Fecundity of the Yellow Swallowtail Butterflies，Papilio xuthus and P.machaon hippocrates, in a Wild Environment

著者 Watanabe Mamoru 雑誌名 Zoological science 巻 3 号 3 ページ 509-516 発行年 1986-06 権利 (C) 1986 Zoological Society of Japan URL http://hdl.handle.net/2241/100341 Fecundity of the Yellow Swallowtail Buｔｔｅｒｆｔieｓ，Ｐａｐｉｌｉｏｘｕtｈｕｓ

and ？． ｍａｃｈａｏｎＨｉｐｐｏｃｒａtｅｓ，１ｎ ａ￥ｆ＼＼ｄ Environment

Mamoru Watanabe and Kazuo NOZATO

DepaΓz所ｅ削げ召lology, Faculty of Education, Mie Univer∫ity,Tsu一shi,Mie jμ， 四d£α&θΓαzθり; of En哨所θ/θ舒，jりｃ�り'θ/≒Agricultuだ，瓦θ油zびniver.∫zり'， Ｎａｎｅｏｋｕ-ｓhi， Ｋｏｃｈｉ７８３，Ｊａｐａｎ

Reprintedfrom ZOOLOGICAL SCIENCE Vol 3，No. 3 (1986) p 509-516

Published by the

Zoological Society of Japan

June 15， 1986 ZOOLOGICAL SCIENCE ３ 509-516 (１９８６) ⑥1986 Zoological Society of Japan

Fecundity of the Yellow Swallowtail Butterflies, I°apilioxuthus and /).ｍａｃｈａｏｎ Ｈｉｐｐｏｃｒａtｅｓ，1ｎ a Wild Environment

Mamoru Watanabe and Kazuo NOZATO^

Ｚ）ｅｐａｒtｍｅｎt ｏｆ Ｂｉｏｌｏｇｙ， Ｆａｃｕltｙ ｏｆ Ｅｄｕｃａtｉｏｎ， Ｍｉｅ Ｕｎｉｖｅｒｓitｙ， Ｔｓｕ-ｓhi， Ｍｉｅ ５１４，

αz�1£αわθΓα1θり; of Entomθlogy, Faculり’θ/≒Agnculturら瓦θ助･びηiversity,

Ⅳangoku一助z，瓦θchi 783, Japan

ABSTRACT―In order to clarify the fecundity and the number of eggs actually deposited of the yellow swallowtail butterflies,乃7ρlllOXUが114sand ？ 謂αchaon, females were captured ｍ the wild and dissected Forewing length was not correlated with age in any of the two species Multiple matings suggested by spermatophore counts were found for P xuthus but not for P machaon in both spring and summer generations Eggs ｍ the ovaries were classified into three categories according to their degree of maturation The fecundity, which was ａ total number of eggs. was 500-600 eggs irrespective of generation and species The small body size of individuals belonging to the spnng generation seemed to affect not the fecundity but the number and the size of mature eggs ｍ the abdomen Oviposition ｍ the spring generation of j）ｘｕthｕｓtook place mainly at noon but that of the summer generation ｍ the morning Considering the oviposition period of both species. the number of eggs deposited ｍ each generation in each species was about 300 eggs, which was about a half of the ｆｅcundity

by dissection and estimates the fecundity and the INTRODUCTION number of eggs actually deposited by each female One of the yellow swallowtail butterflies, Papihθ during her lifespan ｘｕtKｕｓ，has been most intensively studied for Its

population ecology among Japanese swallowtail MATERIALS AND METHODS butterflies [1-2］Its oviposition activity ｍ the

summer generation has been studied by Yamanaka The data ｍ this study were obtained from ｅt al［31 Few studies on the number of eggs ？ｘばhus and P machaon. mainly in Kanagawa, deposited by the female during her reproductive Mie and Kochi Prefectures, warm-temperate zone life. however. have yet been published This may ｍ Japan Both species have more than four be due to her longer longevity, and partly due to generations per year and overwinter ｍ the pupal the difficulty ｍ obtaining eggs under laboratory stage 151 The imago of each species ｍ spring has conditions In addition, no reports have been characteristic wings different from that ｍ the other published on the population ecology ｍ spring seasons Since the adults found ｍ summer are not generation because of their low abundance com- clearly identified Its generation belonged in both pared to summer generation of Ｐ ｘｕthｕｓ species [2], females collected ｍ this study were

