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pidopterologica

A quarterly journal devoted to Palaearctic lepidopterology

Published by Societas Europaea Lepidopterologica

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NOTA LEPIDOPTEROLOGICA

A journal of the Societas Europaea Lepidopterologica

Published by Societas Europaea Lepidopterologica

Vol. 23 No. 1 Basel, 01.04.2000 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

Assistant Editors: Dr. Roger L. H. Dennis (Wilmslow, GB), Prof. Dr. Konrad

Fiedler (Bayreuth, D), Dr. Enrique Garcia-Barros (Madrid, E), Ole Karsholt

(Kabenhavn, DK), Dr. Yuri P. Nekrutenko (Kiev, UA), Dr. Erik J. van

Nieukerken (Leiden, NL), Dr. Alexander Pelzer (Wennigsen, D)

Contents @ Inhalt e Sommaire

Fric, Z. & Konvicka, M. Adult population structure and behaviour of

two seasonal generations of the European Map Butterfly, Araschnia

levana, species with seasonal polyphenism (Nymphalidae) ................. 2

RuTJAN, E. V. A new bagworm species of the genus Dahlica (Psychidae)

MammanaMinoastennyWkrainecs),{., 0 mas an era Ads ae 26

LOELIGER, E. A. & KARRER, F. Unusual demonstration of autosomal do-

minant inheritance of the black coloration of one of America’s swallow-

tails: F, broods of the hybrid Papilio polyxenes asterius with Papilio

DRIEERSOTEOmaSs (PADINONIGAC) ........................00.0000200 000. 40

PLJusHcH, I. G. & DoLinskAYA, I. V. Eggshell fine structure of some

species of Lithosiinae (Arctiidae) of Far East Russia ......................... 50

STEFANESCU, C. New data on the ecology of Thecla betulae in the north-

Edsuoiminedbeman Peninsula (Lycaenidae) ...................................... 64

PUPLESIENE, J. & OLIVIER, A. The karyotype and chromosome number of

Lomvonmmams buzulmavi(lycaenidae) .+,2.............1................... 71

HmemonamAxelScaoLz1(1957-1998) 0... turn. 78

BOOK REVIEWS @ BUCHBESPRECHUNGEN @ ANALYSES .......................... 81

Nota lepid. 23 (1): 2-25; 01.1V.2000 ISSN 0342-7536

Adult population structure and behaviour

of two seasonal generations of the European

Map Butterfly, Araschnia levana, species with

seasonal polyphenism (Nymphalidae)

Zdenék Fric* & Martin KONVICKA**

* Department of Zoology, Faculty of Biological Sciences, University of South

Bohemia, BraniSovska 31, Ceské Budéjovice, CZ-37005 Czech Republic

(contact address)

E-mail: zdenek.fric@tix.bf.jcu.cz

** Department of Botany, Faculty of Sciences, Palacky University Olomouc,

Czech Republic

Summary. Adult population structure and behaviour of the spring and summer broods of

Araschnia levana (Linnaeus, 1758) were studied in order to describe differences in eco-

logical traits between seasonal generations of this polyphenic species. The more numer-

ous summer generation had an unambiguously open population structure, while the pop-

ulation structure of the spring generation tended to be closed. Capture and estimated sex

ratios were male-biased in both generations, probably owing to lower site tenacity of

females. There were marked diurnal changes in behaviour of both generations. Both sexes

fed on flowers during the mornings and late afternoons, while in the early afternoons

males perched in aggregations with some features of typical leks. Females were only

rarely seen during that time, their appearance provoked males to chase them. Butterflies

of the summer generation spent more time nectaring, possibly owing to higher energy

requirements associated with higher mobility. Males of the summer generation spent sig-

nificantly more time flying or chasing as opposed to sitting on their perches, but this

might have been a recording artefact. Seasonal alternation of dispersing and sedentary

generations, combined with a lek assembly, might be the features which allow the species

to respond rapidly to environmental changes.

Zusammenfassung. Struktur und Verhalten adulter Populationen von Frühjahrs- und

Sommergenerationen von Araschnia levana (Linnaeus, 1758) wurden untersucht, um

Unterschiede hinsichtlich 6koiogischer Parameter zwischen den saisonalen Generationen

dieser polyphenetischen Art zu beschreiben. Die individuenreichere Sommergeneration

zeigte eine eindeutige offene Populationsstruktur, während die Friihjahrsgeneration zur

geschlossenen Populationsstruktur tendiert. Gesammelte und geschätzte Geschlechts-

verhältnisse waren von Männchen dominiert, wahrscheinlich aufgrund geringerer

Ortstreue der Weibchen. Es gab signifikante Unterschiede im Tagesrhytmus beider

Generationen. Beide Generationen erscheinen in den Morgenstunden sowie am spaten

Nachmittag zum Nektarsaugen an Blüten, während sich die Männchen am frühen

Nachmittag in typischen Lek-artigen Aggregationen niederlassen. Weibchen wurden zu

dieser Zeit nur selten beobachtet, ihre Erscheinung provozierte die Männchen sie zu

jagen. Die Falter der Sommergeneration verbrachten mehr Zeit beim Nektarsaugen,

möglicherweise um größere Energieressourcen anzulegen, die mit einer höheren

Mobilität korrelieren. Männchen der Sommergeneration verbrachten signifikant mehr

Zeit fliegend oder jagend anstatt sich niederzulassen, aber dies kann auch ein

Beobachtungsartefakt sein. Saisonale Änderungen sich ausbreitender und seßhafter

Generationen, kombiniert mit Lek-Ansammlungen, mögen die Voraussetzungen für diese

Art sein, rasch auf Umweltänderungen zu reagieren.

Résumé. La structure et le comportement des populations d’adultes des générations prin-

tanières et estivales de Araschnia levana (Linnaeus, 1758) ont été étudiées afin de décrire

les différences dans les affinités écologiques entre les générations saisonnières de cette

espèce polyphénique. La génération estivale, qui est la plus abondante, a, sans ambiguité,

une structure ouverte alors que celle de printemps tend à rester fermée. La capture et la

sex ratio des mâles est biaisée dans les deux générations, probablement en raison de la

ténacité des femelles à occuper les sites bas. Il y a des changements diurnes marqués dans

le comportement de chaque génération. Les deux sexes s’alimentent sur des fleurs le

matin et tard dans l’après-midi, alors qu’en fin d’après-midi les mâles s’agrègent et se

perchent avec quelques aspects comportementaux typiques. Les femelles sont rarement

aperçues durant cette période, leur apparition conduisant les mâles à les chasser. Les

papillons de la génération estivale passent plus de temps à s’alimenter, probablement en

raison de leurs besoins de resources en énergie plus intenses liés à leur plus forte mobi-

lité. Les mâles de la génération d’été passent significativement plus de temps à voler ou

patrouiller plutôt que sur leurs perchoirs, mais ceci pourrait être un artéfact d’observation.

L’altération saisonnière de dispersion et de sédentarité des générations, associée à un

comportement de rassemblement, pourraient représenter des caractères permettant à

l’espèce de répondre rapidement aux changements de l’environnement.

Key words: Araschnia levana, adult population, butterfly dispersal, lek, male assem-

blages, mate-locating behaviour, seasonal polyphenism.

Introduction

Seasonal polyphenism in butterflies is an adaptive trait, which

allows species living in areas with changing but seasonally pre-

dictable climates, to avoid costs of genetic polymorphism while

maintaining benefits of two adaptive strategies (Shapiro, 1976,

1980). The European Map Butterfly Araschnia levana (Linnaeus,

1758) (Nymphalidae) is a model example of seasonal polyphenism

in butterflies. Adults of the spring generation (f. levana) are reddish

with a fritillary-like black pattern, whereas the butterflies of the

summer generation (f. prorsa) have white stripes on a black

ground-colour and resemble black admirals (Limenitis camilla

(Linnaeus, 1764)) (Shreeve, 1992a). An occasional third (autumn)

generation (referred to as f. porima or f. intermedia) is intermedi-

ate between the two. Proximately, the polyphenism is controlled by

day length together with temperature and other cues which change

regularly with time of year (Müller, 1955, 1956; Shapiro, 1976).

The species ranges throughout the Palaearctic, reaching Japan

and the Russian Far East to the east (Korshunov, 1972; Korshunov

& Gorbunov, 1995). The western edge of its distribution is situated

in France, where the species ıs expanding westward and has

reached the Atlantic coast but recently (Radigue, 1994-1995). In

southern Europe, there are isolated populations in Spain (Higgins

& Riley, 1980), the southern limits of the continuous European dis-

tribution of the species are found in northern Croatia and Romania

(Niculescu, 1965) down to northern Greece (Koutroubas, 1991).

The butterfly ıs relatively common in Central Europe. However, it

is not as widely distributed as other nettle-feeding Nymphalids.

The preferred habitats are wet meadows adjoining rivers and

streams, river valleys and moist deciduous forests (Hruby, 1956;

Reinhardt & Richter, 1978, Ebert & Rennwald, 1991); it was

reported that the species avoids early successional habitats

(Steffan-Dewenter & Tscharntke, 1997).

Larvae of A. levana feed on nettles (Urtica spp., in Central

Europe only Urtica dioica). They live gregariously after emerging

from the eggs, which are laid in a unique string-like pattern (Porter,

1992) on undersides of nettle leaves. The overwintering stage is the

pupa.

Published studies of seasonal polyphenism in A. levana have so

far focused on the developmental mechanism of the phenomenon

and its relation to temperature change (Süffert, 1924), length of day

(Müller, 1955, 1956; Reinhardt, 1969, 1984) or biochemical factors

(Koch, 1996). On the other hand, field-oriented studies of the

species’ population ecology and behaviour are practically missing.

The few published papers did not study the seasonal generations

comparatively. Instead, they focused on habitat selection or climat-

ic requirements of A. levana in comparison with closely related (in

both taxonomic and ecological sense) species (Reinhardt &

Richter, 1978; Reinhardt, 1984). The European Map Butterfly thus

remains a surprisingly enigmatic species despite fairly good know-

ledge of proximate mechanisms of its polyphenism, extensive

research conducted on other European nettle-feeding Nymphalids

(Bryant ef al., 1997, and references therein) and at least two

authors’ (Brakefield & Shreeve, 1992) discussion of possible

rewards of such studies.

We attempted to fill this gap in our knowledge and stated the fol-

lowing aims: (1) to compare adult population size and structure,

mobility and other parameters of the two seasonal generations of A.

levana, (2) to describe the behaviour of adults of A. levana in field

conditions, to study their distribution, mating activities and changes

of behaviour with time of day and (3) to look for differences, if any,

between the spring and summer generations.

Fig. 1. Map of the study site showing the meadow (white), surrounding woody vegetation

(light grey) and nettle patches (dark grey). The black line shows the position of the tran-

sect route; letters A, B, C, D are the subsites which were recognised for analysis. The grey

circle is the large lime tree overseeing the meadow.

Methods

The lack of field research conducted so far on A. levana could

possibly be explained by methodological problems inherent to

working with common butterflies with open populations (see

Shreeve, 1995). Given our previous experiences with A. levana and

published information on closely related butterflies (tribus

Nymphalini) (Baker, 1972; Warren, 1992; Shreeve, 1995), we

expected an open population structure. However, many of the

methods used in this work are typically applied for butterflies

which live in spatially restricted (closed) colonies (sensu Warren,

1992). When applying these methods, we understood this limita-

tion but believed that our approach might at least indicate the most

important trends, ıf not bring complete answers to the questions

under study.

The fieldwork was carried out in the spring and summer of 1997,

with additional observations in 1998. The study site (Fig. 1) was a

moist meadow with dominant grass Alopecurus pratensis, 1.5 km

south of Hluboka nad Vitavou, South Bohemia (49°02’ N, 14°28’ E,

375 m). The locality is situated within a varied landscape of mead-

ows, fishponds, hedgerows and moist forests known as the Ceské

Budéjovice Basin. The meadow was surrounded by habitats appar-

ently unsuitable for A. levana: there was a mature oak-spruce fo-

rest on two sides, on one side there was a fishpond bank overgrown

by reed, one side was delimited by a dense hedgerow along a water

channel. Nettles (Urtica dioica) formed several distinct patches

within the study site. We divided the meadow into four subsites

which differred in the character of vegetation and distribution of

nettles. Subsite A was a meadow-forest margin oriented to the

South with nettles growing under the canopy; subsite B was a large

nettle patch along a fishpond bank covered by reed; subsite C was

in the centre of the meadow with nettles in sparse bunches inter-

spersed with tall grasses; subsite D was a border between the mead-

ow and a hedgerow with a large lime tree dominating the woody

vegetation.

We applied three methods in the field. Population size and struc-

ture were estimated by (1) standard mark-release-recapture (MRR)

techniques, (2) behaviour and diurnal activity were assessed by reg-

ular observations along a fixed transect and (3) by following of

individual butterflies with portable tape-recorders.

