Table Of ContentVol. 2 o. I 1995 DA TCHE KO et af.: Egg of Russian Theclinae 27
HOLARCTIC LEPIDOPTERA, 2(1): 27-38
EGG STRUCTURE AND NOTES ON
BIOLOGY OF THECLINAE FROM
PRIMOR'E, RUSSIAN FAR EAST
(LEPIDOPTERA: LYCAENIDAE)
ALEXANDER DA TCHE KO, ANDREI SOURAKOV, and THOMAS C. EMMEL
Leninskiy Prospect 33, Inst. ofEvolutionary Morphology and Ecology ofAnimals, Moscow, Russia;
Dept. ofEntomology and ematology, ntversity ofFlorida, Gainesville, Florida 32611; and
Depts. ofZoology and Entomology, Univer ily ofRorida, Gaine ville, Florida 32611, USA
AB TRACT.-Thebiologyandsystemalicsofhairstreakbuuernie ofthegeneraArtropoetes,Araragi.StrymoTlidia. Neozephyrus, Wagimo. UsslIriaTla.
FixseTlia. Thecla, Chrysozephyrus, Japollica. ShiroZl/a, and Alltigil/s from the Rus ian Far Easl are discussed. The life historie and especially the
morphology oftheeggs are useful in drawing taxonomic conclusions.
KEY WORDS: Amur, ATltigillS, ApllllaellS, Araragi, Arhopala, Artropoetes, Asia. behavior, Belulaceae, China, Chrysozephyrlls, ecology, Eumaeini,
Fagaceae,FixseTlia,ho lplant ,immalure ,JapoTlica,Juglandaceae,Korea.larvae,Loxura,Manchuria,morphology,Neozephyrus,Olea eae,Palearctic,
Papilionidae,Rhamnaceae, RhopaJocera, Ro aceae, RUlaceae.Shirozua,StrymoTlidia, sy temalics, Thecla,Theclini, Imaceae, Umbelliferae, ssuri,
Ussllriana, Wagimo.
Primor'e i a mall area ofthe Russian Far East, located in the sandydunehabitatalong thecoast whereQuercusde1l1ata Thunb.
very outheast of Russia, next to Korea and Manchuria. Due 10 dominate. This area i the mo t northern borderofthedistribu
its warm local climate, it contain a high diversity ofanimal and tion of Quercus delltata, which i found mostly in ubtropical
plant life. This tudy con ider the biology and egg morphology Asia. The Prihankaiskiy region is different not only becau e of
of several pecies of Theclini which occur sympatrically in the its climate (autumn in this place is unusually warm), but also
southern Primor'e. becau eofits teppe-type land cape, which i morecharacteristic
There have been no studies on the biology ofTheclini of the ofthe Amur region. Mid-elevation forest ofthe regional Pogra
Far Ea t ince the end ofthe last century (Grae er, I -92). In nichniy mountain chain consists mo tly of Quercus mOllgofica.
a en e, thi ituation i the re ult of the typological concept Our studies were conducted from early April 10 the end of
whichdominated ystematic during thelastcentury. It would be September. They con isted of collecting egg and larvae in
unjust tosay today thatsystematists completely ignore biological nature and breeding butterOie from the egg obtained from the
data: on thecontrary, it is very common to Ii t, forexample, food females in the laboratory. The object ofour tudy were mo tly
plants. Often, thi information i transfered from one work to tree-feeding Lycaenidae of the Theclini and Eumaeini groups.
another, forming whole lists of foodplants, which ometimes are All species of these groups are as ociated with trees or hrubs
noteven found in the region studied. Such works can reduce the and overwinter in the egg stage. Collecting egg in nature
desirability ofusing biological information in taxonomy, due to allowed us to estimate habitat preferences, as well as to obtain
a great amount of misleading information in the literature. information on the oviposition pattern of different species.