Beside the work on P xuthi£s, few studies on the divided into two, spring (Apnl to early June) and population ecology of Ｐ ｍａｃｈａｏｎ Ｈｉｐｐｏｃｒａtｅｓ summer (late June to October) generations Their have been reported ｍ Japan, though many of such habitats consist of margins of forests and open studies have been published elsewhere 141 lands with nectar plants and with host plants of The present study deals with the number of eggs larvae The latter IS Rutaceae (mainly Zanthoxy- in the ovaries of P xuthus and /）ｍａｃｈａｏｎ counted lｕｍ ａｉｌａｎtｈｏｉｄｅｓ)for P xuがzμ∫and Umbelliferae for j）ｍａｃｈａｏｎ Females were collected on var- lOUS occasions such as feeding, roosting, coupling Accepted December 17，1985 Received November l，1985 and ovipositing, from April to October during 510 M Watanabe AND Ｋ NOZATO

1981 to 1984 When the females were caught, their abdomens were immediately fixed into 50% ethyl alcohol and their wing conditions and sizes were recorded The ages were classifiedinto five de- Age ｏ grees from tearing ｃｏｎｄ由onsof the wings, as suggested by Nozato et al[6] All females were dissected to determine how many times they had mated, as indicated by the number of spermatophores in their bursa copula- trix,and to determine the number of eggs lemain- ing ｍ the abdomen Eggs in the ovaries were classified into three categories mature, submature and immature eggs Age l The mature eggs, which were determined by the criterion of Hidaka and Aida [7], were mainly found ｍ the oviducts They were large enough to be ready for oviposition and colored ｍ yellow The submature eggs were mainly found ｍ vitellar- mm of the ovaries They were small and colored lighter than the mature ones The immature eggs which include oocytes filledthe terminal filament. germanium and ａ part of vitellanum of the Age II ovanoles The eggs in ovarioles decreased in size toward the tip of the terminal filament filledwith oocytes Since both immature eggs and oocytes contain no yolk, they look like white The number of mature and submature eggs were directly counted For the immature eggs, their number was assessed for one ovanole on each side,

and the total number was estimated by multiplying Age III by the number of ovaries

RESULTS

In the course of the dissection, we observed ａ phenomenon which was common to both species As shown ｍ Figure 1，the newly emerged or fresh female (age O）had the most abundant fat body which filledm the cavity of the abdomen, while the Age IV oldest female (age ＩＶ）had the smallest one Contrary to this, the volume of the air sac increased ｍ size with ageing of the females

IT tears with frayed scales, Ill notched tears with Fig I Photographs of the abdomens of P xuth£IS, re- frayed scales, IV broken or extensive tears with moving off the cuticle The age class was classified frayed scales The abundant fat body without air sac mainly by the wing conditions, lｅ ｏ newly inhibits to observe the eggs ｍ the age O，while some emerged females with no visible damage and lus- large mature eggs can be seen in the age IV due to trousscales, l fine tears with less lustrous scales, less fat body with a large air sac Fecundity of Swallowtail Butterflies 511

