Flight behavior of Luehdorfia japonica (Lepidoptera, Papilionidae) at the summit area of Mt Egesan, Hiroshima City : 1. Hill-topping and round-patrolling of males and their roles in mating and meta-population convergence

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ecLwt 7'rans.Iqpid.Soe.Japan 57 (3)2:37-254,June2006

Fligh tbehavio rot' Luehdoi:fiaJ'trpon i(cLaepidopte Praap,ilionidae )at the summit
Mt Egesan,Hiroshima City
area of
1. Hill-toppi nangd round-patrolling of males and their roles in mating and
meta-population convergence
Kazuo WATANABE[)! /a'nd* Kazuhiko HiRANo2'
L' Facult yof Integrat eAdrts and Science sH,iroshima University,
Kagamiyama 1-7-1,Higashi-Hiroshima,Hiroshima,739-8S21Japan

!' Graduat eSchool of Biosphere Sciencc, Hiroshima University,
Kagamiyama 1-7-1,Higashi-Hiroshima,Hiroshimu, 739-8521 Japan

Abstract We marked 29 individual s(27 oA 2 ?) of Luehdoclia japoni c(aLeech i)n 2002 at the
summit of Mt Egesan .and analysed their subscquent flig hbtehayior sT.he lif espan at the peak was
estimated as 15-17 days. We describe the diurnal changes of the flig httrack sand discus tshei rim-
plication sfor mating and population convcrgcnce. The flig hptatter 'dn`eryoeu]nodp- sipna ttirmoel. liAnftge"r mak-
ing a short random `"wandering flight "in the morning, a fiyway of with a few
preferenti asltaying spots develops. On certain occasions, they manifest a "perching occupation': at
a specific staying gpot, During thc reund-patrolling, [`spira]ing flight 'ii sevokcd when encountering
another ma!e individua lof the same species, And, if the cncounter is a virgin ferna[ eth,ey copulatc.
Thc fligh tactivities of each individua lat the peak seem to be individua lslpeycitic, and generally
cover a wider area with aging. During the survcy, cvidence of peak-to-peak ruund trips (a tleast
680 m fligh twithin four hours )was obtained. These behavior osf male tndividua plrsoduce a wide-
spread coverage of the peak area by a scattered distributi oonf t'einale-seeki nmagles. This un-
doubtedly is an effective strategy for lowerin gthe probabilit yof female sremaining virgin in the
meta-population.
Key wurds LuehdofVia ,mcta-populution, fligh tbehavior ,hill-toppii irgou,nd patro!lin gt,enitory,
mating, population divergenc peo,pulati ocnonvergence,
Introductien
Many papilion ibdutterfli emsanifest L`hill-topping'7 (Shiel d19s6,7) in their fligh tactivities,
In Luehdonjia ,iaponica, although hill-toppin ghas been well-recognized (Fukud aer at.,
1982), and fligh tbehavior of the species has been systematically describe din a few mono-
graphs (Hiruka w1a9,88; Y Watanabe, 1996; K, Watanabe, 1998) ,we can seldom find de-
tailed analyses of fiigh tbehavior, except for a few data obtained by the mark-recapture
method (Matsumo t1o9,84, 1994; Natsuaki ,1989, 1996; Natsuak iand [Ilakeuc h1i9,99) .In
a specialized fiel dof Kanazawa City ,Centra lJapan ,Matsumoto (1984 ,1994) showed pop-
ulation analyses, male patrolli nbgehavior a,nd several patchy accumulations of individuals
along the ridge ,and proved eight cases of long-distan cfeligh (t526-1, 5m9)6 of marked in-
dividual sI.n field osfOsaka Prefectur eNa,tsuaki (1996 )and Natsuak iand Takeuchi (1999)
proved that males emerged at the bottem of mountains and repeatedly appeared at hil1-tops.
Also ,they observed copulation at hill-to pssu,ggesting that hill-toppi insg a kind of mating
strategy by males to meet females.
'Corresponding author. PrescnL address: 5-9-20 ,Midori ,Minamiku, Hiroshima 7,34-OO05, Japan.