Interest in life history studies is on the rise now in general, due,
undoubtedly, to their importance for conservation. However, the PECIES ACCOU TS
u e of thi kind of information for taxonomic purpo e is more
the exception than the rule (Saigusa, 1993). Life history tudies, Strymo1lidiJI ilicis ilicis (Esper) (Fig. IL) i found only in the
of course, are not a ubstitute for regular taxonomic study of European part of Russia, associated with young oak growth in
adultcharacters,butingroupswithfew morphologicaldifference open area. The female lays egg on the fresh shoots. Fir t
in the imago, they might be very essential. Tree-feeding in tar larvae bore into the buds. The last in tar larva' coloration
Theclinae represent a group where morphological details can be re embles the color of the young oak leaves on which it feeds,
quite useful in taxonomy. sitting on the lower surface of the leaf (Fig. 31). The larva
In 1991-93, we tudied thesystematics and biology ofTheclini pupate on the ground, or occasionally on the shoots. Adult
in two regions of Primor'e: at the very southern end of the hatch at the end of1une.
region, in thedeltaofthe Riazanovka Riverand in the Prihankai Strymo1lidiJI latiorliltior (Fixsen) (Fig. IK). The larva feeds on
kiy area, in the vicinity of the town Barabash-Levada. The Rham/lus ussuriellsis Va sil (Rhamnaceae). Thi species isoften
South Primor'e site includes hilly habitat at IOO-600m elevation, con ideredasubspecie oftheEuropeanSlrymollidiaspilli(Fabri
with domination of Quercus mOllgofica Fi h. (Fagaceae) and ciu ), to which it is closely related. There are significant differ-
PLATE2.(I)Neozephyrtlsu/rrall/orillusFixsen:(a)male;(b)maleunderside;(e)female:(2)N.((ui/a Bremer:(a)male;(b)maleunderside;(e)female;(3)N.saphirilllls
Siaudinger:(a) male;(b) maleunderside;(e)female;(4)N. querCllsL.: (a)male;(b)maleunderside;(e) female; (5) Wagill/osigllaraquercivoraStaudinger: (a)male;
(b) male underside; (6) Fix.<efliaprufliL.• male.
PLATEI.(A)NeozephyrtlssaphirillusStaudinger;(B)Afltigillsbut/eriFenton;(C) eozephyrusquercusL.;(D)N.aquall/arillusOubalolov&Sergeev;(E) Str)'l1/ollidia
exill/iaFixsen;(F)Neozephyrtls taxi/a Bremer;(0) N.japoflicu.r Butler: (H) Thecla betu/ae crassa Leech; (I)AfltigiusaI/ilia Bremer; (J)Japollica olloi Murayama;
(K) Str)'l1/ollidia /alior Fixsen;(L)Srr)'l1/ollidiailicis Esper.
Vol. 2 No. I 1995 DANTCHENKO et al.: Egg of Ru sian Theclinae 31
PLATE4. (I) Tlrecla betulae L. (a) male; (b) male underside: (2)AraragielltheaJanson; (a) upperside; (b) underside; (3)Al/ligiuJallilia Bremer; (a) upperside: (b)
underside; (4) AlltigiuJ butleri Fenlon; (a) upperside; (b) underside; (5) JapalliclI O1wi Murayama: (a) upperside; (b) underside: (6) Jap01,ica lutea Hewilson; (a)
upperside; (b) under ide.
ences, however, in the morphology ofmales, females, and larvae plant is found mainly in the under tory of the forest. Thus S.
of S. s. latior and all other European subspecies of S. spini. eximia is separated from S. latior ecologically. An egg of S.
Beside, the disuibutional areas ofthe e taxa do not overlap, nor eximia was found on Rhamnus ussllriensis Vassil only once(Fig.
do the distribution oftheir food plants: S. spini feeds on Primus 51). In the laboratory, larvae were raised on Rhamnuscatarclica.
spinosa L. (Rosaceae) or Rhamnlls catarctica L. (Rhamnaceae). Adults hatch several days later than S. latior. The larva is hown
These fact are probably the reasons that S. I. lalior is listed in in Fig. 3G.