It IS generally known that the forewing length IS again during the age Ｏ The number of times that ａ good index of the size of ａ butterfly The mean the females had mated increased significantlywith lengths derived from different study areas were their age （Ｐ＜005）Ｏｎ the other hand, most not significantly different from each other All females of ？ machaon seemed to mate once females from different areas were then pooled 64 The diameter of mature eggs ｍ the oviducts IS and 205 m P xuthus, and 22 and 44 m /）ｍａｃｈａｏｎ， shown ｍ Figure 4 It remained constant lrrespec- ｍ spring and summer generations. respectively The size of butterflies ｍ the spring generation was significantly smaller （Ｐく0 005) than that of summer one (Fig 2）Ｔｈｅ wing length was not correlated with their longevities ｍ any of the species This suggests that there IS no apparent relationship between the body size and the longev‘ ity in each generation, as in the case of the black swallowtail butterflies (Watanabe et al , unpub- lished) 一Ｊｏ£ａｏｌＥ」ωａａ ｏｚ

Figure 3 shows that many females of 7）ｘｕthｕｓ こ こ O I Ｈ Ill IV mated more than once Some females of the 0 1 11 111 1V oldest age （IV）ｍａtｅｄ maximum five times ｍ both spring and summer generations The females of age o have single spermatophore ｍ most but o or ２ ｍ ａ few Therefore, females of Ｐ ｘｕthｕｓmated soon after emergence and seemed not to mate

P xuthus Spring Summer Fig ３ Mean number of spermatophores ｍ ？ xuth心 60 and P machaon of respective ages ｍ both spnng and summer generations （士SE）Ｔｏtａｌ number of 50下匹 females examined was 46 and 143 m P xuがtus and 16 and 34 m /）謂αchaon in spring and summer genera-

（ＥＥ） tion,respectively

μ １２ 10 』 ｕi5ｕｄ一 〇ｃ一ＳΦＪｏＬ Ｑり ＱＪ ４ ０ ０ ０ P xuthus にｊ ｊ ｊ ｌ O I Ｈ Ill IV こ ｏ Ｉ Ｈ ｍ IV

Ｐ machaon ／ （ＥＥ）Ｊ８１９Ｅ一石六大

１４ 12 10 ☆ﾃﾞE 』

□1 01 LI」

こ こ ヘノ O I II Ill IV O I H Ill IV Spring Summer こ こ O I H Ill IV Ｏ I II Ill IV Fig ２ Mean forewing length of females in Ｐ ｘｕfhｕｓ Spring Summer and /）陶αｄzαθy?of respective ages in both spring Fig 4 Mean egg diameter in P xuthus and ？ machαθzz and summer generations with the maximum and of respective ages ｍ both spring and summer gen- minimum length Total number of females ex- erations (士SD）Ｔｏtａｌ number of eggs examined amined was 64 and 198 m P ｘｕfhｕｓａnd ２１and ４２ｍ was 44 and 480 m /）χぱ/1£ISand ３０and １１０ｍ Ｐｍａｃｈａｏｎ、ｍspring and summer generation、 re- Ｐ ｍａｃｈａｏｎｍspring and summer generation, rｅ｀ spectively spectively 512 Ｍ Watanabe AND Ｋ NOZATO

tive of age ｍ P xuがlus but increased with age in both generation, while those in age Ill and IV in the summer generation of P machαon (P<0 01) the spring generation were smaller than those ｍ Mean mature egg size ｍ the spring generation of the summer generation （P＜005）Ｔｈｅ fecundi- Ｐ ｘｕthiｔｓwas slightly smaller than that in the ties, which were obtained from the total number of summer generation (P<0 01), while such a differ- eggs, of both spring and summer generations ｍ /） ence was not found m 7）ｍａｃｈａｏｎ ｘzjがz£iswere at 570-600, suggesting that the small The change ｍ the number of eggs found ｍ the body of spring generation may affect not the abdomen of both species are shown ｍ Table ｌ fecundity but the number and the mature egg size The numbers of submature, immature and total ｍ the ovaries eggs decreased with the age except the summer In P machaon. the number of mature eggs of generation of Ｐ ｍａｃｈａｏｎ However, the number spring generation was smaller than that of summer of mature eggs seemed not to decrease with the generation ｍ age Ｏ（005＜Ｐ＜01）ａｎｄ age ｌ（Ｐ＜

