Life history of the Guatemalan Copper, Iophanus pyrrhias (Godman & Salvin) (Lepidoptera, Lycaenidae)

2010·0.81 MB·

TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan
estma Trans.Iepid.Soc.Japan 60 (4)2:59-267,March 2010

Life history of the Guatemalan Coppe4 lophanus pyrrhias (Godman & Salyin)
(LepidopteLryac,aenidae)

Masaya YAGoi) ,rlhkashi MiyAGAwA2), Jun YoKoyAMA3) and Mark WiLLiAMs4)
')The Universit Myuseum, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-O033
Japan; e-mail: [email protected]
2'3-53-i5-402Eifuku,Suginami-ku,Tokyo, 168-O064Japan

3) Department of Biology ,Facult yof Science ,Yamagata Universit y1,-4-1 2KQiirakawa-machi.
Yamagata City,Yarnagata,990-8560 Japan

`' Department of Paraclini caScliences, Faculty of Veterinary Science, University of Pretoria.
Privat Beag X04, Onderstepoor tO.1 1O Republi cof South Africa
Abstract The Guatemalan Copper, lbphanus pyrrhias, is fbund in Guatemala and southern
Mexico. There are no closely re]ated species in the Neotropical and Nearctic regions, and therefore
i ti sof intere stto study this species from a biogeographic paolint of view, In order to elucidate the
lif ehistory of L pyrrhias, the habit sand immature stages were studied on two occasions in the
Guatemalan Highlands. We describe the morphology and behaviour sof al 1lif estages in the
species, and compare our finding wsith previou snotes made and publishe dby earlier researchers.
We record Muehlenibeckia tamn(fblia, occurring along the Andes in Centra land South America, as
itshostplant.

Key words Copper, egg, Guatemala ,immature stages, hostplant ,Iophanus, iophan"s pyrrhias,
larva ,Lycaenidae, Lycaeninae, Muehienbeckia tamnijbiia, pupa.
Introduction
The Guatemalan Copper, Iophanus pyrrkias (Godman & Salvin i)s, distribut eidn Guatemala
and southern Mexico, and is well known as the only Neotropica lspecies in the bycaeninae,
a subfamily of the Lycaenidae that contains some 100 nominal species (Bozan o&
Weidenhoffer ,2001; Yago & Saigus a2,002), Although 16 lycaenin especies occur in the
Nearctic region, L pyrrhias seems to have no allied species in the Americas, according to
morphological characters of the wing markings and male genital i(aMille &r Brown, 1979;
Sibatan i1,974; Yago & Saigusa ,2000; Yago & Saigusa ,2002). TherefOre, as suggested by
previou sresearchers (Sibata n1i9,74; Miller & Brown, 1979) ,it is of much intere stto clari-
fy the historic aplhylogeography of I. pyrrhias .However, ayailable data relating to the life
histor ysheds litt lleight on phylogeneti cand biogeographical aspects of it sbiology .As far
as we know, there are only short bielogica nlotes on this species (Draud t19,20; Miller &
Brown, 1979; D'Abrera,1995;De laMaza, 1987),

For the purpose of elucidating the lif ehistor yof I. pyrrhias, we investigat etdhe adults and
immature stages of the species in Chimaltenango ,Guatemala, where previou sresearchers
have also worked (Mill e&r Brown, 1979) .We fOund eggs, larva eand adults of the species
in the field b,ut the pupal stage was only observed when the species was reared in captivity.
Here we describ ethe morphological and ecological characteristics of L pyrrhias, and com-
pare our finding swith previou sreports. We also discuss the similarities to and differences
from other lycaenine species, based on the morphology of the immature stages, and show
the new data to clarify, to some extent, the phylogeographica alnd evolutionary processes
relating to this species.
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260 Masaya YAGo, Takashi MiyAGAwA, Jun YOKoyAMA and Mark WiLLiAMs
Fig. 1. Views of Volctin Acatenango (righ atnd) Volc6n Fuego (lef ftr)om a market of Antigua.