Tuzov's Synonymic List ofButterfliesfrom Ihe ex-U.S.S.R. as a Strymollidiaw-alblllll w-alblll1l (Knoch)isfound in theEuropean
separate species (Tuzov, 1993). The adults fly in open areas. part of Russia. Here the egg is considered for comparison with
Females lay egg clu ter of two (in the branching points of thin the Far East sub pecies. The egg i shown in Fig. 6H.
branche ) to 80 (at the ba es of the trunk) (Fig. 60). The larva Strymollidia w-albulII slItscJ/Qni (Butler). The taxonomicstatu
bores intoajust-opened bud and laterstays inside thedeveloping of thi taxon is unclear. It is considered a synonym of the
flower clusters. The last-instar larva pupates in the litter on the sub pecies S. w. felltoni in the latest literature. However, we
ground. Underlaboratorycondition, itpupates in thecontainer's consider this taxon as a valid species. The actual position ofthis
layer ofdry leaves, attaching itselfto the leaf with a ilk thread. taxon could be determined only by comparing the biology and
Adults hatch in the middle ten days ofJuly. morphology of all the available taxa in the group. We have not
Strymonidia eximia eximia (Fixsen) (Fig. IE). Larvae feed on observed itsoviposition behavior in nature; however, thisspecies,
Rhamnlls diamamiaca Nakai (Rhamnaceae). Eggs are laid in as with other species ofthe group, is likely to be as ociated with
clusters ofone to four on the uneven surfaceofshoots. The food Ulmlls sp. (Ulmaceae). In the laboratory, larvae were raised on
Ulmlls levis L. Adults feed on Umbelliferae together with S.
PLATE 3. Last instar larvae: (A) Araragi ellthea Janson; (B) NeozephyruJ
exi1llia and S. latior, and fly in the second halfofJuly. The egg
IIltramarilluJ Fixsen: (C) N. aqllau",rilllls Dubatolov & Sergeev: (D) Wagima
siglla/{l Staudinger; (E) Thecla bellllilla laudinger: (F) NeazeplryruJjapolliclIs is shown in Fig. 61.
regilla Butler; (G) Strymollidia eximia Fixsen: (H) S. latior Fixsen: (I) S. iliciJ Fixsellia prtmi prlllli (Linnaeus). Larvae of this species bore
Esper; (J) Amigills bwleri Fenlon; (K) Thecla bellliae L.: (L) AmigillJ allilia into the buds of PrtlllLlS pad,.ts L. (Rosaceae) after hatching from
Bremer; (M)Japollica olloiMurayama;(N)NeozeplryruJsaphirillll., Staudinger:
the egg. Adults fly in open areas in the middle ofJune.
(0) N. /{lxi/a Bremer.
32 DA TCHE KO et aL.: Egg of Rus ian Theclinae HOLARCTIC LEPIDOPTERA
Fixsellia heni heni (Fixsen) lay eggs on Pyrus ussuriellsis eozephyrlls taxi/a (Bremer) (Fig. IF and Fig. 2(2)). Thestudy
Maxim. (Graeser). (Rosaceae) In the laboratory conditions, it of Bremer's type material by Dubatolov and Sergeev (1987)
lay egg and feeds on Ma/us malldshurica (Maxim.) Kom. howed that N. raxi/a is conspecific with N. cogllafLIs (Staudin
(Ro aceae). Adults are often seen on flower in mid-June. ger), which then became ajunior ynonym. Neozephyrus taxi/a
Wagimo signata (Staudinger) (Fig. 2(5». The larva of this feeds on an oak, Quercus mOllgo/ica (Fagaceae). The egg (Fig.
pecies hows more specialized features (Fig. 3D). It is found 5J) i laid next to the bud, the female preferring theside hoot.
only on Quercus 1II0llgolica Fi cher (Fagaceae). Egg are laid Hatching of the larva from the egg is synchronized with the
mostly on the large flower buds found in the upper part of the opening of the bud. Thus it is likely that the fully formed larva
oak-treecanopy, and i~ cluster ofone to ix egg (Fig. SF). The overwinters inside the egg. The first instar larva molt on the
larva of the first instar bores into the bud. Later in tars feed on inner ide ofthe bud cales. The last in tar larva (Fig. 30) tay
flower and fre h leaves. Under laboratory conditions, the larva next to the bud area and phenotypically re embles the bud scale.
pupates on the tern. According to Yokoyama and Wakabaya hi Among congeners, this specie appears earlie t: the end ofJune.