age ｍ all the generations 0 005) Although the numbers of submature eggs InP ｘｕthｕｓ,the numbers of mature eggs ｍ age were significantly different between both genera- ｏ to Ill of the spnng generation were smaller than tions in ages O，l and IV, those of immature egg ｍ those of summer generation （Ｐ＜001）Ｈｏｗｅｖｅr， both were not significantly different each other the numbers of submature and immature eggs ｍ with probabilities less than 0 05 Thus, the fecun- age ｏ to II were not significantly different between dities of Ｐ ｍａｃｈａｏｎ１ｎboth generations seemed to

Table1 Number of eggs in ？ xuthus and P machaon in relationto the age (Mean士ＳＥ） くspringgeneration〉

Age ０ Ｉ II Ill IV - P xuがTUS No females examined 17 19 15 8 5 No mature eggs 178±４９ 256±４８ 239± ３５ 178士 ３３ 160土 ９３ No submature eggs ５４９±62 ５５５±68 314± ７６ １４１士 18 ８６± １９ No immature eggs 525 6±497 443 8±412 280 3± ３８２ 186 0±416 128 0± １９４ No totaleggs 602 9±510 524 8±407 335 6土 ４４８ 217 9±416 152 6土 １９９

Ｐ ｍａｃhａｏｎ No females examined 6 8 5 1 2 No mature eggs 435±119 474±６８ 432土 ６６ (26) 135土 ６５ No submature eggs 380±59 ２２４±64 ３２６士 60 09) ８５士 25 No immature eggs 394 7±315 4075±510 428 8±110 0 (136) 244 0±148 0 No totaleggs 476 2±197 477 3±522 504､6±119 2 (181) 266 0±157 0

くSummer generation〉

Age 0 I II 111 IV

？ＸＭがz附 No females examined 37 64 50 30 24 No mature eggs 468±４６ 583±４０ 474土 ３４ 369士 ３２ 247土 ３７ No submature eggs ４８４±33 ４８０±36 389± ３０ 280± ２４ 172± １６ No immature eggs 475 0±214 460 2±197 367 7± ２１８ 359 8± ３５２ 274 5± ２６４ No total eggs 569 7±214 S67 1±172 453 9土 ２１８ 426 0土 ３５９ 316 5士 ２８９

Ｐ ｍａｃｈａｏｎ Nofemales examined 15 18 6 3 ２ No mature eggs 759±９２ 887±６６ 402士 ９９ 660±285 ７２ 5±235 No submature eggs ４８７±38 ５７３±85 362± ３９ 423±123 480士 20 No immature eggs 500 8±312 480 1±325 362 0± ４３３ 3917±130 1 538 0±234 0 No totaleggs 626 2±307 626 1±326 438 3土 ４２２ 500 0±140 4 658 5±208 5 -

Fecundity of Swallowtail Butterflies 513 ①∧几 1 1 1 1 1 ににt 1 1 二 １ １ １ １ 】 9 11 13 15 17 9 11 13 15 17 9 11 13 15 17 ９ 13 15 17 h「 0 1 11 m・IV

Fig ５ Change ｍ mean number of mature eggs ｍ ａ day of 7）ｘｕthｕｓof respective ages ｍ spring generation （士SE）Ｔｈｅ time means ａ median of ２hr, ｅｇ ｇ means from 8 00 to 10 00，11 does from 10 00 to 12 00 and so on The number of females examined ｍ each age was 17，19，15 and 12 1n age O，I，II and III十IV, respectively

８ ｛り ４ ０ In Ｐ ｘｕthｕｓof the spring generation, as shown ｍ Figure 5，the number of mature eggs decreased toward evening ｍ all age classes Steep declina-