Materials and Methods
The adults and early stages of I, pyrrhia swere searched for in the field by the firs tand sec-
ond authors. The research was conducted at Acatenango (al t2.,200 m), Chimaltenango,
Guatemala from August 1st-7th 2005 and from March 22nd-26th 2006. Vblcan Acatenango
(3,97 6m) is locate dto the west of Antigua and is one of the active volcanoes in the
Guatemalan Highlands (Fi g.1) ,Although there are comparatjvely distin crtainy (May-
October) and dry (November-Ap rseialso)ns in this area, the highland sare characterized by
a consistently cool sub-alpine climate with chilly nighttime temperatures, High altitudes of
the mountain are associated with volcanic ashes and sandy soils, with pine-forest esdlopes
up to the tree lin eat about 3,800 m. The mid-slope of the mountain, where I, pyrrhias and
it shostplan twere observed, is coyered primaril yby humid cloud fbrest sW.e also found
alsohostplants,the The
some other polygonaceous plants ,possibly at study site. scientific
names of plant swere determined by the third author.
When adults or early stages were fbund, we recorded the indiyidua lisn situ using two digi-
tal cameras, a Konica Minolta DiMage X50 and a Nikon D70 with a micro lens (Niko nAF
Micro Niklcor 60 mm). The body lengths of all the larval instar swere measured jus tbefore
diapause .Because we could not find any pupae, larvae were reared at the study site to the
pupal stage in an outdoor cage (25 cm diameter ,32 cm height), which was made of a nylon
net with a circular wire frame at the top and bottom. The cage was hung about 50 cm above
the ground and the bottom was attached to a livin ghostplan tc,overing it sapical part .After
rearing the larvae and recording the pupae, the latte wrere released in the field.
Results
Eggs (Fi g2.). Approximately O.6 mm (n = 3) in diameter ,white, hemispherica lf,iattened
basall yand covered with ridged reticulations. Micropylar area rather small and deeply de-
pressed in central axis of egg dorsall yR.emaining reticulate surface composed of concave
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Life history of lophanus p.vrrhias 261
chorionic cells and thick prominent ridges. Chorionic cells depressed ,cup- or rice-ball-
shaped and usually surrounded by three intersect enoddular processe sand three-sided chori-
onie ridges. In dorsa lview chorionic cells radiate in a spiral shape from the central axis, and
are somewhat 1arge rand deeper on submarginal portion b,ut in later avliew graduall yshal-
lower and smaller toward the lower portion.
The eggs are usually laid singly on the upperside ofcomparatively young leave sof the host-
plant ,Our observations show that the eggs have a comparatively high percentag e(abou 5t0
%) ofparasitism by braconi dwasps in the study site.
Larvae (Fig 3s-5) .Oniscifbrm ,as in most lycaenids r;emarkably flatten eidn earlier instars
body in instarF.irstinstarlarva 2 (n
and with slight color variation each approximately mm
3)inlength, flatteneddorsally line (Fig,
= pale green, and with mid-dorsal somewhat edged
3). Second and thir dinstar sflatten ebdut slightly swollen dorsally ,and showing body color
variations from even]y ligh tgreen to ligh tgreen with red mid-dorsal lin eand a pair of pale
yellowis hsubdorsal line s(Fig 4,). Second insta rabout 3.5 mm (n = 3) and third insta rabout
8.5-9 mm in length .Final insta rlarva eevenly brigh tgreen ,rather slender and somewhat
hemi-cylindric a(lFi g5,). Body length of fina linsta r12.5-13, 5mm (n = 3),
Our observations show that the larva erest mainly on the underside of leave sof the host-
plant throughout the larva lstage. In the fina linsta rthey are occasionally fbund on the up-
persid ewhen the leaf is upright. When feeding on leave sof the hostplan tt,he 1arva eleave
an irregula pratchwork of feedin mgarks on the underside of the leaf and do not distur bthe
upper epidermis of the leaf .However, fina linsta rlarva eoften eat larg eholes in the leaye sor
they may consume the entire leaf .The larva eare not associated with ants in any larva lin-
star.
