the migratory nature (e g.., Iwao, 1956) .In the present pierid,
hewever, the differen cien sthe behavior between solitary and crowded groups were still
unclear like the Japane sleibythei bdutterfl yL,ibythe aceltis studied by Akayama (198 1)I.n
addition, the larval coloration seemed independen tof adult forms in this pierid (Table2).
Determinant of Pzipa lcolo7ation
All of the 10 pupae formed on the wall or Iid of the plasti ccups wrapped in the yellowish-
green paper were greenish, while all 10 pupae in the cups wrapped in the brown paper were
brownish (Figs6- 7T)h.e results show that the pupal celoration of the present species is
determined by ground color of the substratum for pupution. The similar determinatio nof
NII-Electronic Library Service
TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society oofJfap anJapan
IIO Mineru lsHII
Table3. Significanc eofs the differenc eby chi-square test in the proportions of
larvae belonging to eaeh color grade between the groups reared in different
densit yandlor photoperiodic conditions. N.S.i P>O.05 ,': O.e5>P>O.Ol,
**: O.Ol>P>O.OOI ,"'/ P<O.OOI, d.f, :degree of freedom ,10L: leL-14D,
NDL ] natural daylengths.
Paired groups Xz d.f. P
4thinstar
10L-solitary vs NDL-solitary 2.40 1223 N.S.N.S.***
IOL-crowded vs NDL-crowded 1.226.5012.00
10L-solitary vs 10L-crowdecl
NLD-solitary vs NDL-crowded
5th instar
10L-solitary vs NDIrsolitary 5.873.3010.982332.3746 N.S.N.S.*s**
10L-crowded vs NDL-crowclecl
10L-solitary vs 10L-crowded
NDL-solitary vs NDL-crowcled
4th vs 5th instars
10L-solitary 14.0322.07 3424 ******S*k
10L-crowded
NDL-solitary
7.8820.72
NDL-crowded
pupal coloration has been known in butterfli esnsch as Pieri smpae (Ohtak i1,960 )and
inachis io (Maisc hand BUckmann, 1987) .All the 20 pupae were emerged as crocale-form
adults despit ethe pupal coloration, showing that the determinant sof pupal coloration and
adult forms are independen tof each other in this pierid.
Determinantsofadugtforms
Yata and Tanaka (197 9r)eported that the larval short days of 10 and 11 hours induced the
Pomona form, while the crocale-form adults were emerged under the long clays of 13 and
14 hours at 20"C in the Okinawa population of the present species. Fukuda (198 1su)ggest-
ecl from his experiments with the Okinawa and Kagoshima populations that high tempera-
tures and/or high clensitie sshould suppress the Pomona-form production by the short clay.
The results of the presen texperiment did not seem inconsiste nwtith their conclusion.
AIthough no Pomona form was obtained in this experiment, the intermediate form adults
emerged only under a short (10-heu prh)otoperiod (Tabl e1) . By contrast, all the individ-
uals which experienced natural photoperiods during Iarva lperiods transformed into the
crocale forms, However, the rearing densit ydurin gthe larva lperiod appeared to unaffect
the adult forms in the present studv.
Because of a few-day's stoppage of electric power during the "Hari Raya Puasa" festival,
the room temperature became higher in the course of rearing, which may suppress the
development of the short-day individual sinto the pomona form. However, it may be sure
from the results of the present experiment that the larva lphotoperiod should be one of the
most important determinant sof the adult seasonal forms in this pierid , Nevertheles si,t
is strange that we can find crocale and Pomona furms in the same season in Sabah where
NII-Electronic Library Service
TThhee LLeepipdiopdteorpoltoegiroaollogical SSoooiceityety ooff JJaapapnan
PolyPhenism in a Tropica lButterfly ilI
natural daylengths including civil twilights range only from 12．5 to 13，5 hours（Beck 1980）
，