6６ｄ tion was found in 11-13, 13-15, 9-11 and 11-13 m age O， I，IT and III十IV, respectively Thus, the ｓ」コleai ｏｎ females seemed to deposit eggs mainly at noon The mean number of eggs deposited ｍ ａ day was estimated by the difference of the egg number between morning and evening 30, 25， 20 and 20 by

二 ａ female of age Ｏ，I，IT and Ill + IV, respectively 9 11 13 15 17 9 11 13 15 17L｡＿ｔ 1 1 」 h「 Because of the relatively small number of the Ｏ・１ II- IV females collected in the spring generation of Fig ６ Change ｍ mean number of mature eggs ｍａ day Ｐ ｍａｃｈａｏｎ，Figure 6 shows the change ｍ numbers ofP mαｃ/zαθzlof respective ages ｍ spring genera- tion(土SE）Ｔｈｅ number of females examined ｍ of mature eggs ｍ two pooled ages ｍ ａ day The each age was 14 and 8 m age O-I and II-IV, oviposition time ｍ younger females was not respectively detected, while that ｍ the older ones were considered to deposit about 30 eggs ｍ the be similar at 500-600, suggesting that the small morning body of spring generation may also affect not the In the summer generation of Ｐ ｘｕthｕｓ，the fecundity but the number of mature eggs ｍ the younger females had clear tendency that they ovaries deposited eggs mainly in the morning (Fig 7) Females of the oldest age (IV)ｍ both species However, the older females had obscure one, and had ａ considerable number of eggs, which mamly they seemed to deposit fewer eggs than the consisted of submature and immature eggs Since younger females. It might be estimated that the such female was not considered to deposit all of females of Ｐ ｘｕtｈｕｓ11n summer generation depos- her eggs remained, the female seemed to have lted 25, 25， 25， 15 and 20, in age O, I, n, Ill and IV,

more eggs than actually deposited respectively, ｍ ａ day In order to estimate the number of eggs depos- In the summer generation of Ｐ ｍａｃｈｏｏｎ,the lted in ａday, we took no account of the number of obscure oviposition time was also found as ｍ the eggs developed from submature to mature ｍ ａ case of the spring generation (Fig 8)Ｔｈｅ mean daytime When the females were classifiedm each number of eggs deposited in ａ day was 25 and 30 1n two-hour intervals according to the time of cap- younger and older females, respectively ture, the time of decrease ｍ mature egg number may be the oviposition time 514 Ｍ Watanabe AND Ｋ NOZATO

９σｅ 一」コ‘ＩＥ

ｊ ソゾソ①

１ １ １ １ １ 二 1 1 1 1 1 ｌ ｌ １ -よ 」 二 ９ １ 13 15 17 9 11 13 15 17 9 11 13 15 17 9 11 13 15 17 9 11 1S 15 17h「 ｍ ０ 1 Ｈ

Fig ７ Change ｍ mean number of mature eggs ｍ ａ day of Ｐ ｘｕthｕｓofrespective ages ｍ summer generation (土SE）Ｔｈｅ number of females examined in each age was 32，63，48，30 and 24 m age O， I,II, III and IV, respectively