E-mail:[email protected]

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238 Kazuo WA'vANABE and Kazuhiko HmANo
To understand the significance of adu]t fli'g bhethavio rmore pre¢isel yv,v'e need to know the
totality of behavior at the hM-tops by marking most of the flyin gindividuals ,recording
their fligh tactivities, and attempting analytical categorization, The problems to bc ex-
plaine dare how they meet in order to copulate and how they find thei rfood plant ,while at
the same time avoiding the dans,e rof populatio dnivergence. Tb accomplish this ,it is nec-
essary to select a proper examination field ,having an adequate land-struct uwrheere the
Luehdo]:tia-butter cfalni edsispla yrestricted fiigh tbehavio rwithout sutfering any interfer-
ence with other spccies; this enables us to make strict fligh otbservations and analyses. Mt
Egesan of Hiroshima City was considered to satisf}J the condjtions.
Materials and methods
1 . Luehdot:fi aj'qponica at Mt Egesan
1-1.CharacterizationMt Egesan habita(tFig,1)
of as a
A half ¢entury ago, the summit area of Mt Egcsan (mai npeak; alt. 582.3 m) was known as
a habita tfor LuehdoT:t 1i'aaponic ah:owever the butterfl iweerse absent or quit erare for 30
years and final lreycovercd in approximately 1990. Such dynamic changes in a smal1 meta-
populatie nseem to occur naturally in many habita tisn the Chugoku distri c(tK. Watanabe,
1991) .Comparably stable meta-populatiens still exist, about eight krn southeast from here.
However, at the same time, an artificial transplantation of the species by anonymous per-
sons is also not completely rulecl out. In any case, data in this manuscript indicat etshat the
populatio ins quit esmall and isolated.
Since 1980, the peak area above the contour of 500 m has been set up as a municipal
Fig ,1.Mt Egesan as a habitat .(a )Image ot' distribut iaroean of the food plant ,Asarum hexaiobum
(grce n)an.d thc cumulative ovjposition sites (re dpoints )durin gthe las t15 years
<1990-2005 )by the authors and co-worker, Takeshi Kameyama. The oviposition points
move from year to ycar, and have been quitc restrie('ed recently. Peak A i sthe major survcy
poinL The sma[1 arrow at Peak D indicate sthe recaptured point of markcd individu aNlo.
'I'he
21 on April 19, 2002, (b )Hiroshima City is helow Peak A. arrow indieate sthe atomic
bomb target, 12.4 km northwest from the Peak A.
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Fig. 2. Cap-like structure of the summit of Peak .A Lwirh a detscripti oofn detail efdlor aT.he bluc
area is bushes of clouded bhrubs, where the flyin Lgttehdocfiajqpon inecvear enter. Thc
yellow area is brigh tbushcs -･ith spotted semi-opcn spaces where the butterfli eocscasionally
fi yabout (se eFig 7, upper left) .The green area is a dricd grassland ,where thc butterf diey
not like to fly ubout. The centra] hexagon is an arbor established by Hiroshima City Othce.
The ground fioor surrounding the arbor (uppe trhan alt .566 m) is constantly cleaned up by
keepers of the municipa] purk. Numerals in the symbol marks of trcc mean the height (m)
of the tree ,The eastern half of the dome area Chatehcd) is the main tligh tarea. which is
covered with brown-color efdallc lneave sof pine trees .The inserte dphotograph shows the
harche darea and the clouded ghrubs from the arbor.
Fig, 3, Markings of Lttehdotzfiajaponica.
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240 Kazuo W,xTixNAB aEnd Kazuhiko HiRixNo
Egesan park of Hiroshima Clty ,and has experienced continuous artiticial manipulation of
the flor aas a "managed forest" .The fbod plant and 1ai deggs are distribut meadinly in and
near the manipulated area (Fig .1) ,beeause the base area, lower than 500 m, is generally
covered with untreated thicketed bush, where the food-plan Atsarum (Heterotf therxpaa-)
lobum E Maekawa does not propagate well and where adult Luehdt)i:17a-butte crafnnloties
easily invade due to thei rHight habits .Although there were wide-spread distributi oonfs
Luehdoiji ameta-populations in the vicjnities-a few or several kilometer dsistan tfiro mMt
Egesan-until 20 years ago (K, Watanabe et al,, 2000), most of the habita twas destroyed
during the recent 15 years by development of building-l oatnds destructio nof the circum-
stances character of the area.