(1967), this species pupates inside a shelter which the larva Neozephyrllsjapolliellsjapolliells (Butler)(Fig. IG).Theauthors
nibble out ofthe bark. Adult hatch in the middle ofJuly. of the work in which the lectotype of N. raxi/a (Bremer) was
Alltigills attiLia (Bremer) (Fig. II and Fig. 4(3» and designated (Dubatolov and Sergeev, 19 7), proposed toapply the
Alltigills blltleri (Fenton) (Fig. IBand Fig. 4(4». Adults, larvae name N. japollieus to bUllerflies as ociated with pecie ofA/llus
(Fig. 3J,L), pupae and genitalic tructure are very similar. The (Betulaceae). While our purpo e here is not to propo e a
egg ofA. blll/eri i similar to that ofA. alli/ia, though in the egg taxonomic revision. and we cannot confirm the identity of the e
ofA. atti/ia, the projections are longerand the egg is flatter (Fig. names, we can make these observation. To determine whether
SA,B). These features probably have been selected as part ofthe these taxa are good specie, one would have to ee their types,
different ecologie of oviposition. A. allila lays a olitary egg and preferably al 0 compare modern ampIes of population
into the gaps and folds in the bark of Quercus mOllgo/ica from their type localities. There i no firm evidence that the
(Fagaceae). A. butleri, on the otherhand, lay eggs in clusters of population we are dealing with in Primor'e belong to this
6 to 12 eggs under the old bark where it is separated from the species, or even a ingle pecie. The eggs (Fig. 6 ) were
trunk of Quercus mOllgo/ica. In the zones of overlap of Q. collected from several different species ofA/llusand were mixed
delJlata Thunberg and Q. mOllgo/ica, eggs of both species have together. The laterstudyofthestructureof ISOeggs showed that
been found occa ionally on Q. delJlata. This shared all the eggs fall into one of two morphological group, with
intermediate-host situation doe not happen in pecie in which almo t no overlap. The adult bUllerflie arequite variable. Eggs
the female lays egg on hoot or buds. In tho e pecie, are laid on different parts of the plant: shoot, buds, and old
urvivor hip of the larva strongly depends on the preci ion with branches. That behavior i not typical for the group a a whole,
which the female place the egg. Buds of Q. dellla/Q open two for each pecies i u ually very particular about the place in
weeks later than those of Q. mOllgo/ica. Such timing often which the egg is laid by the female. All the e facts uggest that
determinestheecological ties between oak pecie and thein ects there could be everal sibling pecies involved in our ample.
feeding on them. To determine that definitively, one would have to rearseparately
Araragi ellthea ellthea (Janson) (Fig. 4(2». Larvae (Fig. 3A) eggsofdifferent ecotype andcomparepopulation ofbUllerflies
feed on Jug/ails mallchurica (Juglandaceae). The egg are laid obtained from each ecotype. The larva in Fig. 3F was photo
ingly, or ometimes in pairs on young shoots, from the ba e to graphed on acultivated species ofA/lUIS. In nature, however, the
the bud. Larvae, which are green in all ofthe in tar, hatch at the shape ofthe larva is congruent to the hape ofthe leafmargin, a
end of May and feed on the under ide of leaves until the end of feature which i typical for all the larvae of the pecies in the
June. Larvae pupate on the leaves. Adults fly at the end ofJuly. group. The la t instar larva rolls the leafand spends mo tofthe
Japolliea Llltea (Hewitson) (Fig. 4(5» and time in ide the resulting tube. It pupate on the lower surface of
Japolliea olloi Murayama (Fig. 4(6». The relationship between the leaf, or inside the several tied-together leaves. Adults hatch
the e twospecie resembles that between A. allilia andA. blll/eri. at the end ofJuly. They appear as the last species to fly in the
The differences in the e ympatric specie are ecological rather group ofgreen Theclinae in Primor'e.
than morphological. Japollica /urea lays egg (Fig. 6G) solitarily eozephyrllsultramarilllls IlLtramarilll1S (Fix en)(Fig.2(I». In
on buds ofyoung shoot ofQuercus mOllgo/iea (Fagaceae). The Fig. 3B, one can ob erve the pre ence of exual dimorphi m in
female ofJ. olloi lay eggs in a imilarmanner, but uses Quercus the larval stage: the female larva i darker than the male larva.
dellfata and depositsclu ters ofthree to 15eggs. In both species, Eggs are laid in the fold of the bark of thi k branche of
the female conceals the egg after it i laid by rubbing it abdo Quercusdelllata (Fagaceae). Egg (Fig. 5H)can beoccasionally
men against the freshly laid egg and masking it with hairs and a found on Quercus mOllgo/iea as well, e pecially in the areas
waxy ecretion. It makes it almo t impo ible for us to find the where Q. delJlata i ab ent. Thi i a typical case of "areal food
egg on the similarly colored, hairy shoots of the foodplant. plant" (a term proposed by Leonid ikolaevskiy, per. comm.).