20 90 60 30 ０ fertilizeall of eggs Thus, the difference ｍ the number of matings ｍ ？ xuthus and Ｐ ｍａｃｈａｏｎ suggests that each has Its own characteristicrepro- ductive strategy Further analysis may be neces- sary for comparison between these two species As shown ｍ the Result, there IS no difference ｍ bｂｄ ｓjコｌＥＬＵ survival rates with relation to body sizes between two species The female size seemed not also to influence the number of eggs ｍ the ovaries The same has been observed in the black swallowtail Ｏｚ butterflies(Watanabe et al , unpublished) Number of deposited eggs decreased with age ｍ Ｐ ｘｕthｕｓ Theestimated number of eggs depos- lted in ａ day by ａ female was larger than that L_ j l l l 卜白子干ヨ7 9 11 13 15 17 h「 observed ｍ ａlaboratory condition ［1］However, Ｏ－１ II - IV Yamanaka et α/［3］proposed about 40 eggs Fig ８ Change ｍ mean number of mature eggs ｍ ａday actually deposited by ａ female of Ｐ ｘｕthｕｓina ofP machaon of respective ages ｍ summer genera‘ day, but they did not show any age structure of the tion(土SE）Ｔｈｅ number of females examined ｍ each age was 33 and 10 1n age O-I and II-IV, females sampled respectively It has been stillunknown when the number of mature eggs recovered after oviposition In 7） ｘｕthｕｓofsummer generation, the younger females DISCUSSION clearly showed oviposition time ｍ the morning to It IS clear that many females of 7）ｘｕthｕｓmated noon, while the older ones did not show ａ more than once with age, while that of Ｐ ｍａｃｈａｏｎ particular oviposition pattern Figure 7 suggests did once Ａ number of recent studies show the that the recovery of the mature egg number had multiple matings ｍ many butterfly species such as began from late-afternoon or evening Although the checkerspot butterfly， Ｅｕphｙｄｒｙａｓｅｄｉtha［81 the egg maturation during evening and night was The role of multiple matings m butterfly species not clarified,the diurnal oviposition activity pat- has been discussed on sperm competition ［9－10］ tern disappeared with age In the spring genera- and on energy for females ［11］In general, ａ tion, on the other hand, the oviposition time single insemination may be usually sufficient to shifted to noon instead of morning Lower -一一-

Fecundity of Swallowtail Butterflies 515

temperature ｍ the spring than the summer may Maeda, Department of Biology, Faculty of Education affect her diurnal oviposition activity This Mie University,for theirassistanceｍ the field tendency may probably be similar to that ｍ Ｐ ｍａｃｈａｏｎ REFERENCES Few comparative studies between spring and Ｉ Tsubaki, Ｙ （1973）Ｔｈｅ natural mortality and Its

summer generations of the swallowtail butterflies factors of the immature stages ｍ the population of

were reported Badawi [12]observed longer and the swallow-tail butterfly Ｐ叩山ｏ ｘｕthｕｓLinne Jpn Ｊ Ecol , 23 210-217 shorter longevities of 7）ｄｅｍｏｌｅｕｓｍwinter and 2 Watanabe, Ｍ （1981) Population dynamics of the summer generations, respectively In this study, swallowtail butterfly, 7）aoilio ｘｕthｕｓL , in ａ defor- the fecundity ｍ each species of spring generations ested area Res Popul Ecol ， 23 74-93 was similar with that ｍ summer generations, ３ Yamanaka, Ｍ ， Suzuki, Ｙ and Takagi, Ｍ （1978）

though their eggs were relatively smaller than Estimation of egg number oviposited per day by ａ

those of summer generations Wiklund and Pers- female of ｊ）ＯＤｌＵｏｘｕthｕｓLmne (Lepidoptera, Papi- son 113]clarified that smaller eggs were not ｍ any honidae) Kontyu, 46 329-334 way inferior to larger eggs within ａ same butterfly 4 Dempster, J Ｐ and Hall, M L (1980）Ａｎ at- tempt at re-establishing the swallowtail butterfly species, /)αΓargeaりμ?ｒlα at Wicken Fen Ecol Entomol ， 5 327-334 It IS still unknown whether the longevities of 5 Watanabe, Ｍ ， Nozato, Ｋ ， Kintani, Ｋ and Miyai,

females are similar with those of the conspecific S（1984） Seasonal fluctuations of egg density and

