A gynandromorph of Papilio memnon agenor Linnaeus (Papilionidae) from the Lao P.D.R.

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TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
uetwh 71rans.tepidS.oc,Japan 57<2):105-1IO,March2006

A gynandromorph of I)tipi mleimnoon agenor Linnaeus (Papilioni fdraoem) the
Lao RD.R.

Masayuki NiMuRA and Hiroki OTANi
Departmcn tofDcvelopmental Biology F,acult yof Medicine ,Shimane Univcrsity ,Izumo, 693-8501 Japan
Abstract We report here a gynandromorphic specimcn of Papitio tnemnon from the Lao P.D.R.
The right wings in this specimen principall yindicat enormal male color pattern ,whereas the left
wings show a mosaic patter nwith the normal male and femal e(for mdistantian ucosl)or patterns.
The wing shape of this specimen is not similar to thosc of normal male and femal e(E dislaniianus)
specimens, and the size is relatively small in comparison with those of norma[ male and female
specimens. This specimen has female (f .di,stantianu sex)ternal feature sin the right half abdomen,
while male featur eisn the lef t,Although the genit aslegments show the externa] feature sof those
of a normal male, an egg is attached to it.
Key words Gynandromorph, wing mosaic, Pcu)ili moemnon, Lao P.D.R,
Ptipili omemnon Linnaeus (Papilion iids waideel)y distribut eacdross Southeas tAsia, and is
one of the commonest species in the Lao RD,R, lt is sexually dimorphic everywhere, and
the female sare polymorphic i,ncludin gthe non-mimetic form agenor and the mimetic form
distantian ti{n sthe Lao P.D.R (Osad eat at., 1999) ,Gynandromorphic specimens of P.
memnon are very interesti bnegcause of sexual dimorphism ,but there have been only a few
reports, e. g. from Sabah, Malaysia (Miyauch i1,993) and Okinawa, Japan (Ishiz a1k99i5,),
In this report, the structural characteristics of an intersexu amolsaic of Ptipil imeomnon
male and female (f d.istantianu cso)llected in Laos are describe d.This specimen was col-
lectedby Ms SonephetWakahara Mr HiroyukiWakaharainVientianeL,ao RD,R, in
and
VI, 1999 andpassed by them to the author (M, N,),
We numbered the designati oonf veins and spaces according to Sibatan (i1980).
A brief summary
On the dorsal and ventral surfaces of the right wings in this specimen, both fore -and hind-
wings indicate normal male color patter nw,hereas on the dorsa land ventral surfaces of the
lef twings, both fore -and hindwings show a mosaic of normal male and female (f .distan-
tianus) color pattern s(Fig s1A, B). The wing shape of thi sspecimen is not similar to those
of the normal male and female (f .distantianu cso)llected at the samc localit yin Laos by the
author (M, N.), and the size is relatively small in comparison with those of normal speci-
mens (Fig sIA, B). Both lef tand right forewing sare 61 mm in length ,while those of nor-
mal males and temale s(f .distantianu asr)e 65-68 mm (n=2 )an,d 70-76 mm (n=4 )re,spec-
tively. The right side of the abdomen is pale cream in color lik efemales ,while the left side
is dark brown like males, respectively (Fig .IB-2). Although the genita lsegments show
normal male feature esxternally, an egg is attached (Fi gI.B-2, arrowheads). The genital
segments also slightly twist to the right (Fi gI,B-2).
The leftforewing