For the MRR study, the butterflies were netted and marked by

waterproof alcohol-based pens. We used the 1-2-4-7 system

(Ehrlich & Davidson, 1960) with minute modifications. Wing wear

(four categories as in Warren, 1987) was recorded for each captured

butterfly.

Progressive estimates of butterfly numbers, obtained during one-

day marking (Thomas, 1983), were calculated in order to detect

whether the population of A. levana was closed or open. The results

of such estimates should gradually improve in accuracy in a closed

population, but increase meaninglessly if the population is open.

The estimates for individual time intervals were obtained according

to Craig (1953). The analysis was performed only on fine days,

when the weather allowed continuous marking from dawn to dusk

(May 17, 18 and 24, 1997 and May 9, 1998 for the spring genera-

tion and July 17, 28, 30 and August 5, 1997 for the summer gene-

ration).

Population sizes were estimated separately for generations and

sexes. Numbers of individuals were calculated by the Jolly-Seber

method, which allows for open populations (Jolly, 1965).

Computing was done by program JOLLY, model A (Pollock et al.,

1990). Residence times were calculated using the formula — In(@)

(Cook, 1967; Tabashnik, 1980), mean residences per generation

@,’s were obtained by the variance weighted averaging method

(Tabashnik, 1980). Tabashnik’s (1980) joint residence-catchability

test and capture vs. estimated sex ratio test were performed in order

to detect whether differences in numbers of males and females

were caused by different residence or catchability of sexes. The

joint residence-catchability test plots the natural logarithm of

recapture probability ratio against time and allows a distinction

between the effects of unequal residences and unequal recapture

probability (between sexes). The capture vs. estimated sex ratio test

estimates relative catchability of males and females for a sampling

day and gives an average estimate for a set of dates (for details of

the tests, see Tabashnik, 1980; Gall, 1984; Murphy er al., 1986).

Behaviour was observed along a fixed transect route (about 400

m), which crossed the entire meadow (Fig. 1). We walked along the

transect three times per hour and recorded the behaviour of all

observed butterflies and the subsite where they were seen. We dis-

criminated four subsites (see above) and seven types of behaviour:

1 — nectaring, 2 — flight, 3 — perching on herbaceous vegetation, 4

— perching on trees/shrubs, 5 — chasing with other butterflies, 6 —

any behaviour unambiguously connected with reproduction (copu-

lation, egg-laying etc.), 7 — resting. By definition, only males could

have been included in the “perching” categories. A butterfly was

considered as “perching” when sitting on tops of herbaceous vege-

tation or at distal ends of tree/shrub branches; males sitting in lower

layers of vegetation were listed as “resting”. The data were

analysed separately for generation, sex and part of day (morning

vs. afternoon). Altogether, 70 transects were walked for the spring

generation (May 15, 16 and 19, 1997) and 78 for the summer gen-

eration (July 17 and August 6 and 10, 1997).

Early in the course of the study, we noticed an aggregation of

perching males regularly occurring at subsite B during the after-

noon hours (see Results section). In both generations, we followed

the males with a tape-recorder. We picked-up a sitting male and

recorded its behaviour for the longest possible time, 1.e., until the

observer lost contact with the animal. The longest recorded obser-

vations lasted 837 and 641 seconds for males of spring and summer

generations, respectively. The recorded observations were subse-

quently split into 2-minutes intervals for further analysis, and the

observations shorter than 120 seconds were discarded. Flight, chas-

ing, resting and basking were distinguished. The tape-recorded

behavioural categories differred from those used for the transect

analysis, because it was not possible to distinguish between “bask-

ing” and “perching” during the transect walks, while during the

tape-recording we classified sitting with opened wings as “bask-

ing” and sitting with closed wings as “resting”. (Also note, that the

entire activity observed with tape recorders was perching.) For the

spring generation, observations were carried out on May 17 and 23,

16 observations were analyzed and 49 discarded. The summer gen-

eration was observed on July 17 and August 3. Sixteen observa-

tions were analysed and 24 discarded.

All behavioural data were log-transformed to achieve normality

and analysed using the Statistica v. 5 (StatSoft Inc.) package.

Results

Population size and structure. The first individuals of the spring

generation were observed on May 3 in 1997, the last specimen flew

on May 28. The summer generation was on the wing from July 7

until August 28 in 1997. The progressive one-day estimates of po-

pulation size (Fig. 2) indicated that the butterflies of the summer

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Fig. 2. Progressive one-day estimates of population size of the two seasonal generations

of À. levana, With their respective standard errors. Left: spring generation, right: summer

generation. Note that y-axes are not in scale.

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generation had a typical open population structure. The situation

was more ambiguous for the spring generation, for which two rep-

etitions of the experiment indicated a closed and two repetitions an

open population structure.

Table 1 summarises daily catches of butterflies for MRR analy-

sis. The Jolly-Seber estimates of daily numbers (Fig. 3) revealed

that the adults of the summer generation were more abundant than

those of the spring generation. Capture sex ratio males:females was

male-biased in both generations (deviation from 1:1 ratio, spring:

Xa = 45.2, p < 0.001; summer: X’, = 1.64* 107, p << 0.001). The

estimated sex ratios were also heavily male-biased (spring: x’, =

JE NO p <= 0.001;-summer: Yo = 5.3710", p << 0.001). The

prevalence of males did not change during their flight periods in

both generations. However, the estimates for females, and the sex

ratio estimates, were highly unrealistic due to low capture and

recapture rates (see below).

There were no significant differences in single-day residences

between males and females within the two generations (paired t-

test, spring: t« = 1.13, p = 0.30; summer: to, = -1.05, p = 0.33). The

residence statistics (Table 1) reveals shorter residence of males of

summer generation. The joint residence-catchability tests were not

significant for either generation. (Residence time males vs.

females, spring: F14 = 0.003, p = 0.96; summer: F,:, = 0.96, p =

0.40. Relative catchability males vs. females, spring: t« = 1.63; p =

0.18; summer: t:, = 2.41; p = 0.09.) Therefore, the tests failed to

distinguish whether the biased sex ratios were due to unequal resi-

dences or catchabilities. There were no significant differences with

regard to residence and catchability between the generations

(ANOVA comparison of the regression lines from the joint resi-

dence - catchability tests, residence: Fi.) = 0.187, p = 0.68; catch-

ability: Fis) = 0.043, p = 0.84). The capture vs. estimated sex ratio

test did not reveal any significant difference in catchability among

sexes of the spring generation (77) = 0.25, p = 0.62), while males

of the summer generation were significantly more catchable than

females (771) = 106.04, p < 0.001).

Wing wear was steadily increasing in spring males, spring

females and summer males. In these cases, the increases were sta-

tistically significant (slopes of the regression lines of average wing

wears plotted against time, spring males: F 44, = 16.73, p < 0.05;

spring females: Fis = 13.59, p < 0.01; summer males: Fa.) =

132.42, p < 0.01). On the other hand, no significant trend was

found for the wing wear progression of summer females (Fa, ı» =

2.76, p = 0.12).

Table 1. Summary of MRR catches and residence statistics of Araschnia levana.

@;s are the estimated residences for individual periods, @,s are the mean resi-

dences per generation obtained by the variance weighted averaging method.

Div/O denotes the periods for which the residence was not computable from the

data.

Males Females

captured recaptured captured recaptured @:

spring generation

5 May

6 May

15 May

16 May

17 May

18 May

19 May

20 May

24 May

26 May

Anne ee

(Oo U à = Lo a ow

ND © R © © NN © Un

BR = NN 0 BP = D = ©

mean residence (@,)

mean residence time (days)

Summer generation

16 July

17 July

23 July

26 July

28 July

30 July

4 August

5 August

7 August

8 August

9 August

11 August

12 August

13 August

OSCO0-c00000

NDANMNABRHNOOBYNDA

NK ON ON © DD ON BR NN 00 -J ~

NN = WK © OONn OCF OO

mean residence (@,)

mean residence time (days)

©.

Behaviour, distribution. In both generations, there were marked

differences in behaviour at different times of the day (see Figs 4, 5

for illustrative examples). Both sexes spent the morning hours nec-

taring on flowers, whereas most of the individuals disappeared

from the observers’ sight during noon (approximately from

12:00 AM till 1:00 PM) and were spotted only occasionally in the

canopy of surrounding trees. Later, a distinct assemblage of males

was daily observed on the sparse nettle bunches in the central part

of the study site (subsite C). The males perched with closed wings

on top of both nettle plants and grass blades (which were often top-

ping the nettles). The average number of males which were seen

perching in this site per transect was 5.3 in spring (40 “afternoon”

transects, N = 223, SE = 2.14) and 6.4 in summer (44 transects, N

= 275, SE = 3.98). Mutual chases among the males were frequent

(spring: 86 chases tape-recorded; summer: 46 chases tape-record-

ed). The males often attacked other flying insects (one dragonfly,

two bees, 14 peacocks — Inachis io (Linnaeus, 1758) — and 14

pierids). During this time, the females were practically absent from

the meadow. If an occasional female showed up near the aggrega-

tion of males (four such observations were tape-recorded), her

presence provoked repeated bouts of chasing, in which several

males participated. The butterflies flew in ascending spirals during

such chases, reaching about 10 m height above the meadow.

Afterwards, the males either returned to the perching site or disap-

peared from sight.

Table 2. ANOVA results for behaviour of A. levana butterflies along the fixed

transect.

Factor, interaction

Generation

Sex

Behaviour

Time of day

0,74; NS

<< 0,0001

OSE NS

<< 0,0001

0,43; NS

<< 0,0001

Generation-sex

Generation-behaviour

Generation-time of day

Sex-behaviour

Sex-time of day

Behaviour-time of day

ON ON ON ON ON FF ON ni

w w ” M ” M w w

ee ee ee femnh

ph es

”

In late afternoons, both sexes were again seen nectaring on flow-

ers. On fine sunny days, the last individuals were nectaring until

sunset.

Four-way analysis of variance of the behavioural observations

from the transect (with the number of individuals as the frequency

variable) (Table 2) confirmed the diurnal periodicity in behaviour

and distribution of sexes. Sex, type of behaviour and time of day all

had a significant effect on the number of individuals observed on

the transect. Furthermore, the combined influences of factors gen-

eration-behaviour, sex-behaviour, time of day-behaviour and time

of day-sex were also significant (Fig. 6). Butterflies of the summer

generation spent more time feeding; only males were engaged in

perching and chasıng; perching and chasing were significantly

more frequent during the afternoon hours.

Analysis of variance for distribution of butterflies within the

study site (Table 3) revealed that both generation and sex signifi-

cantly influenced butterfly distribution. Feeding subsites of the two

generations differred according to distribution of nectar plants;

males of both generations were more frequently encountered at

subsite C, where they formed the afternoon perching aggregation

(see above).

The analysis of tape-recorded behavioural observations revealed

that the summer generation males spent less time flying/chasing

and more time resting when compared with the spring generation

(one-way ANOVA, Fa 2) = 8.76, p << 0.001).

Table 3. ANOVA results for behaviour of A. /evana butterflies along the fixed

transect.

Subsite

Generation-subsite

Sex-subsite à ;

Time of day - subsite ‚56 5 ; 0,34; NS

Discussion

Adults of the summer generation of A. levana were more abun-

dant and less restricted in their movements. Both generations had

Spring generation - Males

May 16, 1997

IE Resting

I Reproductive

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ico) x N oO co co wv N oOo e [ep] Le] FF © w + [sp] N = oO

SIeNPIAIPUI Jo soquINN SJENPIAIPUI JO sequinn

Time

Fig. 4. Diurnal changes in activity of À. levana, adults of spring generation.

Summer generation - Males

August 6, 1997

f Resting

ff Reproductiv

Chasing

eee ee

tii NE

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Summer generation - Females

August 6, 1997

OTE

STEMI

TE TETE TENTE TETE TETE

IETBIIDENDELDUTERLEBDLIUDEDTDUUDDURUNNEN

[de] wo T oD

sjenpiaipui JO Jequinyy

(TETE EE TETE TETE EEE TETE TEE

Or:Z2l

02:21

00:2)

0r:91

02:91

00:91

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02:S1

00:51

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02:71

00:71

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0z:E1

00:E1

Or:cl

02:21

00:21

Ov: LL

OC:LL

00:L1

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02:01

00:01

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Fig. 5. Diurnal changes in activity of A. levana, adults of summer generation.

6a)

LN (Number of

individuals)

Nectar P-herb -Chasing Resting

Behaviour

6b)

OMales |

EFemales

LN (Number of

individuals)

Nectar Flight P-herb P-tree Chasing Reprod Resting

Behaviour

6c)

4 C} Morning

EJ Afternoon

LN (Number of

individuals)

Nectar Flight P-herb P-tree Chasing Reprod Resting

Behaviour

Fig. 6. Behaviour of A. levana adults as it was observed along the transect route (log-

scale), with respective standard error bars: a — behaviour observed in spring vs. summer

generations; b — behaviour observed for individual sexes; c — behaviour observed in morn-

ing vs. afternoon hours. Note the log-scale of y-axis.

male-biased capture and estimated sex ratios, their behaviour was

characterised by predominance of nectaring in the morning hours

and aggregate perching of males in the afternoon hours.