These fact sstrong]y suggest that tbe collection of butterfl iaends eggs in these areas would
bring about serious damage to the Luehdoi:fi ameta-populatien. Recently ,we, along with
inhabitant sand municipal officers, started a projec tto conserve and propagate Luehdot:tia
.1'aponic ain Egesan Municipal Park (t obe published s)o that the collection of the butterflies
and food plant scan be avoided at least until recovery and stable maintenance of a good
Lttehdoijia-popula tiis orenalized.
1-2 .Characterizat ioofn the meta-population
The meta-population is a typical peri-Hiroshima Bay populatio nwhich uses Asaruni hexa-
tob"m asasole food plant (K .Watanabe, 1991; 1998), Mt Egesan is locate don the border
of the two sub-metapopulations, which dilfe rin their emerging periods ;one is eastern late-
emerging which seems to manifest fiv eto seven days late remergence every spring, and the
other is northwestern early-emerging (Hashimo t19o9,1; K. Watanabe, 1998).
By analysing base sequences of ca 800 nucleotides of the mitochondrial ND5 genes of 10
Mt Egesan individua l(sdat naot shown), the populatio nmanifested typica l"Hiroshima-
[ibtto rtiype", which showed a specific SNP (sing lnuecleotide polymorphism )at the base of
107th, that is ,a co-existence of 107-A type and 107-G t>Jp ein the same meta-population
(Hirowat aandr iWatanabe, 2000).
2, Detailed descriptio nof the survey point (Pea Ak)
The summit of Peak A (al t5.68.8 m) was a cap-like open land (ca 50 m × 30 m) with flat
managed ground (Fig 2.), The western halfwas open grass land and the eastern ha]fwas a
sparse woody area of pine trees (Pinu dsensijlor aan)d plante dPrttnus ),edoensis ,with a
hexagona larbor in the center (Fi g2.). The situation made it possibl eto watch and record
multiple flyin gindividua lats the same time. In this season there were no flyin gbutterflies
in the area other than Lttehdoijiajaponica.
In the summer of 2004, the peak area was completely destroye dat a plane of alt .560 m-
contour by publi cconstruction of a radio wave tower (121. 0m)3 with an adjacent four-
story building for digital-TV broadcasting,
3, Marking of inclividu aanlds fo11owing-u opf their fligh atctivities
Markings were made on the ventral surfaces of hind-rig hwitngs (Fi g3.) using color magic
pens <oil-ty Mpaec,kie-Gokuboso, Zebra Co.). We tried to mark all the individua lthsat vis-
ite dPeak A and adjacent peaks during the survey period s(Apri l3 to 26, in 2002).
Although a few individua lats firs tescaped our capture and remained unmarked, in many
cases they revisited the peak and were marked, As almost all the individua] sobserved at
the summit had markings (Fi g4s, 5), the escaped individua lsesemed to have been rare,
The visiting times and their fligh ttracks were all recorded and traced on a map based on
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FlightBehaviors Luehdoi:t7(a1) 24t
ef
Fig, 2. All the data describe dwas recorded by the juni aoutrhor (KH) ,with the occasional
assistance of the senior author (KW). Identificati oonf markings was made using two meth-
ods. The firs wtas direc tobservation of the resting individua lussing naked eyes and field
glasses (Nikon M:icron 7× 15 CF), The second was confirmation of markings by capturing
the fiyin gbutterfl iweitsh a net, and after recording the markings, the butterfl iweerse re-
]eased .As we ¢ould not distingui sanhy recognizablc elfect on fligh btehavio ras a result of
repeated capturjng and releasing, we concluded that such etfects were negligible,
Results and discussion
1. Fligh tactivity and lif espan at the peak
1-1 . Flight activities
We marked 29 individua l(s27 8'2g) at the summit area (Peak As, B, C) in 2002 (Fi g4.).
The fac ttha t939L ･(2712 9of) the flyin gindividua lwesre males indicat etshat the hill-top-
ping is performed mainly by males, or at leas tthat it was in that year.
Fig. 5a clearly shows that the butterfl iiemsmediately stop fiyin gand disappea rin cloudy
conditions. The regularity is quit eimpressive ,as if the fiigh tactivities are rigidly restricted
not only by brightnes aslone but also by another unknown determining factor ,such as po-
larizeldightpatterns.