Pupae (Fig s6-7) .Simila rto other lycaenin especies; slightly rough, squat, rather gourd-
Iike dorsall ybut somewhat flattene dventra]ly, covered with trumpet-shaped minute setae on
extemal surface except for ventral portio nand wings. Body color reddish brown dorsally
but somewhat paler ventrally, with three rows of fine black spots arranged mid-dorsally,
subdorsally and ventrolaterally, from mesothorax to abdominal segment 9; wings creamy
brown on ground color, variably spotted and mottled with dark brown, which is extensive
dorsally s;piracles black .Pupa attached to silk mat spun on dead leaf by metathoracic silk
girdle and anal hooks. Pupal length 9-9.5 mm (n = 3).
Aduits (Fig s8-10) , Wing shape as in thecline species. Forewing triangular ,apex rather
acute; hindwing tailed at tip of vein 2 and prominently lobed at tornal area. Female wing
shape similar to male, but somewhat rounded and transversely widened in both wings. Male
upperside deep purple with narrow costal and outer marginal black borders ,Female upper-
side dark brown, blue-purpl beasally a,nd with 1arge e,lliptical, fain otrange disca lpatch tra-
versed by two black lines on fbrewing, Hindwing upperside with orange submarginal
lunules in both sexes. Undersides of both sexes cryptic; fbrewing rosy brown with ochreous
patch traversed by three dark brown lines ;hindwing mottled with dark brown and rosy
brown. Forewing length 13-14.5 mm (n = 3) in male and 13 mm (n = 3) in female,
Habitat(Figs11-13).This frequents fields, forest
species grassy pastures, clearings and
fbres tfringe salong a mountain path in a narrow valley surrounded by dense bushes (Figs
11-12) ,The males fiy slowly, close to the ground, in sunlit patches and imbibe fluid sfrom
horse-dungbetween 09:OO 12:OO(Figs8,10).During
mud-puddles or and warm sunny
weather males show territorial behavior ,using shrubs O.5 m to 2 m high as perches .The
hostplan tgrows under bushes or at fores tedges along the mountain path or in the pastures
(Fig .13) .Females are found on or near the hostplant dsuring the day (Fi g9.).
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262 Masaya YAGo, TakashiMiyAGAwA, Jun YOKoyAMA and Mark WiLuAMs
tt./.tt..t..R.l//,,iit
lttt/.-.'t"g
l ttt
6 7
Figs 2-9. Immature and adult stages of IQphanus pyrrhias 2.. Egg, dorsolate rvaielw. 3, Firs tinstar
larva ,dorsa lview. 4. Third insta lrarva ,dorsa lview. 5. Final (fourt ihn)sta lrarva ,dorsal
view. 6. Pupu, doTsa lview. 7. Pupa, later avliew, 8. Male adult settling on a leaf .9. Female
adult basking in the sun.
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Life histor yof Ibphanus pyrrhias 263
Figs 10-l7. Adults, habitat sand hostplan tofs Iophanus p.vrrhias .1O, Maie adu]ts sucking up water
on the ground .11-12. Habitats of male adu]ts, 13. Habitat of female adults with hostplant,
14-15. Hostplan LMuehlenbecki atamnijblia (Polygonace ae1)6.. Rumex acetosetla
(Polygonace a1e7). R.umex obtusijbtius (Polygonaceae).
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Hostplan t(Fig s14-15) .At our study site females were observed layin geggs on compara-
tively young leaves of Muehlenbeckia tamnijblia Meisn. (Polygonace Aalel )e.ggs and lar-
vae that we fbund were also discovere don Muehtenbeckia plants ,Muehlenbeckia tamnijio-
tia is distribut efdrom Mexico to Argentina ,along the Andes mountain range,
Discussion
From two surveys cenducted at our study site we discovere dall of the lif estages of L
pyrrhias ,although pupae were reared but not found in the wild (Fig 2s-10) .The immature
stages and adult female of the species are probably described or illustrate fdbr the firs ttime
in this study.