The lower abundance of adults of the spring generation suggests

that the overwintering generation of A. levana might suffer high

mortality during diapause. Host plants growing in spring usually

have a high nutrient content (cf. Slansky, 1993) and, as a result,

abundant progeny of the spring-flying adults replenishes the popu-

lation numbers. This might correlate with different mobility of the

two generations. Whereas the spring-flying adults might profit

from not leaving the site where they successfully completed their

development (such a site, by definition, provides all essential

resources for them), it may be beneficial for individuals of the sum-

mer generation to leave. Higher population density of the summer

generation may lead to competition for larval resources, this in turn

lowering the emigration threshold (Baker, 1984). Differing mobili-

ty among generations has been assumed for several butterfly

species and has been considered as one of the traits which allow

multivoltine butterflies to become widespread in cultivated land-

scapes (Pollard & Eversham, 1995).

Three, admittedly circumstantial, lines of evidence suggest a

higher mobility of the summer generation. First, the summer gen-

eration had a distinctly open population structure, while the situa-

tion was rather ambivalent in spring. Second, the females of the

summer generation were rarely recaptured, being significantly less

catchable than males. This trait caused the inflated estimates of

their abundance (see Fig. 3). Third, the females of the summer gen-

eration did not show a significant increase of wing wear with time.

This again suggests that they might not spend much time on the

study site and instead might move rather freely across the land-

scape. Nevertheless, none of the arguments can be considered as a

direct proof of higher mobility of the summer-flying adults, which

might be obtained only by direct mark-release-recapture work cov-

ering a broader array of habitat patches (Hanski ef al., 1994;

Sutcliffe et al., 1997; Baguette et al., 1998).

An intriguing problem, that recurrently emerges in butterfly pop-

ulation studies, is the male bias in both capture and estimated sex

ratios (Scott, 1974, 1975; Daily et al., 1991; Sutcliffe et al., 1997).

Sexual differences in both distribution and behaviour are reported

for many butterfly species (cf. Wiklund, 1977, 1982; Wiklund &

Ährberg, 1978; Baguette et al., 1998). The sex-specific differences

in both behaviour and distribution found in this study (Fig. 6, b) are

the most plausible explanation of the biased sex ratios in A. levana.

Males of both generations spent most of the days within the reach

of observers, whereas the females regularly “disappeared” for sev-

eral hours around noon. Furthermore, the males assembled at one

site in the afternoon hours, which rendered them easıly catchable.

On the other hand, the joint residence-catchability tests failed to

show significant differences between sexes ın both residence and

catchability. We assume that the failure was caused by the low

number of females recaptured, ın particular ıf more days elapsed

between the first and subsequent captures. Consequently, the resi-

dence-catchability regression lines were constructed from too few

points: compare the low number of degrees of freedom in our tests

with those in Tabashnik (1980) and Gall (1984). It is notoriously

difficult to separate different dispersal rates from different longevi-

ties (Tabashnik, 1980; Shreeve, 1992b, 1995); our residence statis-

tics (see Table 1) do not suggest any conclusive solutions. Murphy

et al. (1986) highlighted the necessity of nearly complete, labour

intensive marking of the entire population, to obtain accuracy of

such tests. This was admittedly not the case in this study, as the

MRR days alternated with days devoted to behavioural observa-

tions.

Brakefield & Shreeve (1992) suggested the “fascinating” possi-

bility that different wing colours of seasonal generations of A. le-

vana might result into different mate-locating behaviours. They

expected fritillary-like patrolling in the spring generation and

admiral-like perching in the summer generation (vividly coloured

butterflies tend to patrol rather than perch — Shreeve, 1992a). Our

observation showed that in A. levana, males of both generations

perch. Perching is restricted to the afternoon hours; in this respect,

the diurnal changes in behaviour of A. levana resemble those of

other European nettle-feeding Nymphalids (/nachis io, Aglais

urticae (Linnaeus, 1758) and Vanessa cardui (Linnaeus, 1758) —

Baker, 1972; Scott, 1974). However, early morning perching was

reported for some Nymphalids and the time of eclosion of females

was proposed as the explanation for the timing of activity of males

(Rutowski et al., 1996).

The perching activity in A. levana was characterised by the

aggregation of males within one patch of the habitat, while the rest

of the study site was virtually empty. Scott (1974) stated that perch-

ing should result to “distributing males more evenly throughout

mating sites” due to mutual harassment in male to male encounters.

In our study, nothing indicated any spacing apart — or territoriali-

ty — of males, which was described for other perching species (cf.

Baker, 1972; Davies, 1978: Dennis & Williams, 1987; Rutowski et

al., 1996). In agreement with Knapton (1985), we suggest that such

a situation might be interpreted as lekking, bearing in mind that the

term may be used only in a relaxed sense for non-vertebrates

(Lederhouse, 1982; Knapton, 1985; Alcock, 1987; Daily et al.

1991; Shreeve, 1992a).

Typical lekking assemblages of males should, according to the

original definition (Bradbury, 1977), occur at a place which lacks

any larval or adult resources. Furthermore, individual males should

monopolise exclusive patches within the lek and the males with the

best (typıcally central) positions should enjoy most of the matings

(Knapton, 1985; Wickman, 1985; Wickman & Jansson, 1997). The

situation was different in A. levana, where the aggregation occurred

at a foodplant patch. On the other hand, no other foodplant patch

within the study site hosted such an aggregation and males com-

peted for their perches as well as for females (e. g. the mutual pur-

suits). Also, we gained no evidence of females ovipositing (or

hanging around) at the aggregation site, at least during the after-

noon hours. Indeed, this would be disadvantageous for them due to

continuous harassment by males. Further study should focus on

elucidating the questions (1) where the ovipositions do actually

occur and (2) whether the males compete for their positions within

aggregation sites.

Although both the transect and tape-recorder results pointed to

behavioural differences between generations, the evidence is weak

and the interpretation of data is not straightforward. The behav-

ioural differences between the generations documented in the

analysis of the tape-recorder data may have been caused by the dif-

ferent colours of the seasonal morphs: the disruptive wing pattern

of the summer generation specimens might be responsible for the

fast disappearance of flying/chasing males from the observers’

sight. This alone would bias the data towards a higher proportion of

the “resting” category. Moreover, a climatic (possibly temperature)

element might have influenced the behavioural results and this

question deserves a further, temperature-controlled study. On the

other hand, the transect-gained observation that butterflies of the

summer generation spent more time nectaring seems to be robust.

This might be explained by higher energy requirements of the more

active summer generation of these ecothermic insects.

Pollard & Eversham (1995) speculated that widespread British

butterflies, such as nettle-feeding Nymphalids, benefited from

changes brought by modern agriculture (e.g. fertilizers) and

became more abundant recently than they had been in the past.

Recently, À. levana has indisputably become a common species in

Central Europe. Although our evidence is only circumstantial, it is

possible that the habit of alternating sedentary and dispersing gen-

erations, when the sedentary generation stays in more favourable

habitats and the dispersing generation leaves these in search for

new space, combined with a lek-assembly mating system (which

ensures that the sexes meet despite their dispersing habits), might

allow the species to respond rapidly to changing natural conditions

and to fluctuate in range and abundance. Such fluctuations are

indeed documented in literature (Hruby, 1956; van Swaay, 1990;

Benes & Kuras, 1997) and there is evidence of recent range expan-

sion and occasional migrations along the southern, western and

northern edge of the range (Koutroubas, 1991; Radigue,

1994-1995; Bradley, 1998).

Acknowledgements

We would like to thank J. BeneS, T. Kuras, K. Spitzer and

E. Tkadlec for valuable comments on earlier drafts of the manu-

script and to T. Grim, V. Novotny, and J. Zrzavy for consultations.

M. Duchoslav, A. Faltynkova and M. HaraStovä kindly provided

much-needed encouragement at some stages of the work. Two

anonymous referees much contributed to the clarity of the paper

and we are grateful to them. The study was partially funded by the

University of South Bohemia studentship to Z. Fric.

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Nota lepid. 23 (1): 26-39; 01.1V.2000 ISSN 0342-7536

A new bagworm species of the genus Dahlica

(Psychidae) from southeastern Ukraine

Evgeniy V. RUTJAN

Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine,

UA-01601 Kiev MSP, Ukraine

Summary. A new bagworm species, Dahlica karatyshica sp. nov., is described from the

steppe of Ukraine. It is characterised by its small size, the reduction of some veins, an

average genitalic index equal to 1.1 and a habitat preference for xerophytic vegetation,

which is not typical for other known species of the genus.

Zusammenfassung. Eine neue Sacktragerart, Dahlica karatyshica sp. nov., wird aus der

ukrainischen Steppe beschrieben. Charakteristisch für diese Art sind eine geringe

Korpergröße, die Reduktion einiger Adern, ein durchschnittlicher Genitalindex von 1.1

sowie die Anpassung an xerophytische Vegetation, welche in dieser Gattung

außergewöhnlich ist.

Résumé. Une nouvelle espèce de psychide, Dahlica karatyshica sp. nov., est décrite de la

steppe d’Ukraine. Elle est caractérisée par une taille réduite, la réduction de certaines

veines, un indice génitalique de 1.1 et une adaptation à une végétation xérophytique, qui

est atypique dans ce genre.

Key words: Lepidoptera, Psychidae, Dahlica, new species, steppe, Ukraine.

During two collecting trips in May 1997 and in April 1998 to the

virgin steppe of Kamennye Mogily Nature Reserve (SE Ukraine,

Donetsk oblast’ [province]), 38 males and 6 females (only two

female specimens are included in the type series — see below) of

a new bagworm species were found. One more male was captured

in May 1998 in the neighbouring Lugansk oblast’ (Proval’ska

Steppe Nature Reserve). By an aggregate of characteristics (lack of

epiphysis, lack of medial cells on both fore- and hindwing, female

pupal antennae equal to legs), the new species is conditionally

assigned to the genus Dahlica Enderlein, 1912 (type species:

Tinea triquetrella Hübner, [1812]), to which it looks much more

closely related than to the representatives of other Dahlicini genera

like Siederia Meier, 1953, Brevantennia Sieder, 1953, Eosolenobia

Filipjev, 1924 and Praesolenobia Sieder, 1954. The new species

rather occupies an intermediate position between Postsolenobia

Meier, 1958 and Dahlica (cf. Meier, 1958); its generic position still

requires clarification.

Terminology and principles of diagnosis are accepted herewith

according to Sauter (1956), Galliker (1958) and Arnscheid (1985)

unless stated otherwise; the taxonomic structure of the genus

according to Sauter & Hättenschwiler (1991, 1996).

Fig. 1. Dahlica karatyshica sp. nov. holotype à.

Dahlica karatyshica sp. nov.

Holotype d, “Ukraina, distr.[ictus] Donetsk, res.[ervatum] Kamennye Mogily, 26. IV.

1998, leg. E. Rutjan, ex pupa 28.1V.1998”: designated with label on printed form

“Holotypus” (in italics) with handwritten insertion: “Dahlica karatyshica Rutjan, sp. n.,

6, Ukraina, res. Kamennye Mogily (recto) and “Coll. Schmalhausen Inst. Zool., Kiev”

(verso). Paratypes: 35¢, 29, “Ukraina, distr. Donetsk, res. Kamennye Mogily,

26.1V.1998, leg. E. Rutjan, ex pupa 26.1V-3.V.1998”; 3, “Ukraina, distr. Donetsk, res.

Kamennye Mogily, 9.V.1997, leg. E. Rutjan”, d, “Ukraina, distr. Lugansk, prope

Sverdlovsk, pag.[us] Provalje, 10.V.1998”. — Holotype and a part of the paratypes are

deposited in the Lepidoptera collection, Schmalhausen Institute of Zoology (Kiev), a part

of the paratype series is forwarded to Zoological Museum, Taras Shevchenko National

University (Kiev) and Zoological Institute, Russian Academy of Sciences (St.-

Petersburg).

Male (Fig. 1). Smallest among Dahlica species: wingspan

7-9 mm. Head (Fig. 2) covered with grey ruffled hairs. Labial palpi

2,71

Fig. 2. Dahlica karatyshica sp. nov. paratype d: head structures (antennal scales and

bristles removed).

approximately 21, shorter than eye diameter. Antenna with 24-26

segments, ca. 0.6 of forewing length. Each antennal segment bear-

ing a belt of light-coloured scales at its top and bristles equal or

somewhat longer than the length of the bearing segment. Space

between eyes exceeding 0.33 of their diameter. Ocelli lacking.