However, these cryptic eggs do not e cape parasitic wasps: 40% When a bUllerfly evolve together with its food plant, it acquires
ofthe eggs collected in nature were parasitized. These observa extreme ecological and biochemical compatability with it. A
tions aresimilarto thoseofSaigusa(1993), whostudied Japane e new, closely related plant might ub equently move into thearea.
populations ofthe e species. We support hi view ofthis group Thi new planl doe not always become a new fo dplant of our
as being differentiated on the basi of ecological rather than bUllerfly,even though it larvaearepotentiallycapableoffeeding
morphological characters. A larva ofJ. olloi i een in Fig. 3M. on it. It might be u ed by a bUllerfly as an alternative source of
Vol. 2 o. I 1995 DA TCHE KO et al.: Egg of Ru sian Theclinae 33
food in case ofacrisis with the original foodplant population. A which ha red and blue patche on the front wings. 0 type
imilar example of such an occurren e in onh America would specimens are known. The pecies described as C. smaragdill/ls,
be the Schaus swallowtail (Papi/io aristodemus pOllceallus C. brillialltilll/s and C. korshullovi all fit the original de cription
Schaus) (papilionidae) in southern Aorida. BUllernies of thi ofC. aurorinus. Thu . C. brilliantinus is not even mentioned in
specie normally lay egg only on torchwood (Amyris e/emifera) Tuzov's (1993) list becau e its taxonomic tatus i not clear. In
(Rutaceae), which i rather rare in its u ual hardwood hammock ouropinion, it is a good species, who e name hould be revived.
habitat. Larvae are capable of feeding on wild lime (Zalltho Ussuriana michaelis michaelis (Oberthiir). Larvae feed on
xylem Jagara) (Rutaceae), present in the area in abundance. Fraxillus rhYllchophylla Hance (Oleaceae). Egg (Fig. 6E) were
However, wild lime i utilized as a foodplant in nature only obtained under laboratory conditions. The female' ovipo itor i
occa ionally. twice a long a its abdomen and normally i hidden inside the
eozephyrus quercus (Linnaeus) (Fig. IC and 4(4». Thi abdomen. Before laying an egg, a female look for a deep gap
specie is found in the European pan ofRussia. The larva feed in the bark. She in ert her ovipositor into thi cavity and lays
on Quercus robur L. (Fagaceae); the female lays olitary egg, an egg. The last in tar larva of a Chine e population of this
usually in thecorner formed by the union ofthe bud and thestem specie is illustrated in Koiwaya (\993). We think that the
(Fig. 71). The fir t instar larva bores into the bud, which i till population examined by Koiwaya is imilar to our population.
closed. In laterin tar, the larvae(Fig. 8B)strongly resemble the Adults ny at the end ofJuly, feeding on Umbelliferae. BUller
scales of the bud and stay next to them. Flight period ofadults nies spend their non-feeding time in the canopy.
varies geographically, but u ually occurs in the econd halfofthe Shirozua jonasi (Janson). According to our observation,
summer. In the present work (sec following), this specie i females lay eggs alway on the ant paths on the trunk ofQuerclls
compared with its eastern analogous species, NeozephYl"LIs mongo/ica (Fagaceae). Larvae of the last instar feed on aphids
saphirillus ( taudinger). on leaves and branche. They are colored purple-brown and
Neozephyrus saphirinus ( taudinger) (Fig. IA) is very similar con tantly are tended by ants. Placed in the cage together with
to N. quercus in appearance and biology (Fig. 2(3) and Fig. 3 ). larvae of other specie of butternies, however, the larvae of
The food plant ofN. saphirillus is Quercus delltata (Fagaceae), Shiro::'/la jonasi attacked molting and sick individual and con
which leafs out two weeks after the leaf-sprouting ofQ. mOllgo/i sumed them. Adult hatch in the fir t ten days ofAugust.