with monthly mean temperatures in lowlands being as constant as between 26 and 28℃
（Japan Meteorological Agency，1984）． Although the present species is one of the wel1 −
known migrants in the monsoon Asia¢ ． g．， Wiliams，1930；Yamada ，1973；Yukawa ，1983；
Ishii，1989）， the pomona−form adult is especially inferre dto liv eand to migrate longer than
the crocale −form adult because the ovarian development of the former is retarded after
emergence （Yata and Tanaka，1979）． It might be possible， therefore， that most pomona−
form adults are emerged in the regions of high altitudes where the temperatures are low
enough for the pomona−fQrm production and migrate into the lowlands as Sandakan to be
mixed with the crocale −form adults emerg，ed there，
Acknowledgetnents
Ithank Dr O， Yata of Kyushu Universit yand Mr H， Fukuda for their kind advices ． My
thanks are also due to Mr H． Matsuka who took photographs of Iarvae and pupae of
Catopsil ipaomona inserted in this report ． I am deeply indebte dto Dr T． Hidaka of
formerly Kyoto Universit yfor giving me the opportunity of pursuing the present study ． I
alsQ appreciate the kind assistance and help offered by Mr S． Lantoh and the other staffs
，
of the Entomologica lsection of the Forest Research Center Sabah Malaysia．
， ，
References
Akayama ， A ．，1981． Phase variation in a butterf，i Lyibythea celtis FruhstQrfe（rLepidQpter， aLibytheida）．e
1（oηbノ舜， TokPo 49；334−343． ’
Beck， S． D．，1980．加∫翩 Photoperiodisin．2nd ed．387 pp． Academic Press， New York．
Fukuda ， H ．，1981， SeasQnal且uctuations of two adult forms of Catopsil pioamona jn Taiwan and southcrn
Kagoshima ， and factors affecting the manifestation of the wing forms．［Kagoshimachttokoleoκ召納｝・u
Kiyo］（10）；53−75（in Japanes），e
Ishii、 M ．，1989． Observation sin the migratory Hight sof Catopsil ibautterfli（eLsepidopter， aPier［dae）in
Iw a oS， aSb．a，h1，9 5M6a．l ayPshieano． men7aンδ Gof（z p4h0a：se8 1v−ar8ia3tio， 重ユ in insects． Seibzftsufea｛g（Saukpzp1．｝：37−42（il lJapanes）、e
Maisch， A ． and D． BUckmann ，1987． The cQntrQI of cuticular rnelanin and ユutein incorporation of the
rnorphological cQlour adaptation of a nymphalid pupa ，∬nachis io． ノ． Jnsect 1〕h）’siol，33：393−402．
Ohtaki， T．，1960． Humoral control of pupal coloration in the cabbage 、、・hite butterf，i yPガ8廊 rctpae ．
ノln」・fotnes Zool．ゴaP ．33：97−103．
Owel1， D． F．，1971、 丁穿毎）i（・al BzttteZfii．e2s14pp 、 Clarendon Press， Oxford．
Yamada ， Y．，1973． Migration of Catopsili apomona Pomaona Fabricius in west Java．7｝1δCa 24：78−79（in
Japanese．
♪
Yata，0． and H．Tanaka ，1979． The e仕ect of photoperiod on the dimorphism of the Lemon Migrant ，
CatoPsili apontona Fabricius． 7｝16 Ga 30：97−106．
Y Wui kl alwCiaaarmi，s ，t JCa，， ，J． 1Ba9、v，8a31．．9 3乃0A，n δ コoGb「Vshez’er v3aル3tiFo ：nユ瓶8海5−o o1麗l8l6 o．fth cB 鋸mig飽ra壇torガy8s．47且3 ipgph．ts0 1ofi vCea ratn〔）dps iBloi ypado， mEodnian bpuormgo．na （Fabricius）in
摘 要
熱帯産ウス キシ ロ チ ョ ウにお ける幼虫， 蛹， 成虫の表現型多型の決定要因 （石井 実）
ー ・
マ レ シア， サバ 州産のウスキシロ チョ ウにお ける成虫の翅多型， 幼虫 蛹の色彩多型の決定要囚を明
一
NNI工I工-EElleoetcrotniroonic LLiibrbarryary Service
TThhee LLeepipdiopdteorpoltoegiroaollogical SSoooiceityety ooff JJaapapnan
112 Minoru ISHH
らか にするために，幼虫を密度と日長を変えて，室温 （22−34℃）で飼育した．その結果，幼虫期の日長
は成虫の翅多型に影響を与えることがわかっ た．すなわ ち， 自然日長区 （約 13時間）ではムモ ン型しか
得られなかっ たが， 10時間日長区で はい くらかの 中間型が羽化した，また， 幼虫を高密度 （200ml の容
器に 5個体）で飼育する と，体側が強く黒化する個体が多 く出現した．これ は多 くの昆虫で知 られ る相
一
変異現象の 種と考えられる．単独飼育区と比べ ると，高密度区の幼虫期間はやや短 く，蛹は小型であっ
た，さらに， 本種の蛹の色彩は蛹化する部位の背景色で決 まるこ とがわか っ た．すなわち， 緑色の紙を
背景に蛹化した個体は緑色，茶色の紙を背景に蛹化した個体は褐色であっ た．本種にお ける幼虫，蛹，
成虫の表現型多型がすべ て独立の要因に より制御され てい ることは興味ふかい。
（Accepted Januar y24，1994）
Published by the Lepidopterological Soc量ety of Japan，
c／00gata Building，2−17， Imabashi 3−chome Chuo −ku Osaka 541 Japan
， ， ，
一
NNI工I工-EElleoetcrotniroonic LLiibrbarryary Service