Thorax and tegulae lıght grey. Wings (Figs 3, 4) narrow: forewing

length 31,, times, and that of hindwing 4 times exceeding its width.

Hindwings |, narrower than forewings. Fringes wide, more than

half of wing width; on forewing, along external and anal margins

paler than the wing ground-colour, on hindwing and along costal

margin of forewings the fringes are grey. They consist of long thin

two- and three-dentate scales of various length and ciliae (Fig. 3,

fui; Fig. 4, cf). Forewing pattern consisting of grey-brown lace-

like reticulation, darker in basal area. Pale spots grey-yellow.

Cloaking scales of forewing (Fig. 3, a) classified into classes [V—V

according to Sauter (1956), whereas according to Solyanikov

(1990) they comply with nos. 44.5. Underlying two-dentate

cuneiform scales (Fig. 3, b) of various length. Area between sub-

costal vein and costal margin covered with short two-dentate

cuneiform scales only (Fig. 3, c). Area between veins A, , ; and

anal margin covered with scales with rounded tip (Fig. 3, d).

Hindwings covered exclusively with cuneiform two-dentate scales

(Fig. 4, a). Forewing with 10 veins going out to the margin, of

which A, is less well expressed (n = 4; Fig. 3). Radial accessory cell

developed only in part of the examined specimens. Veins R, and R;,

R, and R;, and M, and M, merged pairwise. External margin of dis-

cal cell slight. Hindwing with 9 veins (Fig. 4). Intercalar cells lack-

ing. Vein branching not observed. Veins M, and M, completely

merged. Legs dark. Foretibia (Fig. 5, a) without epiphysis. Midtibia

(Fig. 5, b) with one, and hindtibia (Fig. 5, c) with two pairs of

spurs. Abdomen dark, covered with sparse light scales. Genital seg-

ments densely covered with long light-coloured scales.

Male genitalia (Fig. 6). Length of valve exceeding 3 times its

width in middle part, and 5 times exceeding saccus with vinculum.

Base of sacculus situated at the middle of inferior margin of valve.

Cucullus length 2.5 times its width. Tegumen with uncus 3.2 times

longer than vinculum with sacculus. Genitalic index approx. 1.1

(n = 3). Androconial scales two times shorter than total length of

sacculus and vinculum.

Male pupa structure as shown in Fig. 7.

Female (Fig. 8). It was impossible to describe the female alive.

Of 6 available female specimens only 2 could be kept intact. Body

length with protruded ovipositor 3.7 mm. Eyes small, diameter 2.6

times less than the space between them and approximately equal to

antenna scape length (Fig. 9). Labial palpi well distinctive. We did

not manage to determine the exact number of antennal segments,

but their number certainly exceeds 8. Tarsi of all legs with 4 seg-

ments (Fig. 10). Foretibia without epiphysis (Fig. 10, a). Mid- and

hindtibia bearing one pair of spurs (Fig. 10, b, c).

Female genitalia (Fig. 11). Ovipositor long and sclerotized.

Apophyses thin and long. Free ends of posterior apophyses (when

Ovipositor protruded) reaching the ostium (Fig. 11, b). Postvaginal

plate heart-shaped (Fig. 11, b). Thorns of intersegmental membrane

single and pointed, both long and short (Fig. 11, c). Postvaginal

plate membraneous area with merged and singular thorns, short and

wide (Fig. 11, d). Hairs of 7th segment normal.

Female pupa. Length of antennae covers exceeding length of

forelegs covers (Fig. 12).

Distribution. Up to the present, D. karatyshica sp. nov. is known

only from the type locality (SE Ukraine, Donetsk and Lugansk

oblast’s [provinces]).

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Fig. 5. Dahlica karatyshica sp. nov. paratype d : a — foreleg; b — midleg; c — hindleg.

Fig. 6. Male genitalia Dahlica karatyshica sp. nov. paratype d : a — ventral view: b — lat-

eral view.

Bionomics. Flight period from late April to the middle of May,

with peak in late April-early May, in a single brood. It is known to

occur in a Stipa capillata+Festuca sulcata+Festuca varia steppe

association of the solum petrosum variant, where together with

Reisseronia staudingeri (Heylaerts, 1879), it inhabits granite out-

crops on river banks (see Rutjan, 1998). Cases were collected on

slopes with a north-east exposure, attached to the sides of large

stones, with the caudal end freely hanging down. The pupal stage

lasts no less than 10 days. Under laboratory conditions males

emerged during the evening hours (20:00-22:00), but started flying

only the next morning (ca. 7:00). Females emerged in the morning

(5:30-8:00). Life span of males (as observed in captıvity) was 2,

and that of females about 3 days and nights. The larvae most prob-

ably feed on lichens. According to field observations, D. karatyshi-

ca Sp. nov. is a stenotopic species, occurring in places with a xero-

phytic vegetation.

Fig. 7. Dahlica karatyshica sp. nov. male pupa exuvium.

Type locality. SE Ukraine, Donetsk oblast’ (province),

Kamennye Mogily Nature Reserve.

Etymology. The name is derived from Karatysh, a small river at

the type locality.

Similar species. By size and genitalic index the new species is

close to Postsolenobia thomanni (Rebel, 1936), Dahlica (?) pal-

lidella (Zagulajev, 1997) and Siederia transsilvanica Herrmann &

Weidlich, 1999, but differs from these by a number of characters.

Fig. 8. Dahlica karatyshica sp. nov. paratype © (reference bar | mm).

Fig. 9. Dahlica karatyshica sp. nov. paratype 2: head structures.

Fig. 10. Dahlica karatyshica sp. nov. paratype 2: a - foreleg; b — midleg; c — hindleg.

From P. thomanni it differs by (1) lack of discal and anal dark spots

on the forewing; (2) presence of only three radial veins on the fore-

wing; (3) development of radial accessory cell on forewing in some

individuals; (4) absence of more than four-dentate scales; (5) habi-

tat: steppe xerophyte places, in contrast to mesophyte mountain

woods (cf. Sauter, 1956).

From D. (?) pallidella it differs by (1) only three radial veins and

two medial veins developed on forewing; (2) lack of intercalar cell

on hindwings and only two medial veins being developed; (3)

antennae exceeding half of forewing length; (4) space between eyes

exceeding eye diameter; (5) labial palpi 2 times shorter than eye

diameter; (6) tegumen with uncus domed (cf. Zagulajev, 1997).

—

Fig. 11. Female genitalia Dahlica karatyshica sp. nov. paratype 2: a — lateral view; b —

ventral view; c — intersegmental membrane thorns; d — thorns of postvaginal plate mem-

braneous area.

DATE TP Rune,

EST PES PSE 2 PP TNT

SS.

Fig. 12. Dahlica karatyshica sp. nov., 2: pupal skin, anterior part, ventral view.

From S. transsilvanica it differs by the lack of the epiphysis and

more than four-dentate scales, number of antennal segments 24—26

(cf. Herrmann & Weidlich, 1999).

From species with a genitalic index close to 1 (Eosolenobia man-

nil (Zeller, 1852), Dahlica fumosella (Heinemann, 1870), D. larel-

la (Chretien, 1906), D. incospicuella (Stainton, 1843) (cf. Sauter,

1956; Arnscheid, 1985), D. sauteri (Hättenschwiler, 1977), D.

rianella Hättenschwiler, 1981, D. achajensis (Sieder, 1966) and D.

colchica (Kozlov, 1985) D. karatyshica sp. nov. is distinguished by

its smaller size and the following characters: (1) venation; (2) num-

ber of antennal segments; (3) differences in cloaking scale classes;

(4) number of tarsal segments in female; (5) shape of thorns on

intersegmental membrane; (6) male and female genitalia structures.

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Nota lepid. 23 (1): 40-49; 01.1V.2000 ISSN 0342-7536

Unusual demonstration of autosomal dominant

inheritance of the black coloration of one of

America’s swallowtails: F, broods of the hybrid

Papilio polyxenes asterius with Papilio machaon

gorganus (Papilionidae)

E. A. LOELIGER* & F. KARRER**

*Hofdijck 48, NL-2341 ND Oegstgeest, The Netherlands

**Rebbergstrasse 5, CH-4800 Zofingen, Switzerland

Summary. From two inter se crossings of F, hybrids between the black American Papilio

polyxenes asterius Cramer, 1782 (Papilionidae) and the yellow P. machaon Linnaeus,

1758 from Europe, 62 F, offspring were obtained, 79% being black and closely resem-

bling P. polyxenes asterius and 21% yellow and almost identical with the European phe-

notypes of P. machaon. These percentages are consistent with the 3:1 ratio expected

according to Mendel’s law for the inheritance of an autosomal dominant trait. Earlier evi-

dence of such inheritance was based on backcrossing only of black F, hybrids of the black

American and the yellow European P. machaon strain with European P. machaon. One of

the yellow F, offspring was backcrossed with a specimen of P. machaon sphyrus Hübner,

1823, generating 32 healthy and exclusively yellow P. machaon-like butterflies, by this

confirming Mendel’s law. Possible causes of breeding healthy F, generations are dis-

cussed. Keeping down pathogenic germs when rearing the broods appears to be the most

likely explanation of our success.

Zusammenfassung. Aus zwei Kreuzungen der F,-Hybriden zwischen schwarzen nord-

amerikanischen Papilio polyxenes asterius Cramer, 1782 und gelben P. machaon

Linnaeus, 1758 aus Europa untereinander resultierten 62 Individuen einer F,-Generation,

von denen 79% schwarz gefärbt waren und P. polyxenes asterius sehr ähnelten, wahrend

21% gelb gefärbt und nahezu mit europäischen Phänotypen von P. machaon identisch

waren. Diese Prozentsätze sind konsistent mit einem bei autosomaler Vererbung eines

dominanten Allels gemäf der Mendelschen Regeln erwarteten 3:1-Verhältnis der

Phänotypen. Frühere Hinweise auf einen solchen Erbgang basierten ausschließlich auf

Rückkreuzungen schwarzer F,-Hybride des schwarzen amerikanischen und des gelben

europäischen P. machaon-Stammes mit europäischen P. machaon. Einer der gelben

F,-Hybride wurde mit einem Falter von P. machaon sphyrus Hübner, 1823 rückgekreuzt;

diese Nachkommenschaft bestand aus 32 gesunden, ausschließlich gelb gefärbten

Individuen (Phänotyp wie P. machaon), womit die Vererbung der Flügelgrundfarbe nach

Mendelschen Regeln wiederum bestätigt wird. Mögliche Gründe für die erfolgreiche

Aufzucht lebensfähiger F,-Generationen werden diskutiert. Die Unterdrückung der

Ausbreitung von Pathogenen während der Zucht erscheint als die wahrscheinlichste

Erklärung für den beobachteten Erfolg.

Resume. A partir de croisements entre hybrides F,, issus d’accouplements entre la forme

noire américaine Papilio polyxenes asterius Cramer, 1782 (Papilionidae) et la forme jaune

européenne de P. machaon Linnaeus, 1758, une génération F, de 62 individus a été

obtenue dont 79% sont noirs et ressemblent à P. polyxenes asterius et 21% jaunes et

presque identiques au phénotype européen de P. machaon. Ces pourcentages concordent

avec le rapport 3:1 attendu des lois de Mendel pour la transmission autosomique d’un ca-

ractère dominant. Les mises en évidence précédentes reposaient sur des rétrocroisements

entre hybrides F, issus de la forme noire américaine et d’une souche européenne de P.

machaon avec P. machaon d'Europe. Un exemplaire jaune de la génération F; fut croisé

avec un exemplaire de P. machaon sphyrus Hübner, 1823, donnant 32 papillons

vigoureux, exclusivement jaunes, du type P. machaon, confirmant ainsi les lois de

Mendel. Les causes possibles de l’élevage convenable de générations F; sont discutées.

L'absence de germes pathogènes lors des élevages apparaît la principale explication de

notre succes.

Key words: Lepidoptera, Papilio, hybrids, crossing, coloration, inheritance, Europe,

America.

Introduction

The black coloration of some American swallowtail butterflies

(P. polyxenes asterius Cramer, 1782, P. brevicauda Saunders, 1869,

P. ‘nitra’ Edwards, 1883 — the latter is now considered as a geneti-

cally integrated morph within P. zelicaon Lucas, 1852, cf. Sperling,

1987) was repeatedly shown to be dominant over the yellow of

others (P. machaon Linnaeus, 1758 and P. zelicaon), due to one sin-

gle gene dominating over yellow (Clarke & Sheppard, 1955;

Fisher, 1977; Blanchard & Descimon, 1988). Evidence for this was

the 1:1 segregation of the F, generations of backcrosses between

any random individual of an F, hybrid of the black/yellow cross

with a genuine yellow individual. Confirmation was provided by

the 3:1 ratio of black:yellow after sib matings of black individuals

after one or more backcrosses with a yellow one. But sib matings

of the F, of swallowtail hybrids remained unsuccessful as in the

early fifties (Clarke et al., 1953-1955; Aubert et al., 1997); only

after at least one backcross were fertile sib matings obtained

(Clarke & Sheppard, 1955; Aubert et al., 1997). In contrast to these

results, the findings presented here demonstrate Mendelian autoso-

mal dominance of black coloration by means of F, X F, inter se

matings. Possible explanations for our success will be discussed.