1-2. Timing of emergence and lif espans
After the firs etmergence (Apri 1l) at the peak, four fres hunmarked individua lwesre cap-
tured as late as April 19 (more than half a month later )(Fig 4.). The four might be heralds
of a late emerging populatio nas previousl ysuggested (Hashimot o1,991; K. Watanabe,
1998). However, we still cannot determine whethcr or not there are two such populations
with di'ffer egnetneti cbackgrounds .It is in any case interestin tghat the Mt Egesan popula-
tion display sa long duratio nof the adult emergence period ,even though Mt Egesan, in
southern Hiroshima Prefectur eh,as a temperate climate and is not a snowy provinc ewhere
the different imealtling of drifte sdnow would expand the duratio onf adult emergence.
In the cases of Nos 9 and 1O, they were observed for at least 15 days at the peak (Fig 4s,
5b). Their wing conditions were completely fresh on the firs tday and heavi]y stained on
the las tday, And the fligh atctivity on the last day loeked quit eattenuated, indicati ntghat
the lif espan of the individua lisn the longes tcase was around 15-17 days. This was consis-
tent with the data of Matsumoto (1984 i)n Kanazawa City .Additiona ldata recerded in
2003 and 2004 also confirm the fac t(Hiran eot al., 2006).
2. Are the fligh btehavior sat the summit individual slpyecific"?
From 20 individua lmasrked from April 3 to 13, eight individua lwesre observed at the
peak, for period sexceeding tive days (Fig 4s, 5) .It is remarkable that a censiderable num-
ber of early-emerging individual sstay around the peak area. On the other hand, individuals
marked on April 19 were not observed for sucb a long time (Fi g4.). The question arises
whether the late-emergi nigndividua lhsave a tendency fbr stronger migratory activity.
Howcver, at the same time, seasonal changes in temperatures and daylight conditions during
these two weeks may not be negligible and should be taken int oaccount.
The data showed that some individua lrespeatedly appeared at the peak; on the other hand,
other individua lapspeared only once or a few times ,indicati ntghat fligh htabit sat the peak
are individual lspyecific, and tha thabit smight change with aging (Fi g5.b), For example,
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FlightBehaviers Letehdo (1) 243
of ,Lfia
although individu aNlo. 4 (flyi tnhegre for 1 1 days) had a tendency of occasional appear-
ance at a young age, he changed his habit to stay fora long time at the peak in his older
age. No, 9 stayed at the peak on the afternoon of April S, and after a long (1 3days) ab-
sence, again appeared ther eon April 19. No, 1 1 showed a tendency ef appearing there re-
peatedly with a certain absent period (mor ethan 20 min) in the same day, No. 14 appcared
in the afternoon at a younger age, but late rin the morning (Fi g5.b).
When we compare the fligh ttrack of No. 4 and No. 14 in Figs 6a, b, we can recognize that
No. 4 tends to fly out over wider ranges of eastern clouded bushes from 11: Oe-13: OO (see
green arrows in "ALL" of Fig. 6a) .As for thc fligh htabit ofNo. 14, we could nut recognize
such a tendency (Fig 6.b), suggesting that the fligh btehavior sare individual slpyccific,
As human visual range is restricted to about 4 m in height and 90P in width, strict implica-
tion of the absence of butterfli ersemains a subject of discussion .One of the point sinvolves
how fai /they are flying from the peak. We will reply, in part ,to the questio nin Section 4.
3. Daily progression of fligh atctivities at the peak
Out of many flight-tr adcakta ,we show examples of No, 4 in Fig ,6a, and No. 14 in Fig. 6b.
3-1 . Acquisiti oonf flight-tr acocnksistency
Many males finis htheir dail yfiigh atctivity at the hill-t oopf Peak A and take an overnight
rest there on the leaves of pine trees .One author (KW) during these 15 years aL Peak A has
witnessed several cases in which iner tmales have fa]le nas a result of weak fiigh ftrom pine
trees to the ground befor etheir start of dail yactivities at around 8: 30 am (K .Watanabe,
1998),
After random preparativ efligh tfsor a short time, the butterfi isetsart to manifest a typical
fligh ptattern ,called "wandering fiigh t("Y ,Watanabe ,1996; K, Watanabe, 1998) ,in which
they fi yat a heigh tef about 20-70 cm from the fleQ rsurface as if they are wandering
around. The fia tfloo rconsisted of decp yellow-brow ndead leave sof pine trees showing a
dappled color ("Kanokomadar ain" Japanese )as a result of the sunlight through the crown
leaves,
The butterfi ireepseatedly rest C5-1 0seconds) on the ground followe dby wandering flight
of (1O-3 s0econds) for about 15 min. Eventuall yt,he butterfl iteesnd to come along to sunlit
areas ln otherwise shady areas, 1[b visit a few preferent iaandl specialized spots successively
means the fligh ptath becomes straight The consistcnt fiigh ttracks seem to be produced by
"looking" at the brightnes psatter nin fron tand by joini inn ga lin ea few brigh tspots. In
preparati vfeligh tt,he butterfl ileosok as if they are oriented only to the ground surface, but
in straight fiigh tthey have to watch in front using their compound cyes (Arikaw a2,001).