Concerning the habita tof I, pyrrhias, Draudt (1920 )and D'Abrera (1995 n)oted that it oc-
curs at altitudes between 6,000 and 7,OOO feet (1,830-2, 1m)3 0in montane Guatemala, fiies
close to the gound and is fbnd of alighting on pig droppings .According to De la Maza
(1987 )this species occurs in mountainous, fagaceous-pinaceous, temperate forest sof
Chiapas (Mexico )in August. As mentioned in the results section of this study, we observed
the species in almost the same situation and habitat sT.hus, previous reports are not inconsist-
ent with our own observations.
We investigat etdhe lif ehistor yof L pyrrhias in August 2005 and March 2006, In the first
survey, in summer, the egg, larva land adult stages were observed in the fiel d(Fig s2-5,
8-1O) .In the second survey, in early spring, the second author could not find the adults due
to a long spell of bad weather but confirmed the egg and larva lstages on the hostplant.
Although De la Maza (1987 )noted that the fligh tperiod of this species is restricted to
August, the actual fiigh pteriod seems to extend over multiple broods.
In lycaenin especies, the most detaile dstudy on morphology of immature stages is that by
Wright (l983 )H,e investigat etdhe ultrastructure of eggs, larva eand pupae of a Nearcti cly-
Eipidemia (Boisduv&alLe Conte),
caenine species, qpixanthe using a scanning electron mi-
croscope. Other detaile dstudies of immature stages are those of the South African L.vcaena
orus (Stol alnd) [,ycaena clarki Dickson by Clark & Dickson (1971) I.n addition, scanning
electron micrographs of eggs were publishe dby Miller & Brown (1979 f)or the Nearctic
Elpidemia helloides(BoisduvalE)P,idemia dorcas (Kirby) theHolarcticL(ycaena
and
phlaeas (Linnaeu sOt)h.er studies of the immature stages of Lycaeninae includ ethose in
New Zealand by Gibbs (1980 [)Helle siataustius (Fabrici uHest)te,ia .feredayi (Bates a)nd
Botdenaria boldenarum(White)t]hose inthe Palaearcticby Henriksen & Kreutzer (1982)

[Heode svirga"reae (Linnaeus H)e,odes tityru s(Poda von Neuhaus), POIaeochrysophanus
hippothoe (Linnaeu san)d L. phlaeas] .Fukuda et al. (1986 )studied the immature stages of
L. phtaeas and Igarash i& Fukuda (1997 )the Palaearct icT. dispar ,and the SE Asian
Hetiophorus kiana (Grese-smi tahnd) Heliophorus ila (de Niceville) .Yligo et al. (2005 r)e-
ported and illustrate dthe early stages of the SE Asian Heliophorus indicu s(Fruhstorfer)
and Heliophorus delacour iEliot.
Our study showed that, compared with the immature stages of some lycaenin especies from
these previous studies, the egg of l. pyrrhias resembles those of Heliophorus species in the
smaller chorionic cells and thicke rchorionic ridges (Igara s&h iFukuda, 1997; Ylaig oet al.,
2005). However, it dilfer isn having more chorionic cells and somewhat smaller nodular
processes .In appearance, the lary aof I, pyrrhias is very similar to those of Heliophorus
species in body shape, in that younger instar sare flatten eddorsall yand the fina linsta ris
somewhat slender. As fbr the pupal morphology, 1. pyrrhias has many trumpet-like setae in
common with other species in the Lycaeninae. The pupal shape and color patter ndoes not
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Life histor yof lophanus p.vrrhias 265
resemble that of other lycaenin especies, although the dark brown body color is shared with
the pupae of L. phataeas, H. tityru sand T, dispar ,Although we do not know whether the
character states mentioned above are apomorphic or not, overall L pyrrhias appears to be
more similar to Heliophorus species than the ether lycaenin egenera for which the im-
mature stages are well known. However, the pupal morphology of Heliophorus species is
quit edifferen tfrom that of L pyrrhias ,especially in that the body color is light green and
the abdomen is roundly swollen (Igara s&h iFukuda, 1997; Yago et al., 2005). The under-
sides of the wings of both sexes and the upperside of the male of L pyrhias are similar to
those of Papua New Guinean Melanotycaena species, and the female upperside is close to
those of Chinese-Himalayan Helleia species. Future research by us will therefore focus on
the immature stages of both Metanot.vcaen aand HeUeia species.