Material and methods

Pupae. Most of the non-hybrid pupae were obtained commer-

cially, those of P. machaon gorganus Fruhstorfer, 1922 and P. ma-

chaon sphyrus Hübner, 1823 from caterpillars collected in the

Czech Republic and Sicily, respectively. Only a few specimens of

P. machaon gorganus came from caterpillars collected in

Switzerland. Pupae of P. polyxenes asterius came from Brandon

province, Manitoba, Canada.

Hand-pairing and oviposition. The technique of hand-pairing,

consisting of holding the male and female butterflies together, in

order that their genitalia interlock and insemination can take place

(Clarke, 1952; Clarke & Sheppard, 1956), was applied by only one

of us (EAL), with the following modifications: first, the butterflies

were held with their wings downwards instead of upwards, which

enhances the view of the genitalia; second, no rubbing after mutu-

al approximation of the genitalia was applied. Under optimum con-

ditions — healthy well-fed butterflies, a temperature of 20°C or

higher, sunshine in the room where experiments took place, early

afternoon (13:00-14:30) — the copulation usually took place with-

in one minute and lasted 30-120 min. Before and after copulation,

the butterflies were kept in round gauze-covered cages 65 cm wide

and 85 cm high. In the event of successful mating eggs would be

laid after an interval of 24 hours; for oviposition the females were

placed in small cages and at the start of incoming daylıght host

plants were positioned in direct sunshine. The number of eggs

deposited on the first day after mating easily reached 50, the total

number being about 250.

Rearing the larvae. In view of the excellent pupae material

obtained from outdoor caterpillar collection, eggs were not disin-

fected. Larvae, host plants and other material were treated only

after thorough hand washing. L,-L; were kept indoors, each larva

on a separate twig of fennel seedlings (Foeniculum vulgare),

preferably not more than 5 specimens per plant. Older caterpillars

were either placed separately on one large fennel twig kept in water

indoors or transported outdoors onto a 10 m° large bed of full-

grown fennel covered with fine gauze for protection against preda-

tors. Contact between caterpillars was minimal due to the size of

the bed and the plants. Signs of sickness were not observed and loss

of caterpillars was negligible.

Results

In 1996 and 1997, hand-paired mating of the P. machaon gor-

ganus male with the P. polyxenes.asterius female and vice versa

was obtained repeatedly, but not ali females displayed oviposition

of fertile eggs. Table 1 shows the results of two such crosses. The

F, hybrid imagines all resembled P. polyxenes asterius with a nor-

mal sex ratio. F, inter se matings resulted in abundant oviposition

by several females. Development and hatching of the L, larvae did

not proceed normally however: in 1996, embryogenesis of the more

than 100 eggs of one female stopped in about 70% of cases after

they had turned brown, and of the remaining eggs which turned

black only hatched 1:3, which means that only 10% came through.

In 1997, the rate of successful hatching of the F, generation was

substantially higher, with normal pupation of most caterpillars. The

F, imagines eclosed after a few weeks, slightly more than */, of

them again being practically indistinguishable from P. polyxenes

asterius, those remaining closely resembling P. machaon gorganus

(Table 2). The sex ratio was not different from 1:1. These figures

are consistent with Mendelian-type autosomal dominant inheritan-

ce of the black coloration of P. polyxenes asterius. In a final exper-

iment, one of the machaon-like males of the F, imagines was suc-

cessfully mated with a female of P. machaon sphyrus (Eller, 1936).

Table 1. Results of backcrosses of F, imagines of the hybrid P. machaon gor-

ganus 3 x P. polyxenes asterius 2 with a P. machaon gorganus 3 and 9,

respectively.

P. polyxenes P. machaon-like

asterius-like (black) (yellow)

Meee ae

m 6 x (m/a) 2 2 1 2 3

Percentage

m — P. machaon gorganus; a — P. polyxenes asterius

* The deviation from the hypothetical expectation of 13:13 is so small that the

probability of its occurring by chance is > 0.50 (chi-square, degrees of free-

dom = 1).

Table 2. Results of F, inter se matings of the hybrid P. machaon gorganus & with

P. polyxenes asterius ®

* The deviation from the hypothetical 46.5:15.5 ıs so small that the probability

of its occurring by chance is > 0.50 (chi-square, degrees of freedom = 1).

In spite of the low vitality of the female, sufficient progeny were

obtained to confirm Mendelian-type inheritance, all 32 imagines

being yellow with a sex ratio of 1:1.3.

Discussion

The first report on spontaneous mating in captivity of a P.

machaon male with a P. polyxenes asterius female appeared at the

beginning of this century (Floersheim, 1910). The author, a well-

known entomologist, observed the copulation in his butterfly house

on July 29, 1908. He pointed out: “The day was a gloriously fine

one, and the pairing, which took place about mid-day, lasted about

four hours”. The resulting hybrid imagines all closely resembled P.

polyxenes asterius. The backcross with a P. machaon unfortunate-

ly ended in drowning of the female. More than 40 years later, after

the introduction of hand-pairing (Clarke, 1952), the hybrid was

obtained many times (Clarke & Knudson, 1953; Clarke &

Sheppard, 1953), leading within a couple of years to the conclusion

that black coloration of some of the American P. machaon-group

species is determined by a single gene which is dominant over yel-

low (Clarke & Sheppard, 1955). This conclusion was, however, not

based on straightforward inter se crossing of the F, hybrids, since

the latter remained unsuccessful. It was deduced from the results of

backcrossings which resulted in a black:yellow ratio of 1:1. More

recently, this Mendelian-type dominance of black coloration was

confirmed by backcrossing over four generations black F, females

with wild P. machaon males, which yielded imagines with a sex

ratio opposite to that predicted by Haldane’s Rule (Blanchard &

Descimon, 1988; Sperling, 1993). This back-cross evidence of

autosomal dominance of black over the yellow coloration was also

shown by us (Table 1). Numbers were too small, however, to assess

male:female ratios. But, contrary to other authors, we were also

able to deduce the Mendelian-type autosomal dominant inheritance

of black from offspring of inter se crossings of F, x F, hybrids

(Table 2). To corroborate the results, one of the yellow F, males was

backcrossed with a P. machaon sphyrus female, yielding 32

healthy, exclusively yellow imagines.

We believe that our results of inter se crossing are indeed large-

ly due to particularly healthy broods from outstandingly viable

pupae as well as careful breeding under strictly aseptic conditions,

handling the caterpillars as explained under Methods. Infection

with viruses is a well-known complication of swallowtail rearing

and is probably enhanced by inbreeding. Such an infection killed

the whole F, brood in the laboratory of a Japanese researcher (Ae,

1964) and is obviously also well known to British scientists (Clarke

& Larsen, 1986). This increased susceptibility to infection quite

certainly explains the failure to obtain viable offspring of F, x Fı

hybrid crossings of P. machaon with P. hospiton instead of P. poly-

xenes asterius (Clarke & Larsen, 1986; Aubert et al., 1997). The

French researchers also suspect another infectious agent, Le.,

Bacillus thuringiensis, which is contained in insecticides thereby

threatening caterpillar rearing.

Another cause of unsuccessful endeavours to obtain F, x F,

adults might be the fact that the other investigators worked with

P. machaon from southern Europe, while our material came from

Switzerland and from northern Europe. The British authors, in their

early work in the fifties, used the insular P. machaon sphyrus from

Malta (Clarke & Sheppard, 1955), the French investigators pupae

from Cahors (Lot). Both report the presence of some caterpillars,

the French even an L,, all of which died, however. Later endeavours

to obtain imagines from F, x F, crosses in which P. hospiton was

used instead of P. polyxenes asterius (Clarke & Larsen, 1986;

Aubert ef al., 1997), were equally disappointing. The British

authors observed a large variation in the hatching ratio of their

10 broods with low numbers of eggs, averaging only slightly more

than 10, clearly pointing to low quality of the commercially

obtained material. The hatching ratio varied from 0 to 56%, about

15% on the mean. Only one deformed female imago was obtained

from two typical hybrid caterpillars. In comparison to the British,

the French researchers obtained about 10 times as many eggs from

12 females. They report the occurrence of brown crowns on the

eggs as a sign of embryonic development in about 10% of cases,

and about 5% turned black, indicating full development of the

caterpillar, none of which hatched, however. Since, in their later

work, British researchers used P. machaon from Poland — the

French did not specify — our supposition that P. machaon strains

from southern Europe might be responsible for the low hatching

percentages and brood failures becomes very unlikely. Equally

improbable an explanation offers our modification of the handpair-

ing method, the disappointing results of the French researchers

having been obtained with broods from spontaneously copulating

butterflies (Aubert et al., 1997).

The above deliberations suggest that the success of obtaining

healthy F, adults from inter se crossing of swallowtail butterfly

hybrids is largely due to healthy livestock and germ-free handling

of the imagines, ova and larvae. This hypothesis is supported by the

report of an additional 8 healthy F, imagines in 1991 by another

well-known, highly experienced and exceedingly precise amateur

entomologist (Harbich, personal communication 1996).

Lastly, the results of an unintentional cross between a commer-

cially obtained P. polyxenes asterius male (pupa provided by Mr

John Reichel, Vancouver, CA) with a P. machaon gorganus female

(Kutscher, 1992) are also interesting in this context. The fairly

abundant offspring consisted of black as well as yellow specimens

in a proportion of about 1:1, suggesting heterozygosity of the black

male from Canada, possibly a natural hybrid specimen from the

cross of one of the yellow American swallowtails in the Manitoba

region. According to Sperling (1987 and personal communication

1998) Kutscher’s specimens are, judged from colour photographs,

“very reminiscent of ‘kahli and ‘avinoffi which are natural

hybrids”. As Brandon (Manitoba) was the origin of our P. polyxe-

nes asterius pupae material and lies close to the P. machaon x

P. polyxenes asterius swarm in central Manitoba, we cannot

exclude the possibility that it contains some introgressed machaon

genes (Sperling & Hansen, 1994). Relevant discussion is far

beyond the scope of the present communication, however.

We conclude with the remark made by investigators of swallow-

tail butterflies of the Americas in the Do it yourself chapter of their

recent handbook which states that: “Each cross needs its own ten-

derness that must be discovered empirically” (Tyler er al., 1994).

New data (addendum)

In 1999, we repeated our experiments using Japanese P. ma-

chaon hippocrates Felder & Felder, 1864, kindly supplied by

O. Yoshiaki (Tokyo), instead of the European P. machaon gor-

ganus; the black American P. polyxenes asterius originated from

the same area as in the earlier experiments. Hybridization was

again easily achieved, resulting in 44 F, specimens, but there were

only 10 females, that emerged only one to four weeks later than the

males. This low number and late emergence of females interfered

with inter se mating, and only two females produced enough eggs

for meaningful statistical analysis. Of the 186 eggs deposited by the

first one, 37 turned brown macroscopically. Inspection by means of

a low-power binocular microscope identified another 20 with signs

of embryonic pigmentation. Only one egg turned dark brown,

which is usually observed shortly before the hatching of the larva.

The latter did not come out, however. A second female produced

143 eggs, none of which displayed any signs of embryogenesis.

These additional observations suggest a larger phylogenetic dis-

tance between the American and the Japanese than between the

American and the European swallowtail species. They indicate that

the insular Maltesian material of P. machaon sphyrus used by the

British researchers could at least partly have been responsible for

the failure to produce viable offspring of the F, inter se crossings.

Clarke, Mittwoch & Traut (1977), in a study that we had over-

looked so far, present data that agree remarkably with our own:

from 32 matings of F, imagines, that are hybrids between a

P. machaon male from Finland and a P. polyxenes asterius female

from the east of the United States, these authors obtained 79 F,

imagines of which 78% are black and 22% yellow, with a normal

sex ratio in both. Considering our results — 79% and 21% respec-

tively — the close similarity is striking, the deviation from the

expected 3:1 ratio in particular. Considering the observation by

these authors that many larvae had died from disease, one is tempt-

ed to suppose that larvae predestined to develop into black ima-

gines resist influences of nurture easier than those becoming yel-

low imagines.

Acknowledgements

We thank W. Kutscher (Pohlheim, D), H. Harbich (Salz, D), and

F. A. H. Sperling (Berkeley, USA) for fruitful discussions by letter;

Mrs. G. P. Bieger-Smith for correction of the English and Mrs.

L. A. Nijssen-Kosters for typing the manuscript. We are also grate-

ful for the many helpful comments made by three anonymous

referees. Finally, we wish to gratefully acknowledge O. Yoshiaki

(Tokyo) for providing us with Japanese pupae material, as well as

bibliographic information.