».
ca, a pecie analogous in phenology to the food plant of N. Thecla betu/acrassa Leech (Fig. IHand Fig. 4(I Larvae feed
quercus in Europe. So despite the morphological re emblance on Prulllls padlls L. and PrIIllllS spinosa L. (Rosaceae) in the
between the two bUllerny pecie, N. saphirillus adults appear European part of Russia. Our population feed on apricot,
later in a panicular ea on due to their association with the Armelliaca mallchurica Maxim. (Ro aceae), and ometimes on
lower-leafing Q. dentata. The egg is shown in Fig. 5 . PI"LIIlUS asiatica Kom. Egg (Fig. 5C) are laid olitarily, or
eozephyrus aquamarilllls DubatoIov & ergeev (Fig. ID). ometime in pairson the tern orat the branchingpointofyoung
Egg (Fig. 5 ) were obtained from a live female and rai ed on plant, or ometime on the lower branche of the older tree .
European Quercus robur (Fagaceae). We have not seen type The larva comes out of the egg, a in most member of this
material of thi species, so the identification of this specie is group, in early pring and feeds on the leave from ju t-opened
ambiguou. The population of female adults obtained from our buds. Later instar feed ,on the leave, sitting on their upper
laboratory rearing is very close in appearance toN. aquamarinus. urfaces. The mature larva (Fig. 3K) pupates in the liller on the
However, malesare 0 unu ual that theydo not fit thede cription ground. This pecie occur all over Ru ia, but the subspecies
ofany known taxa, including N. aquamarillus. Their genitalia, T. b. crassa i restricted to the Amur and Ussuri di tricts.
however, fit the de cription of N. aquamarillus. Under the TheclabelL/lillaStaudingerissimilartoT. bew/amorphological
laboratory condition, female laid eggs primarily on the buds. ly and biologically. The larva(Fig. 3E) feeds on Ma/us manelll/
Fir t instar larvae are similar to other species ofthis group. The rica Maxim. (Rosaceae) and, starting from thesecond instar, rolls
la't instar larva (Fig. 3C) is dark grey, and most ofthe time this the leaf, forming a tube. inside which it spends mo tofthe time.
larval stage stays on thick branches, feeding only at night. The larva pupate on the ground. Adults ny in August in the
Neozephyrus sp. This species is clo e to Favollit,s macrocercus understory on the edge of the forest. Fig. 6J hows the egg.
Wakabayashi and Fukuda from Korea, but that pecies ha not Artropoetespryeripryeri Murray lays egg in groups of3-5 on
been fund on the territory of Russia. Adults and immature old hoots of I-3m high Syrillga amurellsis Rupr. (Ro aceae).
tage ofour population are significantly different. The discus Adults ny in early July in open areas of the fore t.
sion of the taxonomic status of this species will follow. Eggs
werecollected at theedgeofthe fore ton large, 7-8m high, tree DlSCU SIO OF EGG DlFFERE CES
ofQuercus mOllgo/ica (Fagaceae). Theegg is hown in Fig. 6L.
Chrysozephyrus brilliLlntinus (Staudinger). The eggs are laid In thi review of a eries of clo ely related Theclini pecies
solitary on the nower buds of Quercus //Iongo/ica (Fagaceae) in from the Primor'e region ofRussian Far East, we found that the
the open areas and in the forest. The larva (Fig. 8C) of thi egg morphology i the mo t u eful character for differentiating
pecie feed on oak eventhough other Theclini species of this and grouping pecie. We u ed a Scanning Electron Microscope
group(e.g., C. smaragdinL/s. C. yoshikoae. C. /inae, and C. gaOl), ( EM) to examine the egg surface and micropylar region in
are found on Ro aceae. Chrysozephyrus brilliami/llls, described detail. The following imilarities and differences were found.