Hitherto, a species ofRumex (Polygonacea hea)s been recorded as a hostplant of L pyrrhias
(Mille&r Brown, 1979).However, larvae females discovered
eggs, and egg-laying were
only fbr M"ehlenbeckia tamndelia (Polygonace ian eth)e presen tstudy (Fig 1s4-15) .At our
study site two introduce dRumex species, Rumex acetosella L. and Rumex obtusijblius L.,
were found in the habitat of L pyrrhias (Fig s16-17) but these piant swere not utilized as
hostplants by L pyrrhias .In Guatemala there are native Rumex plants ,such as Rumex mexi-
Meisn. Rumex Moris(=Rumex berlandierMieisn.),
canus and chr:ysocarpus which are en-
demic to Guatemala, Mexico and the southern part of the USA (Stand] e&y Steyermark,
1946). A wonhwhile extension of this study would be to investiga twehether I. pyrrhias
feeds on these endemic plants .Judging from the known hostplan tI,bphantt spyrrhia smay
be closely related to Papua New Guinean Metano4}Tcaena species. In fact ,Sibatan i(1974)
noted that the Melanolycaena species feed on another plant of the same genus,
Muehlenbeckia monticola Pulle & Danser. In addition, he also suggested that the
Melanol: caena species are similar to J. pyrrhias in regard to external morphology of the
adults, particularl ythe wing markings and the male forele gtarsus.
Miller and Brown (1979 )noted that ,as suggested from it sdistributi oLn ,pyrrhia sis the
most primitiv esurviyor of the Cretaceous-[Ibrt iianrvyasio nof the New World, and that it is
fbund in the same region where Anetia (th emost primitiv edanaid) and Baronia (th emost
primitiv peapilioni do)ccur. However, de Jong (2003 )doubted this hypothesis, and denied a
Gondwanan origin for Australian butterflie sb,ased on fossi land molecular evidence. In any
case, phylogeneti crelationships among lycaenin egenera are not yet clear, even though
there are analyses of the Nearctic genera (Mille &r Brown 1979) and the Holarctic genera
(Zhdanko ,2000) that are based on morphological characters. Given the paucity of infbrma-
tive biologica alnd morphological data gained from the presen tstudy of L pyrrhias, further
work is needed in order to elucidate the historic aplhylogeography of the species, in particu-
lar by analysis of DNA sequence data within the subfamily Lycaeninae.
Acknowledgments
We would lik eto express our thanks to Dr. M. Kondo (Univers oift [y[bkyo) for her critical
reading of an earlier draft W.e are also deeply gratefu tlo Mr. N, Nagahata (Saitam af)or his
valuable infOrmational inputs.