References

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Nota lepid. 23 (1): 50-63; 01.TV.2000 ISSN 0342-7536

Eggshell fine structure of some species of

Lithosiinae (Arctiidae) of Far East Russia

Igor G. PLsusuch & Irina V. DOLINSKAYA

Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine,

UA-01601 Kiev MSP, Ukraine

Summary. Eggs of 7 species belonging to 6 genera of the Lithosiinae (Arctiidae) from

the Primorskiy Krai (Far East Russia) were examined with the use of scanning electron

microscopy. Descriptions and a comparative morphological analysis are provided for all

of these species. The diagnostic characters for genera and species are chosen.

Zusammenfassung. Eier von 7 Arten aus 6 verschiedenen Gattungen der Lithosiinae

(Arctiidae) aus dem Primorskiy Krai in Fernen Osten Rußlands wurden rasterelektronen-

mikroskopisch untersucht. Fur jede Art werden die Eier beschrieben und eine verglei-

chende morphologische Analyse gegeben. Diagnostische Merkmale für die Gattungen

und Arten werden begrundet.

Résumé. Des œufs de 7 espèces, appartenant à 6 genres de Lithosiinae (Arctiidae), en

provenance du Primorskiy Krai (Extrême Orient russe), ont été étudiés par moyen d’un

microscope électronique à balayage. La description, ainsi que l’analyse comparative de la

morphologie des œufs etudiés est présentée pour chacune de ces espèces. Des caractères

diagnostiques des œufs pour les genres autant que les espèces sont établis.

Key words: Lithosiinae, Arctiidae, egg, exochorion sculpture, morphology, Far East

Russia.

Introduction

The knowledge of the egg chorion of Arctiidae is still insufficient

and fragmentary. The eggs of some members of the subfamily

Arctiinae have been studied in some detail (Döring, 1955; Sarlet,

1967; Hinton, 1981; Orlando & Evoneo, 1984), whereas those of

the subfamily Lithosiinae remain poorly studied (Döring, 1955;

McFarland, 1972; Lafontaine er al., 1982; Garcia-Barros, 1985,

1986). Thus, the aim of our study was, even if partly, to fill this gap.

Material and Methods

The work is based on the original material collected by the

authors in the Primorskiy Krai (Far East Russia). The eggs were

obtained from females captured in the field. The moths were placed

into pasteboard boxes of a size of 100X90X50 mm, overlaid with

lichens. Females also laid the eggs at random in the box, without

choosing the special substrate. The eggs were examined with the

use of scanning electron microscopy (SEM). Both dry egg chorions

that were collected after hatching, and fresh specimens fixed with

glutaraldehyde-osmium tetroxide or with alcohol were examined.

Fixed eggs were first dehydrated in increasing acetone concentra-

tions, and then critical point dried.

The Lithosiinae species were identified with the aid of Daniel

(19395 1951, 1952, 1954) and Inoue (1982). The systematic

arrangement follows Dubatolov er al. (1993).

Description

Stigmatophora laecrita (Swinhoe, 1894)

Material. 22, Russian Federation, Primorskiy Krai, Ussuriisky rayon, Gorno-

Tayozhnaya Stantsia (Mountain-Taiga Station), 22.VII.1995 (eggs laid the same day;

10 eggs examined).

Egg spherical (Figs 1, 2), height 0.5 mm, diameter 0.5-0.55 mm.

Chorion white, hyaline. Exochorion sculpture cellular in the micro-

pylar area and with clearly marked aeropyles on the remaining egg

surface. Micropylar area more clearly expressed, represented by

2-3 rows of polygonal cells (Figs 2, 3). Micropylar rosette (50.8-

58.9 um in diameter) with 10-11 petalled cells, 3 times longer than

wide, joined along ”/;—/, of their length. Central portion of the

rosette like a small star-shaped depression (7.6-9.3 um in diame-

ter), 6.3-6.7 times smaller than rosette diameter, with 5 micropylar

openings. The remaining egg surface smooth, with hardly notice-

able cells but clearly expressed aeropyles, bordered by roller-like

margins (9.8—12.0 um in diameter) (Fig. 4).

The larva nibbles out a large rounded opening at the apical area

of the egg (Figs 1, 2). Caged females laid the eggs in single-layered

clusters.

Miltochrista miniata rosaria Butler, 1877

Material. 42, Russian Federation, Primorskiy Krai, Ussuriisky rayon, Gorno-

Tayozhnaya Stantsia (Mountain-Taiga Station), 6.VII., 8.VII.1995 (eggs laid the same

days; 10 eggs examined).

Figs 1-6: 1 — S. laecrita (x 240), 2 — S. laecrita (x 240), 3 — S. laecrita (x 1500), 4 —S.

laecrita (x 1200), 5 — M. miniata rosaria (x 220), 6 — M. miniata rosaria (x 200).

Figs 7-12: 7 — M. miniata rosaria (x 780), 8 — M. miniata rosaria (x 780), 9 — P staudin-

geri (x 540), 10 — P staudingeri (x 1100), 11 — P. staudingeri (x 720), 12 — L. quadra

(x 190).

Egg high, oblong-ovoid, narrowing towards the base (Fig. 5).

Height 0.7-0.85 mm, the largest width 0.3-0.45 mm. Egg pale

orange. Chorion pale brown, hyaline. Exochorion sculpture clearly

expressed, cellular (Fig. 8). Micropylar area represented by 2-3

rows of polygonal cells with flat cell margins (Figs 6, 7).

Micropylar rosette (51.9-68.6 um in diameter) with 7-8 wide cells,

1.5-2 times longer than wide, joining along '/,—’/, of their length.

Central portion of the rosette like a small star-shaped depression

(4.6-7.1 um in diameter), 9.7-11.2 times less than rosette diameter

with 3-5 micropylar openings (Fig. 8). The cells across the remain-

ing egg surface are concave and show sharply uplifted cell margins

reaching 1.3-1.6 um (Fig. 8).

The larva nibbles out a rounded opening at the apical area of the

egg. In captivity, the eggs were laid in single-layered, loose clus-

ters.

Paraona staudingeri Alpheraky, 1897

Material. 32, Russian Federation, Primorskiy Krai, Ussuriisky rayon, Gorno-

Tayozhnaya Stantsia (Mountain-Taiga Station), 4, 9, 11.VII.1995 (eggs laid the same

days; 10 eggs examined).

Egg spherical, height 0.5 mm, diameter 0.65 mm. Egg pale

green, immediately before eclosion the egg is green. Chorion

white, hyaline. Exochorion sculpture cellular in the micropylar

area, remaining egg surface faintly folded and with moderately

expressed aeropyles (Fig. 9). Micropylar area more clearly marked,

represented with 2—3 rows of polygonal cells. Micropylar rosette

(72.6-91.5 um in diameter) with 11-13 cells, 2-3 times longer than

wide, with margins fusing along ’/;—/, of their length. Central por-

tion of the rosette looking like a small star-shaped depression

(5.0-6.3 tum in diameter), 14.5 times smaller than the rosette dia-

meter, with 5 micropyles (Fig. 10). The remaining egg surface is

slightly folded, and presents hardly noticeable cells with slender

cell margins (0.2-0.3 um) and more clearly identifiable aeropyles

(1.95-2.9 um in diameter) bordered by roller-like margins

(Fie? BP):

The larva nibbles out a rounded opening at the apical area of the

egg. Captive females laid the eggs in single-layered clusters.

Lithosia quadra (Linnaeus, 1758)

Material. 22, Russian Federation, Primorskiy Krai, Ussuriisky rayon, Gorno-

Tayozhnaya Stantsia (Mountain-Taiga Station), 7.VII.1994 (eggs laid the same day:

10 eggs examined). 5

Egg spherical, height 0.5 mm, diameter 0.5-0.6 mm. Egg pale-

green. Chorion white, hyaline. Exochorion sculpture clearly

expressed, pitted-cellular (Fig. 12). Micropylar area represented by

2 rows of pitted cells. The micropylar area sculpture differs from

that on the remaining egg surface by the lack of aeropyles (Fig. 13).

Micropylar rosette (67.0-85.0 um in diameter) with 8-9 wide cells,

1.5-2 times longer than wide, with margins fused along 7/;—7/, of

their length. Central portion of the rosette looking like a small star-

shaped depression (7.4-8.5 um in diameter), 9-10 times smaller

than the rosette diameter, with 5 micropyles (Fig. 14). Remaining

egg surface with wide cell margins (10.2-10.8 um) and minute

aeropyles (2.8-3.1 um in diameter) at the margin junctions

(Preset).

We consider this type of sculpture to be transitional between the

pitted and the cellular sculpture. Its cells are not polygonal (as in

the micropylar area or in the Notodontidae) (Dolinskaya, 1987a, b),

but are rather rounded or oval with unclear margins.

The larva nibbles out a rounded opening at the apical area of the

egg. Captive females laid the eggs in single-layered clusters.

Agylla gigantea (Oberthür, 1879), A. collitoides (Butler, 1885)

Material. A. gigantea 29, A. collitoides 22, Russian Federation, Primorskiy Krai,

Ussuriisky rayon, Gorno-Tayozhnaya Stantsia (Mountain-Taiga Station), 4, 7.VII.1995

(eggs laid the same days; 10 eggs of each species examined).

Egg spherical, height 0.45-0.55 mm, diameter 0.55-0.7 mm.

Egg of A. collitoides pale-green with faint bluish hue. Chorion

white, hyaline. Exochorion sculpture cellular in the micropylar area

and remaining egg surface folded with sharply expressed aeropyles

united in the cells (Figs 16, 17, 20). Micropylar area represented by

2-3 rows of oblong polygonal cells (Fig. 18). Micropylar rosette

(71.7-89.0 um in diameter) with 11-13 petalled cells, 2-3 times

longer than wide, joined along ?/.—/, of their length. Central portion

of rosette looking like a small star-shaped depression (8.2-

11.7 um in diameter), 7.6-8.7 times smaller than rosette diameter,

with 4-5 micropylar openings (Fig. 20). Remaining egg surface

folded, looking sharply expressed, united in the cell aeropyles, bor-

dered by large roller-like margins (4.0-5.7 um in diameter) and

with hardly noticeable cell margins (Figs 20, 21).

The larva nibbles out a rounded opening at the apical area of the

egg. Captive females laid the eggs in single-layered clusters.

Note. No significant differences have been found between both

species.

Eilema griseolum (Hubner, [1803])

Material. 32, Russian Federation, Primorskiy Krai, Ussuriisky rayon, Gorno-

Tayozhnaya Stantsia (Mountain-Taiga Station), 16, 18, 23.VII.1995 (eggs laid the same

days; 10 eggs examined).

Egg spherical, height 0.55-0.65 mm, diameter ca. 0.7 mm. Egg

yellowish white. Chorion white, hyaline. Exochorion sculpture

weakly expressed, consisting of hardly noticeable cells (Fig. 22).

(Clear border between micropylar area and remaining egg surface

lacking). There is a progressive transition in egg surface appear-

ance from the micropylar area towards the egg equator. Sculpture

more clearly defined in the first two rows (1-2 rows) of the

micropylar area, then gradually disappearing (Fig. 23). Micropylar

rosette (66.8—116.0 um in diameter) with 14-17 petalled cells, 2—3

times longer than wide, joining along °/, of their length. Central

portion of rosette looking like a small depression (6.7-8.7 um in

diameter), 10.0-13.3 times smaller than rosette diameter, with 4-8

micropyles (Fig. 24).

The larva nibbles out arounded opening at the apical area of the

egg. Eggs laid in single-layered clusters.

Discussion

Based on the foregoing data, we can reach the following conclu-

sions.

Eggs of the studied lichen moths are radial symmetric (upright

type) with the micropylar area located at the pole opposite to the

surface. Most of the studied Lithosiinae genera have spherical eggs

(Stigmatophora Stgr., Eilema Hb., Agylla Wlk.), while oblong-

ovoid-shaped eggs occur in Miltochrista Hb.. The size ratio of the

eggs varies: spherical — height 0.45-0.65 mm, diameter 0.5-

0.7 um; oblong-ovoid — height 0.7-0.85 um, largest width

-0.3-0.45 um.

The coloration of the eggs depends of their contents, and is most-

ly of a greenish tint (Paraona Moore, Lithosia quadra, A. colli-

toides). Some species have yellowish-white (E. griseolum) or pale

orange (Miltochrista) eggs. The chorion is hyaline, mostly white

(otherwise pale brown: Miltochrista).