The wing shape and venation show the same as either male or female (Fig 2sA-F). The
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106 Masayuki NiDviuR Aand Hiroki O'rANi
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Fig .1.A gynandromorphic specimen of Papilio memnen agenor Linnaeus, 1758. A, Dorsal sur-
face ,B. Ventral surface, A-2. A magnified view of the boxed urea in A. The bright-blue
scales are locallz cond the wing surfaces along by the vcnations (CuA, )in the tnale areas as
in normal niales (arrowhea dsB-)2,. A magnified view of the boxed area in B, The right
sidc of abdomen is pale creani in color lik ethose of females ,while the lef tside of it is dark
brown lik ethose of male respectively, Although the genita lsegments show the external
featur eost' those of iiormal rriulcs, an egg is attached on i t(arrowhea dM,)], vein Mi; CuAi:
vein CuAi.
ratio of forewing ,outer marginal (ti opf forewing to end of vein 1A+2A), and dorsum (vein
1 A+2A) length sbetween the gynandromorph ,and male and female specimens col]ected at
the same locali tarye 2.02: 1.36: 1, 2.01 :l,37: l and2.05: 1.43: 1, respectjvely,
1. Dorsal surface
A boundary lin efrom M, to the base (continu iinngt othe disca lcell) on the wing subdivides
the dorsa ]surface int otwo halves ,The costal side of the two halve sshows male color pat-
tern (Fig s2A, C), whereas the dorsa lside shows mainly female (Fig 2sA, B) andanumber
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A Gynandromorph of Papitio memnon 107
of (approxima t12e) lunyconnected patches of male patter nrandomly spread across the area
(Fi g2.A). Although the bright-bl usecales are localize odn the wing surfaces along the
veins in the male area as in nermal males (Fi g2,C), the number of scales is smaller than in
(Fig1.A-2,
norma] males arrowheads).
2. Ventral surface
As on the dorsa lsurface, a boundary lin efrom M, to the base (continu iinntog the discal
cell) on the wing subdivides the ventral surface int otwo halves .The costal side of the two
halves shows mainly male color pattern (sFig s2D, F), and a number of (approximat e1l5y)
unconnected patche sof female patter nrandornly spread across the area (Fig s2D, E),
whereas the dorsa lside shows mainly male color patter n(Fig s2D, F). This is the opposite
situation to that on the dorsa lsurface (Fig 2sA, D).
Sibatan i(1987 h)as been analyzing spontaneous wing homoeosis in Lepidopter aand he has
recorded more than 350 cases, and noted that they are generally confined to either the ante-
rior or posterio hralf (putat icvomepartments) of a wing surface, with a boundary fallin ogn
the space M,-M2 and continuing basad into the discal cell towards the base on both wings
(Sibat a1n9i80,, 1987). The presen tresults for the lef tforewing are practicat lcoynsistent
with those of Sibatani.
The lefthindwing

The wing shape is mainly similar to that of the female (Fi g2sG-L). However, outer margin
from M2 to Sc+R shows a male-like pattern ,whereas from M, to 1A+2A a female-lik epat-
tern (Figs2G-L),