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摘　　要
グァ テ マ ラ クロ ベ ニ シ ジ ミの 生活史（矢後勝也 ・宮川　崇 ・横山　潤 ・Mark　Williams）
一
ベ ニ シジミ亜科 （あるい はべ ニ シジミ族）で唯 の中米種であるグァ テマ ラクロ ベ ニ シジ ミは，メキシ
コ南部か らグァ テ マ ラの 高標高地にの み生息 し，北米に産するベ ニ シ ジ ミta　16種とは近縁性が見られ
ない稀種で ある．その ため．本種の 進化や系統地理の 解明は非常 に興味深い が，この不可思議な分布の
謎を解くの に重要な系統学的 生物地理学的研究の基礎となる生態情報や幼生期の形態情報がほ とん
，
ど知られてい ない ．そ こで筆者らは．本種の 生活史解明 を目的と して 2005年と2006年の 2度にわた り，
グァテマ ラ高地におい て本種の幼生期および成虫の探索を行っ た．その結果，現地での袋掛け飼育によ
ー
る蛹期を除く全ス テ ジの野外観察 撮影に成功した．全幼生期および♀成虫の図示，さらに幼生期の
記載はおそら く本稿が最初となる．本種は深い 低木林に囲まれた浅い 峡谷の 山道沿い にある放牧地周
辺や草地，伐採地あるい はその 脇の ブッ シ ュ に生息 し，♂は午前中に地面近 くをゆ っ くり飛翔，水たま
りや馬の 糞尿で 吸水する姿が 目撃され，午後になる と低木上で 占有行動が見られた．♀は♂よ りも個体
数は少なく，食餌植物の近 くで終日観察された，また野外観察から本種は多化性であることが示唆され
た．
本種の卵はchorionic 　cell が小さ くchorionic 　ridges が厚い 点で．ウラ フチベ ニ シ ジ ミ属の種 にやや似る．
幼虫は若齢時で より扁平で，終齢ではやや筒状かつ 細長 くなる点で，やはりウラフチベ ニ シジミ属の種
に類似する，ただ し，蛹はべ ニ シジミや旋 046 ∫  r岬 などの真性ベ ニ シジ ミ類のよ うに全体の 起伏が
弱 く，色彩は褐色で 細かい斑点を散布する点で，黄緑色で丸 く膨張 した腹部を持つ ウラフチベ ニ シジミ
属 と大 きく異なる．♂成虫の 翅表や♂♀両方の翅裏はパ プアニ ュ ーギニ ア固有のパ プアクロ ベ ニ シジミ
属2種にかなりよく似るが，♀翅表の楕円形中央橙色斑の現れ方は中国西部一ヒマ ラヤの高地帯に固有
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TThhee　 LLeepipdiopdteorpoltoegiroaollogical　 SSoooiceityety 　ooff　 JJaapapnan
Life　hjstor　yof　Jophanus　P．vrrhias 267
の Helleia属 グル ープに類似する．・・方，本種の 食餌植物 として タデ科の ギ シギ シ属の一種がこ れ まで
記録されて い るが，今回の 調査で は タデ科の Muehlenheckia　tamnifotia の みか ら卵，幼虫，♀の 産卵行動
が観察 された．こ の情報か ら判断すると，本種 と同 じくMuehtenbeckia属を食するパ プア クロ ベ ニ シジ
ミ属の種との近縁性が認め られ るが，成虫の形態で も翅の斑紋や前脚の跚節をは じめ とする多くの 共
通 した形質を本種とパ プア クロベ ニ シジ ミ属が持ち合わせるこ とが知られる．総合的に見ると，本種は
ウラ フチベ ニ シ ジミ属やパ プア クロ ベ ニ シジミ属，中国一ヒマ ラヤ固有の Helleia属との 類似性が認め
ー
られるが，今回得 られたデ タから共通形質が派生的な状態であるかどうかの判断がで きず，類縁関係
の議論は難しい ．また，上記のパ プアクロ ベ ニ シジ ミ属やHellei　ak グループを含む多くの種の幼生期
が図示されてい ない ため これらの 生活史の解明が今後の課題 となる．
，
Baronia やAnetia（最 も原始的なアゲハ チ ョ ウ科とマ ダラチ ョ ウ科）が同地域の 中米に生息する こ とか
ら，グァテマ ラクロベ ニ シジミは新世界へ 中生代白亜紀一新生代第三 紀に侵 入した最も原始的な遺存種
である とMiller＆ Brown （1979）は指摘 してい る．しか し，　de　Jong（2003）は化石証拠や最近の 分子系統
学的研究からこの仮説を疑問視 して い る．いずれにせ よ，本種が生 じた歴史的背景は未解明の ままであ
るため，今後はDNA による分子系統解析も用い るこ とで，ベ ニ シジミ亜科全体の系統地理を明らかに
してい く必要があるだろ う．
（Accepted　June　25，2009）
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