The exochorionic sculpture consists of a reticulate pattern,

formed by the cell margins. It may be cellular over the whole egg

surface, with sharply pronounced cell margins (Miltochrista) or

cells developing only in the micropylar area, and the remaining egg

surface with clearly expressed aeropyles united in the cells

(Stigmatophora, Paraona Agylla). Sometimes cells are hardly

noticeable (Zilema). The sculpture may also be peculiar, pitted-cel-

lular (L. quadra). A similar type of sculpture is present in some

species of the family Lasiocampidae (Euthrix potatoria askolden-

sis Oberth., Paralebeda plagiera W\k. — authors’ original data,

unpublished yet).

The cell margins of some Lithosiinae eggs are very narrow

(0.2-0.3 um), weakly marked, or lacking. However, in some

species they are sharply pronounced, reaching 1.3-1.6 mm

(Miltochrista) or 10.2-10.8 um (L. quadra).

The aeropyles can be either sharply expressed, bordered with

large roller-like margins (up to 4.0-5.7 um in Agylla spp., or

9.8-12.0 um in Stigmatophora), small (2.8-3.1 um in L. guadra),

or even not identifiable under SEM (as in Eilema, Miltochrista).

The micropylar area is represented by the rosette and 2—3 rows

of polygonal cells with clearly expressed cell margins. The size of

the micropylar rosette varies from 50.8-68.6 um (Stigmatophora,

Miltochrista) to 71.7-91.5 um (Agylla, Paraona), and the number

of rosette cells amounts to between 7-9 (L. quadra, M. miniata

rosaria) and 14-17 (Eilema). The central portion of the rosette

looks like a small star-shaped depression, 6.3-14.5 times smaller

than the rosette diameter. The number of micropylar openings

varies from 3 to 5.

Studied species of Lithosiinae are similar to Arctiinae (Pljushch

& Dolinskaya, in press) in having the same oviposition behaviour

in captivity. The eggs are laid in single-layer tight clusters, where

FRA RARE ARTE |

188. @U

18 AU cK

Figs 13-18: 13 — L. quadra (x 780), 14 — L. quadra (x 1200), 15 — L. quadra (x 780),

16 — A. gigantea (x 160), 17 — A. gigantea (x 210), 18 — A. gigantea (x 540).

Figs 19-24: 19 — A. gigantea (x 950), 20 — A. collitoides (x 1000), 21 — A. collitoides

(x 780), 22 — E. griseolum (x 150), 23 — E. griseolum (x 280), 24 — E. griseolum

(x 1000).

they are pressed to one another (Stigmatophora, Paraona, Eilema,

Lithosia, Agylla) or not (Miltochrista).

The larva nibbles out a uniformly large, rounded opening at the

apical (micropylar) area of the eggs, more rarely in the apical-la-

teral part.

Comparative table of the character states in studied species of

Lithosiinae.

Species >

Shape (A — spherical, B — oblong-

ovoid).

Egg colour (A - different green tints, B

— pale orange, C — yellowish white)

Chorion colour (A - white, B — pale

brown)

Exochorion sculpture (A — cellular

only in the mere area, the

remaining surface looks like aeropyles,

B - cellular over the whole NE sur-

face, C — hardly noticeable cells, D -

pitted-cellular

Ess margins (A - hardly noticeable, B

— clearly noticeable)

Aeropyles (A — moderately or clearly

noticeable, B — not visible, C — sharply

noticeable)

Oviposition (A — the eggs pressed to

one another, B — not ae

1 — S. laecrita, 2 — M. miniata, 3 — P. staudingeri, 4 — L. quadra, 5 — A. gigantea & A. col-

litoides, 6 — E. griseolum. * Available for A. collitoides only.

The comparative analysis pursued herein allows the evaluation of

the following characters (Table).

1. The egg shape can be used to identify some genera

(Miltochrista) or for grouping of the genera (see below).

2. Such characteristics as the way the larva nibbles out the eclo-

sion opening and the oviposition behaviour in captivity are not as

reliable as in tiger-moths.

3. The egg coloration can be used to identify species or genera.

4. The chorion coloration can be used to identify some genera

(Miltochrista).

5. The exochorion sculpture allows to identify species and ge-

nera. It is useful for grouping of the genera (see below).

6. The characters of the micropylar area, such as the number of

micropylar area rows, the number of rosette cells, and the number

of micropylar openings, are the only quantitative characters repre-

senting a continuous clinal variation and are subject to intraspecif-

ic variation. Therefore only the extremes of a cline could appear

useful for diagnostic purposes.

As a result, all examined genera of Lithosiinae may be placed in

two groups according to the egg characters (with three subgroups

in one of these):

(1) Miltochrista — egg oblong-ovoid, chorion pale-brown.

(2) Stigmatophora, Paraona, Lithosia, Agylla and Eilema — egg

spherical, chorion white.

(a) Lithosia — exochorion sculpture pitted-cellular.

(b) Stigmatophora, Paraona, Agylla — exochorion sculpture

looking like clearly expressed aeropyles united in the

cells (Stigmatophora — exochorion sculpture smooth with

clearly expressed aeropyles; Paraona — exochorion sculp-

ture faintly folded with moderately expressed aeropyles;

Agylla — exochorion sculpture clearly folded with sharply

expressed aeropyles).

like hardly noticeable cells, aeropyles not visible.

All the examined genera can be placed also into four groups

according to the exochorion sculpture:

(1) Exochorion sculpture cellular only in the micropylar area, the

remaining surface looking like clearly expressed aeropyles

united in the cells — Stigmatophora, Paraona, Agylla.

(2) Exochorion sculpture cellular on the whole egg surface —

Miltochrista.

(3) Exochorion sculpture pitted-cellular — Lithosia.

(4) Exochorion sculpture weakly expressed, looking like hardly

noticeable cells — Eilema.

Acknowledgements

We wish to express our thanks to Dr M. Ponomarenko and

Dr E. Belayev (Institute of Biology and Pedology, Vladivostok,

Russia) and to Dr M. Omel’ko (Gorno-Tayozhnaya Stantsia

[Mountain-Taiga Station], Primorskiy Krai, Russia) for their assis-

tance during our field work in Primorskiy Krai and granting us their

stationary facilities. We are obliged to Dr A. Lvovsky (Zoological

Institute, Russian Academy of Sciences, St. Petersburg) and to Mr

I. Kostjuk (Zoological Museum of the National Shevchenko

University in Kiev, Ukraine), who kindly assisted with the litera-

ture. Mr I. Kostjuk provided access to the collection material under

his care. |

References

DANIEL, F., 1939. Gedanken zu einigen Arctiiden-Formen (Lepidoptera). —

Mitt.münch.ent.Ges. 29: 354-368.

DANIEL, F., 1951-1954. Beiträge zu Kenntnis der Arctiidae Ostasiens unter

besonderer Berücksichtigung der Ausbeuten von Dr. h. c. H. Höne aus diesem

Gebiet (Lep.-Het.). III Teil Lithosiinae. — Bonn.zool.Beitr. 1951 2(3-4):

291-327, Taf. I; 1952 3(1-2): 75-90, Taf. II; 1952 3(3-4): 305-324; 1954

5(1-2): 89-138, Taf. III.

DoLiNskayA, I. V., 1987a. Morphology of the egg of Notodontid moths

(Lepidoptera, Notodontidae) of the fauna of the USSR. Comm. 1. — Vestnik

Zoologii 1: 65-74. [In Russian].

DouinskayA, I. V., 1987b. Morphology of the egg of Notodontid moths

(Lepidoptera, Notodontidae) of the fauna of the USSR. Comm. 2. — Vestnik

Zoologii 2: 50—60. [In Russian].

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Verlag. 154 S.

DuBATOLOV, V. V., TSHISTIAKOV, Yu. A. & VIDALEPP, J., 1993. A list of the

Lithosiinae of the territory of the former USSR (Lepidoptera, Arctiidae). —

Atalanta, Würzburg 24(1/4): 165-175.

GARCIA-BARROS, E., 1985. Identificaciön de la larva y datos biolögicos de eilema

uniola Rambur, 1858. Caracterisatiôn provisional de la larva del genero

Eilema Hübner (Lepidoptera, Arctiidae). — Bol.Asoc.Esp.Ent. 9: 223-237.

GarciA-Barros, E., 1986. Descripciön de las fases preimaginales y biologia de

Paidia murina (Hb., 1822) (Lepidoptera, Arctiidae). — Graellsia 42:

175-187.

Hinton, H. E., 1981. Biology of Insect eggs. 3 vols. — Oxford, Pergamon Press.

PSP:

INOUE, H., 1982. Arctiidae. In: Moths of Japan 1: 638-659; 2: 74-79, 136-137,

334-342; pls. 154-162, 346-348.

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and life history of Acsala anomala (Arctiidae: Lithosiinae). — J.Lep.Soc. 36

(3): 218-226. |

MCFARLAND, N., 1972. Egg photographs depicting 40 species of Southern

Australian Moths. — J.Res.Lepid. 10(3): 215-247.

ORLANDO, S. O. & EvonEo, B. F., 1984. Caracterizacäo morfolögica do ovo €

lagarta de Eupseudosoma aberrans Schaus, 1905 e E. involuta (Sepp., 1852)

(Lepidoptera, Arctiidae). — Revta bras.Ent. 28(3): 285-288.

PLJUSHCH I. G. & Do inskaya I. V., 2000. External morphology of the eggs of

some tiger-moths (Lepidoptera, Arctiidae). — Lambillionea (In press).

SARLET, L. G., 1967. Iconographie des œufs de Lépidoptéres belges. Deuxième

partie. Heterocera- Bombycidae — Sphingidae. Premier supplément. —

Bull.Ann.Soc.ent.Belg. 103: 294-301.

Nota lepid. 23 (1): 64-70; 01.1V.2000 ISSN 0342-7536

New data on the ecology of Thecla betulae in

the northeast of the Iberian Peninsula

(Lycaenidae)

Constanti STEFANESCU

Can Liro, E-08458 Sant Pere de Vilamajor, Spain

E-mail: canliro@teleline.es

Summary. Observations of 3rd and 4th instar larvae of Thecla betulae associated with

two ant species, Formica rufibarbis and Lasius grandis, are reported for the first time

under field conditions. New data on secondary host plants other than blackthorn (Prunus

spinosa) are also given for two populations from the northeast of the Iberian Peninsula.

Zusammenfassung. Vergesellschaftungen der Raupen (3. und 4. Larvalstadium) von

Thecla betulae mit zwei Ameisenarten, Formica rufibarbis und Lasius grandis, werden

erstmalig aus dem Freiland beschrieben. Ferner werden fiir zwei Populationen aus dem

Nordosten der Iberischen Halbinsel neue Beobachtungen zu sekundären Wirtspflanzen

neben der Hauptnahrungspflanze der Raupen (Schlehe, Prunus spinosa) mitgeteilt.

Résumé. Des observations relatives aux 3iéme et 4iéme état larvaire de Thecla betulae en

association avec deux espéces de fourmis, Formica rufibarbis et Lasius grandis, sont rap-

portées pour la première fois en conditions naturelles. De nouvelles données concernant

des plantes-hôtes secondaires autres que le prunellier (Prunus spinosa) sont également

présentées pour deux populations en provenance du nord-est de la péninsule ibérique.

Key words: Lycaenidae, Thecla betulae, myrmecophily, Formica rufibarbis, Lasius

grandis, host plants, Prunus domestica, Catalonia, Iberian Peninsula.

Thecla betulae (Linnaeus, 1758) is an Eurasian lycaenid occur-

ring in most of Europe and through Asia to Korea (Tolman &

Lewington, 1997). In Spain it is restricted to the northern part of the

country, from Galicia to Catalonia. The presence of isolated popu-

lations in Avila and north Extremadura has been questioned recent-

ly by Garcia-Villanueva et al. (1997) and, therefore, the southern-

most Spanish strongholds are probably found in Catalonia, a region

where its distribution is fairly well known (Viader, 1994;

Stefanescu, 1997).

Most ecological knowledge on European populations of this

species comes from extensive studies carried out by J. A. Thomas

in the early 70’s (Thomas, 1974). Though additional data have been

gathered by other authors since then (e.g. Ebert & Rennwald,

1991), the published information from southern European coun-

tries is almost inexistent.

In this note some new data on the ecology of T. betulae in the

northeast of the Iberian Peninsula are given, ın particular concern-

ing its myrmecophilous behaviour and its feeding habits.

Myrmecophily in T. betulae

Many lycaenids associate with ants during their immature stages,

though the degree of myrmecophily varies among the species

(Fiedler, 1991a). In the Western Palaearctic region, however, the

vast majority (more than 75%) of lycaenıds are myrmecophilous, at

least towards the end of the larval stage (Fiedler, 1991b).