From the whole day fiight-tr adcakta (Fig 6sa, b), we can recognize the fligh ttrack is never
random but frequentl hyas a consistent fiigh rtoute. And, the fligh rtoutes of No. 4 and No.
14 were well-overlapped, Further w,e can recognize that there were a few preferenti aac-l
cumulating spots which corresponded to the brighte rareas (1- 2m in diameter )in the
course of the fligh ttrack,
3-2. Round patrolli nwigth resting points
With time ,fiigh atctivities became more active and fiigh atreas wider, and the butterfii eosc-
casionally fiew away from the observation area. Importantl yi,n due time, they came back
again to the peak (homing- i(nF)ig 6sa, b), This resulted in them travelin gin a circular
loop.
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TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society oofJfap anJapan
244 Kazuo WATANABE and Kazuhiko HiRANo
(a)
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Fig.S.Detaile tdiming ef the emergence of the marked individua alt sPeak A. Ca )Diurna lchanges
summarized for al] survey days .(b )The data of (a )was rcwritten for each individua l(Nos
1-29), The top lin cexpresses the time of appearancc of the butterfii east Peak A, 9: OO
means from 9 o'cloek to 9 o'cloek 19 minutes 59 seconds, and 9: 20 means from 9 o'clock
20 minutcs to 9 o'clock 39 minutes 59 seconds. W: weather (conditi oofn ssunlight). When
the sunlight was shaded by clouds, thc boxcs are sbadowed. Day: Survc ydays in Apri],
No.: Identificati onunmber ormarked individuals.
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TThhee　 LLeepipdiopdteorpoltoegiroaollogical　 SSoooiceityety 　ooff　 JJaapapnan
Fligh　tBehavior　sof 乙uehd ‘尹’プ1α （1） 　　　 542

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In　the　loopin　gobserved 　in　the　case　of　No ．140n 　l　l　o ’clock 　April　12，　the　butterf　lHycw
away 　fro皿 apoint （Qver 　bushcs　composed 　of　E‘4びα一Cα〃zellia−，Rhododendr（ノn　with 　a　height
of　ca 　2，5　m ；right 　photograp　hin　Fig．7），　travele　din　a　circle 　along 　a　route （about 　50　m ），　and
舳 ．ally　came 　back　f±om 　another 　route （the　northeastern 　small 　path；lef　tphotograph　in　Fig．
7）．In　the　case ，血e　butterHy　spellt　ccrtain 　times　at　six　precis　eresting 　point；sthree　were
brigh　tareas 　at　the　hil．−1top，　and 　the　other 　three　werc 　brigh　tareas 　along 　the　eastern　slanting
surface 　over 　the　bushcs（Fig．7）．　 At　these　points　the　butterf　isypent 　about 　10　secondsT3
1ninutcs ，　displayin　ag　few　repeated 　rests　and 　fligh，t（scα 2−20　scconds 　rest　and　cα 3−20　sec一
一
NNI工I工-EElleoetcrotniroonic　 LLiibrbarryary 　Service
TThheLee piLdeoppteirdoloopgitcaelrSooclleotygical Society oofJfap anJapan
246 Kazuo WATANABE and Kazuhiko HiRANo
(a)
Fig. 6.Detaile dflight-tr aocfk tswo exainples (No .4 and No. 14) m cuch day and the cumu]ative
sum of them (ALL) .(a )The case of individua lNo. 4. (b )Thc case of individu aNlo. 14.
Blue lettcr gand line swcrc fiig httracks of 9: OO-10: 59, green werc 1L :OO-12 :59, and red
were 13: OO-14 :59.
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