1. Dorsal surface
A boundary line from M, to the base (exce pfotr the discal cell) on the wing subdivides the
dorsa lsurface into two haive s(Fi g2.G), The costal side of the two halves includin gthe
disca lcel] shows male color pattei n(Fig 2sG, I) ,whereas the dorsa lside shows mainly fe-
male color pattem and a number ef (approxima 1t8e) lunyconnected patches of male pattern
randomly spread across the area (Fig s2G-I) ,The tail is incomplet esince the size is smaller
than in normal female specimens collected at the game localit y(th elengt hand width of the
tai lon the gynadromorph is shoTter and narrower than in normal female specimens) <Figs
2G, H). As in the forewing ,the bright-bl usecales are localize don the wing surfaces along
the veins in the male area as in normal males. However, hair sseen on the basal area of nor-
mal male hindwing are hardly detecte don this specjmen.
2, Ventral surface
The ventral surface shows equally a number of unconnected patche sof female and male
color patterns (approxima t49e.l8 y% for female areas) (Fig 2sJ-L). In the spaces Sc+R-Rs,
Rs-Mi, M2-M3, M3-CuAi, CuAi-CuA2, and disca lcell, the area consisting of orange colored
scales, neither of males nor females, are detecte dat the border of pale colored patche sof
female pattern (sFi g2.J, arrowheads),
The forewing
right
Both dorsal and ventral wing surfaces show the male color pattern swith the bright-blue
scales (Fig s3A-F), However, the dorsal surface is differen tfrom typical males and shows
Ies sdark brown color (Fig 3sA, C), The ventral surface shows almost normal male color
pattern (sFi g3.D-F). The wing shape is slightly differe nftror nboth male and female speci-
mens collected at the same locali t(yFig 3sA-F). The ratio of forewing, outer marginal (tip
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108 Masayuki NiMuRA and Hiroki OTANi
Ml MIF
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Fig. 2. The lef twings of gynandromorph (A, D, G, J) ,normal femal e(f d,istantianus )(B ,E, H, K),
and normal rnale (C, E I ,L) specimens fi'o mthe same locali tyA-.C. Dorsal surfaces of the
lef tforewings on gynandromorph (A) ,norrnal fernal e(B) ,and normal male (C) .D-F.
Ventra lsurfaces of the lef tforewings oii gynandromorph (D) ,nornial femal e(E) ,and nornial
male {F) .G-l. Dorsal surl'aces of the lef htindwing son gynandromorp h(G) ,normal female
(H) ,and nonnal male CI) .J-L, Ventra lsurfaccs of the lef thindwings on gynandromorph
(J) n,ormal femal e(K) ,and normal male (L) .J, arrowheads. Areas that censist of orange
colored scales, neitber of males nor female s,Sc+R: vein Sc+R; Rs: vcin Rs; M/: vein Mi;
M2: vein M]; Mi: vein M" CuA,: vein CuA,; CuA,: vein CuA,; D: discal cell; T: tail.
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A Gynandremorph of Papitio memnon 109
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Fig. 3.The right wings of gynandromorp h(A ,D, G, J) ,normal female (f .distantian"s )(B ,E, H,
K), and normal male (C ,E I, L) specimens from the samc locali tyA.-C. Dorsa lsurfaces of
the right fbrewings en gynandromorph (A) ,norrnal femal e(B) ,and normal male (C) ,D-F.
Ventral surfaces of the right forewings on gynandromerph (D) ,normal female (E) ,and nor-
mal male (F) .G-I. Dorsa lsurfaces of the righ thindwing son gynandromorph (G) ,norrnal
female (H) ,and nernia] male (I )J,-L. Ventra lsurfaces of the right hindwtngs on gynandro-
morph (J) n,ormal fema]e (K) ,and normal male (L) .T: tail,
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110 Masayuki　N−・tuRA 　and　Hiroki　OTANI
of　forewing　to　vein 　l　A ＋2A ），　and 　dorsum（vein 　IA＋2A）length　sbetween　gynandエomorph
，
male ，　and 　fbmale　spccimens 　are　1．9：1．25：1，2．02：1．38：1alld　2．08：1．42：1，respectively ，
The hindwing
　right 　
Both　dorsa　land　ventral 　wing 　su血 ces　shQw 　the　typical　male 　color 　pattern　wsith 　the　brigh−t
blue　scales （Figs　3G−L）．　As　the　right 　forewing，　however　the　ground　color 　of　wing 　surfaces
，
is　les　sdark　brown．（Figs　3G 1　J　L）．　 The　wing 　shape 　is　morc 　silnilar　to　that　of　female
specimens （except　for　tai）lcol，lec，ted， 　at　the　sarne 　locali　ttyhan　to　Inale 　specimens （Figs　3G−
L）．
Acknowledgments
We　deep］y　thank　Dr　Kazuo　Watanabe（Hiroshima　University）for　valuable 　advice ，　and　Mr
Hiroyuki　Wakahara　and 　Ms　Sonephet　Wakahara（Lao　RDR ．，　Vienti且ne ），　who 　kindl　ygave
me 　the　gynandromorphic 　specimen 　of　P．　memnon 　from　Lao ．　P．D ，R，
References
Ishizak，　iT．，1995．　 Nagasaki−ageha 　no　seimozaiku （Gynandromorphi　cmosaic 　I’apilir丿mentnon ），　 Choken
　　Fieid・10（ll6）：18（jnJapanes）e．
Miyauchi，　N．　and 　M ，　Harada，1993，　Gynandromorph 　of　Papiti　o」nemnoft 　and 　Pa配翩 ca　pedonga．　Buttenyflies
　　5：57（in　Japanes）e．
Osada，　S，，　Uemura ，　Y ，　and 　J．　Uehara，1999．　An　ilrustra　tCehdeckli　sotf　the　Buttenyfl　iOesf　Laos　P，D ．R，240
　　pp，，143　pls，　Mokuyo−sha ，　Tokyo ．
Sibatani，A ．，198（1．　Wing　homDeosis　in　Lepidoptera：asurvey ，ρ8vψρ．　Biot．79：1−18，
　　　　，1987．Homoeosis　of　dorsa　land 　ventral 　wing 　surfaces 　in　bulternie．s ．1．　tiber、　Arts　Kansai 　med ，
　　 〔ノniソ．11：11−12．
｝商 要
・
ラオス産ナガサキアゲハ 雌雄モ ザイクの記録 （二村正之 大谷　浩）
ラオス産ナガサキアゲハ の雌雄モザイクを記録 した．本雌雄型個体は 1999年6月，ビエ ンチャ ン市内に
て若原ソン，若原弘之両氏によっ て採集されたものである．前翅長は左右ともに61mm で あ り，同地
域から採集された正常雌 （70−76　mm ，　i・t＝4）および雄 （65−68　mm ，　n＝2）と比べ て小型である，
本個体の右前翅および後翅は背側 （表）．腹側 （裏）面 ともに概ね雄の斑紋を示すが，左前翅及び後翅は
両側面ともに雌雄モ ザイクを示す，左前翅には両側面にM1 脈から中室 を貰い て基部とを結ぶ境界が
認め られる，背側面はその線を境に前縁側が雄の 後縁側が雄と雌とのモザイクを，腹側面は前．縁側
，
が雄 と雌のモ ザイク を 後縁側が雄の斑紋をおのおの示す．また雄の斑紋部分には青白色の鱗粉が散在
，
してい る．翅形は前翅が左右ともに正常個体との差はほとんど認め られないが，後翅は雄と雌のい ずれ
とも異なっ てい る，
（A¢cepted 　Novcmber 　15 2005）
，
Publjshed　by　the　Lepidopterologi　Scoacliet　yof　Japan，
5−20，Motoyokoyama 2、HachiojiTokyo 、192−0063 Japan
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