males According to Abu Ｙａｍａｎ[14], adult survival rate ｍ Japanese black swallowtail butter- females of P demodocus survived for 7-12 days flies Jpn J Ecol ，34 271-279 and males for 5-10 days The males of the 6 Nozato, Ｋ ， Kintani, Ｋ ， Miyai, Ｓ and Ban, Ｙ （1985） Studies on ecology and behavior of Japanese black swallowtail butterflies also survived Japanese black swallowtail butterflies IV Estima- for 13－18 days [6]Thus, it IS likely to assume that tion of life span of male ７）ａｏｉｌｉｏｈｅｌｅｎｕｓｎｉｃｃｏｎｉ-

the longevities of the adult swallowtail butterflies ｃｏｌｅｎｓButler and j）ρΓotenθΓdemetrius Cramer

were about two weeks If the longevity coincides (Lepidoptera Papilionidae) Appl Entomol Zool ， 20 494-496 with the duration of oviposition, P ｘｕthｕｓwould 7 Hidaka, Ｔ and Aida, S （1963）Ｄａy length as the deposit 345 and 330 eggs ｍ spring and summer mam factor of seasonal form determination ｍ generations and P machaondid 450 and 420 eggs, Ｐｏｌｙｇｏｍａ ｃ-ａｕｒｅｕｍ(Lepidoptera, Nymphalidae)

respectively, by multiplying the estimated egg Zool Mag ， 72 77-83

number deposited ａ day by the duration of each 8 Labine, Ｐ Ａ （1966） The population biology of age (about 3 days ｍ each) However, there must the butterfly， Ｅｕphｙｄｒｙａｓ ｅｄｉtha IV Sperm precedence―a preliminary report Evolution, 20 be some bad weather days during their oviposition 580-586 periods, such as rainy, windy and cool cloudy, ａ11 9 Parker, Ｇ Ａ （1970) Sperm competition and Its ０f which inhibit females from oviposition Then, evolutionary consequences ｍ the insects Biol

about 300－400 eggs were estimated to be actually Rev ， 45 525-567

deposited by the spring and summer generations of 10 Rutowski･Ｒ Ｌ （1982) Epigamic selection by both species This was considered to be ａ so-called males as evidenced by courtship partner preferences realized fecundity Thus, such value was a half of m the checkered white butterfly（乃ens /7Γotodice) the fecundity in P xuがtus and Ｐ ｍａｃｈａｏｎ Anim Behav ，30 108-112 11 Boggs, Ｃ Ｌ and Gilbert, ＬＥ（1979) Male con- tribution to egg production ｍ butterflies evidence for transfer of nutrients at mating Science ２０６ ACKNOWLEDGMENTS ８３－84 ’

We thank to Dr K Kintani, Division of Entomology, ^^ Badawi･Ａ（1981) Studies on some aspects of the National Institute of Agro-Environmental Sciences, Dr biology and ecology of the citrus butterfly PaD111０ N Suzuki, Department of Biology, Faculty of Scienc。 ｄｅｍｏｌｅｕｓＬ ｍSaudi Arabia (Papilionidae･ Lepi- Kyushu University and Dr Ｃ Wiklund, Department of doptera) Z ang Entomol ・91 286-292 Zoology, University of Stockholm for their critical 13 Wiklund･ Ｃ and Persson, Ａ （1983) Fecundity, reading Thanks are also due to Mr Ｍ Taguchi, Hashi- and the relation of egg weight variation to offspring moto High School, Mr Ｋ Kobayashi and Miss T fitness ｍ the speckled wood butterfly /）ａｒａｒｇｅ 516 Ｍ Watanabe AND Ｋ NOZATO

αど1μ?na,or why don't butterflyfemale lay more citrus leaf caterpillar, PａＤｉｌｉｏ ｄｅｍｏｄｏｃｕｓ Esp eggsりOikos, 40 53－63 (Lepid ， Papihonidae) in Saudi Arabia Ｚ ang 14 Abu Yaman, IK（1973) Biologicalstudieson the Ｅｎtｏｍｏｌﾌﾟﾌ１ 7n(s-383