by Staudinger, was ynonymized by Shirozu and Yamamoto Echinoid eggs, with sharp pointed spine and bearing aero
under the name C. aurorillus (Oberthiir). Chrysozephyrus pyles (Fig. 7A) on circular rib, are found in the following
allrorillUS wa de cribed from Oskold Island from a female, species: Neozephyrus quercus (Fig. 5E, O. 5mm), W. signata
PL TE5.Micrographsofeggs(magnification: IOOx).(A)AlI/igiusblllieriFenlon:(B)A.atriliaBrelller:(C)Theclabe/ulaeL.:(D)Chrysazephyrusbri//ialllinusSlaudin
ger;(E)NeazeplryrusquercusL.:(F-G) IVagilllosigllalllquercil"OraSlaudinger:(H)Neozeplryrusui/mlllarinusFixsen;(I)S/rJlllollidiaeximiaFixsen;(1)Neozeplryrus
taxi/a Bremer: (K)N. aquamarinus Dubalolov & Scrgeev:(L) N. saplririllusSlaudinger.
Vol. 2 o. I 1995 DA T HE KO el al.: Eggs of Ru sian Theclinae 35
(Fig. 5F, 0.95mm), N. u/lramarillus (Fig. 5H, 0.89mm), N. laxi/a which he proposed on the ba i ofegg tructure corre pond 0
(Fig.5J, 0.95mm), N. aquamarilllls (Fig. 5K, 0.90mm), N. well with modern systematics that it is almost inexplicable why
saphirillus (Fig. 5L, 0.75mm), N. japollicus (Fig. 6B, 0.70mm), egg structure is not commonly u ed in Lycaenidae y tematic .
and in the species nova of Neozephyrus (Fig. 6L, 0.90mm). In Doherty wrote: "When the ovation i tudied, the e genera fall
N. quercus, though, the spines are wider and Ie pointed. The intoconvenientgroups,defined bycon tant and peculiarforms of
micropyle of the e egg are 4, 5, or 6-petaled ro ette (Fig. egg. I am aware that this classification i not likely to become
7E,F,G). a popular one: the student will always prefer to separate his
Theclo est in appearance to thi group i theegg ofChrysoze genera by an artificial key based on venation. But that the e
phyrllsbrillialllilllls (Fig. 50, 1.10mm). However, the tubercule groups of mine are natural one, and indicate in mo t ca e the
there are not pointed. The micropylar area i a 6-petaled roseUe truth ofdescent, I do not doubt."
situated in the crater, not porous and situated in the middle as in Speciesstudied in the pre entwork fall nicely intothreegroup
other specie. The egg is larger than in other pecies. Cell are identified by Doheny:
tetragonal in all ofthe above specie. I. Arhopala (eggs with tetragonal sculpture):
In Antigillsalhilia (Fig. 5B, 0.87mm) and A. butleri (Fig. 5A, a. semispheric egg with strongly developed pines: S. iLicis, S.
0.85mm), tubercules become even wider and shoner, exposing eximia, S. lalior, N. laxila, N. japonicus, N. ullramarinus, N.
quercus, N. saphirinus, N. aqllamarinus, N. sp., W. signala, C.
rounded p rous hexagonal cell (Fig. 7H). Eggs in these specie
brilliafllinus.
are much nauer.
b. disc-shaped egg with well developed ribs: U. michaelis, S.
Absence of pine is ob erved in Thecla belulae (Fig. 5C,
w-album.
0.90mm) and Thecla belulilla (Fig. 6J, 1.0mm). The micropyle
2. Aphnaeus/Loxura (eggs with hexagonal sculpture)
in T. belu/ae is a 3-petaled opening in the middle of the porous a. semispheric egg with well developed ribs: A. aI/ilia, A. bUlleri,
ro eUe, with pores hidden in the deep inclined pockets (Fig. 7C). A. ell/hea, T. bellllae, T. bellliina.
Cells are hexagonal. b. smooth, di c- haped egg: J. IlIIea, J. onoi, J. sapesiriata.
Theegg ofAraragi elllhea i very di tinct (Fig. 6C, O. Omm). 3. Thecla (trigonal or isometric hexagon): S. jonasi, F. herzi, F.
The chorion bear long tubercule with wide triangular tips, and pmni.
with the aeropyle opening in the middle. The micropyle is a Itseem remarkable that 0 many pecic are 0 similarin their
6-petaled ro eue with pore similar to tho e in T. bell//ae (Fig. lifecycle. We found egg ofeightspecies ofTheclini on asingle
7B). Cell are hexagonal. tree of Quercus lIIolIgolica. Thi i an excellent example of a
Egg of Ussllrialla michae/is (Fig. 6E, 0.63mm) are spineless, highly pecializedcommunity, inwhichevery pecie i very well
with ribs and tetragonal porou cells. The micropyle is not adapted to its particular niche. Thi excellent degree ofadapta
distinct. tion can be een in the oviposition preferences ofevery specie,
A imilar, only larger egg, is found in Fixsel/ia prlmi (Fig. 6F, regarding the place where the egg is laid on the branch, as well
0.90mm. The egg of F. heni (Fig. 6K, 0.93mm) is much more as the di tance from the ground of the branch chosen for
di tinct, with candle- haped tubercule on the cro - ections of ovipo ition.
the ribs, bearing aeropyle and a ro eue-shaped micropyle. Cell The larval behavior is even more pecialized, with the variou
are mo tly triangular and highly porous. taxa falling into distinct groups: larvae feed at night (N. orienla
The egg ofJapolJica /utea i pineless with hexagonal, highly lis) or during the day; larvae feed on the nower (w. signata, N.
porou cells (Fig. 6G, 0.93mm). The micropyle i a 5-petaled laxi/a, C. brillialllil/lIs, N. oriel/la/is) or on the leaves (A. atti/ia,
roseUe. A. blllieri, J. /lIIea); when feeding on the leave, larvae occupy
Long and sharp pine found on the egg ofSlrymollidia latior upper ide (T. bell/lae, F. prul/i, S. ilicis, A. allilia, A. blllieri) or
are typical for this group (Allyn, 1984). Unlike eggs of Neoze underside ofthe leaf(J. hllea); orne larvae are a ociated with
phyrus, StrymolJidia egg have high ribs with numerou harp ants and aphids (S. jOl/asi), while others feed olitarily.
spines, which do not seem to bearaeropyles. Tetragonal cells are We lack dctailedcomparativeinformationon theadult biology,
porcle s. The micropyle i a4-petaled rosette (Fig. 60, 0.80mm). but from what we have observed to date, there are certain
TheSlIymollidia eximia(Fig. 51, 0.63mm)egg is similarto the particularities in the type of the forest in which each of the
above, but ha a 5-petaled roseUe and horter spine. species is found, as well as in utilized nectar sources and the
Eggs ofStrymol/idia IV-a. w-album (Fig. 6H, 0.86mm) and S. night period.
IV-a. sUlschal/i (Fig. 61, 0.86mm) are very di tinct. The lauer
ACKNOWLEDGME TS
might be a good species, even based on this character alone.
Both specie' egg have low rib, with cell edges forming
We would like to thank L. ikolaev kiy for discu sion of
tetragonal ribs with low thin pines. TheeggofS. IV-a. slllschalli
taxonomy and biology of the group and for providing us with
i more rounded, with more distinct rib, than in the European
valuable information; .Sergeev, A. Chuvilin, V. Siniaev, and S.
ubspecie. The micropyle isa 5-petaled roseue.
Kunitsin for help in collecting the eggs; V. Konovalov for his
DISCUS 10 help in our housing at Riazanovka; and U. Eit chberger and O.
Gorbunov for their help in obtaining the literature. Chri tine M.
The first auempt to u e egg hape for y tematics was by Eliazar kindly prepared revi ions of the manu cript. Thi
Doherty (I 86). Unfortunately, his greatcollection wa lost. Hi research and its publication was supported by the Lepidoptera
approach to systematic was unconventional for the time, when Re earch Fund ofthe Univer ity of Florida Foundation.
sy tematics was ba edentirely on adult morphology. The y tem
PLATE6.Micrographsofeggs(magnification: 100x).(A)Neozephymsl/irralllarilll/sFixsen;(B)N.japollicl/sregillaButler;(C)AnmrgiemheaJanson;(D)SrrYlllollidia
lat;orFixsen;(E) Ussl/riOlUllllichaelisObenhtir:(F)Fixsell;apmlliL.:(G)JapOllictlll/rea Hewitson;(H)Srrylllollidia IV-a. IV-albl/III Knoch;(I)S. IV-a. sl/rschalliTut!;
(J) Theclabellllilla Staudinger: (K) Fixsell;a heniFixsen; (L)NeozephyrllS sp.