The strength of ant-associations is related to the presence and

secretory activity of so-called myrmecophilous organs: multiple

minute pore cupola organs (PCOs), a dorsal nectar organ (DNO)

and paired tentacle organs (TOs) (cf. Fiedler er al., 1996). Both

DNO and TOs are absent in so-called myrmecoxenous species (1.e.,

those not attended by ants in nature) and, conversely, re-inforce and

stabilize the associations between larvae and ants in truly myrme-

cophilous lycaenids (Fiedler, 1991a; Fiedler er al., 1996). PCOs

occur in both lycaenid larvae and pupae of all studied European

species (e.g. Malicky, 1969). PCOs produce secretions highly

attractive to ants (possibly amino acids and pheromones; Fiedler et

al., 1996), though differences in the function of these organs have

been found (Fiedler, 1991a).

Within the tribe Theclini, species in the predominantly Holarctic

subtribe Thecliti (cf. Eliot, 1992) show reduced myrmecophilous

organs. TOs are entirely missing and the presence of a fully func-

tional DNO has not yet been confirmed without doubt for any

Thecliti species (Fiedler, 1991a). In Thecla betulae only PCOs are

present (Malicky, 1969; Fiedler, 1991b), and hence just a weakly

myrmecophilous relationship is to be expected.

In fact, no records of ant-associations of T. betulae larvae under

field conditions appear to have ever been published. In his thorough

review, Fiedler (1991a) scored T. betulae as a weakly myrme-

cophilous species because, as indicated by Malicky (1969), some

old records vaguely stated that occasionally ant-associations seem

to occur. However, intensive field work in England gave no addi-

tional evidence of this association (Thomas, 1974, 1975). On the

other hand, fully grown larvae are known to be very attractive to a

variety of ants in captivity (Taylor, 1915; Malicky, 1969; Fiedler,

pers. comm.), and pupal ant-associations in the wild with Lasius

niger or related taxa have already been described (Thomas, 1986;

Thomas & Emmet, 1989).

In May 1998 and 1999, I was able to make some interesting

observations of ant-associations of 3rd and 4th instar larvae of T.

betulae under field conditions for the first time. On 29 April 1998

three larvae in their 3rd instar were found resting on the underside

of leaves of two blackthorn bushes (Prunus spinosa) at Can Liro

(Sant Pere de Vilamajor, Barcelona province, 41°41’16 N 2°23’07

E, 310 m), in NE Spain. Both blackthorns were ca. 1.5 m tall and

grew in a hedgerow between cultivated fields. Egg-laying by T. be-

tulae had been recorded on both bushes the previous September

and October and seems to occur regularly every season (pers. obs.).

T. betulae larvae were monitored until they left for pupation by

the end of May. On 16 May a last instar larva was seen attended by

a worker of Formica rufibarbis. On 27 May another full-grown

larva was found attended by two Lasius grandis workers (Fig. 1).

In the first case, the association lasted only for a short time (approx.

1-2 min.), but on the second occasion L. grandis workers attended

the larva for at least 15 min. and showed no signs of disturbance

while the pictures were taken.

On 11 and 21 May 1999 a 3rd and a 4th instar larvae were found

on another blackthorn in the same hedgerow. Both larvae were

monitored almost daily until they left for pupation on 28 May and

2 June, respectively. The first larva was found attended by one or

two L. grandis ants on four out of eight occasions while in the 3rd

instar, and on eight out of ten occasions while in the 4th instar. The

second larva was seen attended by L. grandis two out of ten times,

but in one of these there were four mutualistic ants.

Thomas & Emmet (1989) suggested that the lack of field obser-

vations of ant-associations with 7. betulae larvae may be a conse-

quence of their habitat. In the temperate-zone woodlands the num-

ber of ants foraging on bushes and lower trees is limited and thus

‘(SOIICIIN ‘IN :yde18ojoyd) SI9Y10M

PUDAS snısp7 OM] AQ PapUd) 2D/H12Q DJ20YI JO BAIL] aeysur YANO “| ‘SA

the probability of encounters between ants and T. betulae caterpil-

lars is much reduced (see also Malicky, 1969 and Fiedler, 1991a for

a similar but more general reasoning). In the area where the fore-

mentioned observations were made, however, blackthorns and a

nearby peach tree Prunus persica were intensively visited by

Lasius grandis and Formica rufibarbis throughout April-May and

June (but not later on in the season), attracted by aphids that con-

centrated by that time on the underside of some young leaves and

possibly also by extrafloral nectaries occurring on these plants (cf.

Tilman, 1978). My own data seem to indicate that these associa-

tions may be quite common, though full-grown larvae have been

found several times at the same hedgerow and also at nearby places

without mutualistic ants.

Host plant use

Most European records mention blackthorn Prunus spinosa as

the most usual food plant of T. betulae (e.g. Tolman & Lewington,

1997) and in Spain this was the only species recorded so far

(Munguira et al., 1997).

In 1998, however, one larva and several eggs were found on plum

Prunus domestica at two different sites. On 23 April a 3rd instar

larva was found on the underside of a leaf in Can Riera de Vilardell,

an agricultural area surrounded by evergreen oak forest near the

village of Sant Celoni, Barcelona province (41°41 N 2°32’ E, 250

m). In addition, during the winter, several eggs were found on var-

ious plums growing in the vicinity of Can Liro.

Finally, on 7th October 1995, also at Can Liro, a female was seen

in Oviposition behaviour on a peach tree Prunus persica. She

alighted on the tree and immediately started walking on a twig, tap-

ping with her forelegs and antennae. However, no egg was finally

laid. Although no actual oviposition record was obtained, the dis-

tinctive behaviour shown by this female seems indicative that

peach trees may be sometimes used as a secondary host plant in this

population.

These observations confirm that several woody plant species

other than blackthorn are used as host plants by T. betulae in

Europe. In Germany, for example, Prunus domestica and other

related shrubs and trees of the family Rosaceae (Prunus insititia,

P. avium, P. padus, Crataegus monogyna and Chaenomeles japon-

ica) had already been cited as food plants (Ebert & Rennwald,

1991; K. Fiedler, unpubl. data). In Britain, P. insititia had also been

recorded as a food plant in the wild (Thomas & Emmet, 1989), and

a couple of other Rosaceae species (mostly in Prunus and related

genera) are documented as host plants of Eastern Asıan populations

(e.g. Dantchenko et al., 1995).

Moreover, there is at least one confirmed record of oviposition

on Betula pendula (Betulaceae) from Germany (Ebert &

Rennwald, 1991), that indicates that even plants not belonging to

the Rosaceae can be occasionally selected by ovipositing females.

This record, together with others obtained from the Russian and

Japanese literature (eggs and/or larvae found on several plants

belonging to the Betulaceae, Caprifoliaceae, Corylaceae,

Grossulariaceae and Salicaceae: Lukhtanov & Lukhtanov, 1994;

Korshunov & Gorbunov, 1995) led Fiedler (1991a) to categorize

T. betulae as a moderately polyphagous species.

Acknowledgements

Prof. Dr. K. Fiedler kindly read a first MS and provided much

information and valuable comments to this paper. My wife, Marta

Miralles, took the excellent picture of the Thecla betulae larva

attended by Lasius grandis workers and also found a 3rd instar

larva feeding on Prunus domestica. Xavier Espadaler (Universitat

Autonoma de Barcelona) identified Formica rufibarbis and Lasius

grandis ants.

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Tuomas, J. A., 1975. The ecology of the brown hairstreak butterfly. —

Rep.Inst.terr.Ecol. 1974: 24-25.

THoMAS, J. A., 1986. RSNC Guide to butterflies of the British Isles. — Country

Life, London. 160 p.

THOMAS, J. A. & Emmet, A. M., 1989. Thecla betulae. In: Emmet, A. M. & Heath,

J. (eds.): The butterflies of Great Britain and Ireland. Vol. 7, pt 1. — Harley

Books, Colchester: 123-126.

TILMAN, D., 1978. Cherries, ants and tent caterpillars: Timing of nectar produc-

tion in relation to susceptibility of caterpillars to ant predation. — Ecology 64:

1411-1422.

TOLMAN, T. & LEWINGTON, R., 1997. Collins Field Guide. Butterflies of Britain &

Europe. — Harper Collins Publishers, London. 320 p.

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Butll.Soc.catal.Lep. 74 51-60.

Nota lepid. 23 (1): 71-77; 01.1V.2000 ISSN 0342-7536

The karyotype and chromosome number of

Polyommatus buzulmavi (Lycaenidae)

Jurate PUPLESIENE & Alain OLIVIER

* Institute of Ecology, Akademijos 2, LT-2600 Vilnius. e-mail: jurate.puple-

siene@ village.uunet.be |

** Luitenant Lippenslaan 43 B 14, B-2410 Antwerpen. e-mail: alain.olivier@

village.uunet.be

Summary. The karyotype of Polyommatus buzulmavi Carbonell, 1992 from the Dez

Valley (Turkey, Hakkari province) is described and figured: the haploid chromosome

number n = 45 has been determined and several B-chromosomes have been found. This

number departs significantly from the number n = 23, which has been found in several

other species of Polyommatus s. str. so far.

Zusammenfassung. Der Karyotyp von Polyommatus buzulmavi Carbonell, 1992 aus dem

Dez Tal (Türkei, Hakkari Provinz) wird beschrieben und abgebildet. Als haploide

Chromosomenzahl wurde n = 45 festgestellt und es wurden mehrere zusätzliche B-

Chromosomen gefunden. Diese Zahl unterscheidet sich erheblich von der Zahl n = 23 bei

anderen Arten der Gattung Polyommatus Ss. str.

Résumé. Le caryotype de Polyommatus buzulmavi Carbonell, 1992 de la vallée du Dez

(Turquie, province de Hakkari), est décrit et figuré: le nombre haploïde de chromosomes

n = 45 a été déterminé et plusieurs chromosomes surnuméraires ont été trouvés. Ce nom-

bre diffère de façon significative du nombre n = 23, qui a été trouvé chez plusieurs autres

espèces appartenant au genre Polyommatus s. str.

Key words: Lycaenidae, Polyommatus buzulmavi, karyotype, Turkey.

Introduction

Carbonell ([1992]: 223-224) described Polyommatus buzulmavi

after a series collected by himself and a few colleagues in the Dez

and Zap Valleys, Hakkari province, Turkey. This butterfly can eas-

ily be told apart from the syntopic Polyommatus icarus

(Rottemburg, 1775), being highly characteristic by ıts much larger

size, its lighter blue colour and the light grey colour on the under-

sıde. In the same paper, on pp. 224-226, Carbonell described a dis-

tinct subspecies, Polyommatus buzulmavi narlica, after six males

collected about 10 km N. of Catak, Van province, Turkey, and list-

ed a series of characters, purported to differentiate the Catak mate-

rial from that from the Dez Valley. De Prins & van der Poorten

(1993: 12-13) described the female of this species and expressed

doubts about the validity of the distinction of narlica. They further

placed P. buzulmavi in the subgenus Meleageria de Sagarra, 1925.

Hesselbarth er al. (1995: 684-685) were able to examine larger

series of P. buzulmavi from both Hakkarı and Van provinces and

found the characters, purported to differentiate P. buzulmavi narli-

ca, to fall within the range of variation of nominotypical buzulmavi:

consequently they established the former name as a junior subjec-

tive synonym of the latter one. They further placed this taxon again

in subgenus Polyommatus Latreille, 1804. Both external and geni-

talic characters support its placement within last-named subgenus

(J.G. Coutsis, pers. comm.).

The biology of this taxon is still unknown, and so was, up to the

present, its karyotype and chromosome number.

Material and methods

On 23.VII.1999, the second author of the present publication

collected one single topotypical male of P. buzulmavi for karyolo-

gical study. The testes were fixed a few hours after collecting, when

the butterfly was still alive and, later on, a slide mount (AO 99173)

was prepared by the first author, who also made the photographs

reproduced here as Figs 1—3 (methodology followed as described

in detail in Olivier et al., 1999). The photo negatives were scanned

and processed with Corel Photo Paint. The specimen and the slide

mount are deposited in the Vlaamse Lepidoptera Collectie

Antwerpen (VLCA).

Karyotype and chromosome number of Polyommatus buzul-

mavi

The testes of the only specimen gave good results. Nine cysts

were found that contained spermatocyte divisions. 32 cells were at

the M-I stage, 2 cells at the M-II stage and 11 cells were in the early

prometaphase. The chromosomes of 8 M-I cells were counted and

4 M-I plates were photographed. The karyotype of P. buzulmavi is

asymmetric (Fig. 1). Bivalents fall into two main groups, that are

12.

Fig. 1. Karyotype of Polyommatus buzulmavi, slide AO 99173, M-I, n = 45, Turkey,

Hakkari province, Dez Valley, 1500-1700 m, 23.VII.1999, leg. A. Olivier (VLCA).

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Fig. 2. Karyotype of Polyommatus buzulmavi, slide AO 99173, M-I, n = 45 + B°+2B,

the arrow indicates the B bivalent, Turkey, Hakkari province, Dez Valley, 1500-1700 m,

23.VIL.1999, leg. A. Olivier (VLCA).

Fig. 3. Karyotype of Polyommatus buzulmavi, slide AO 99173, M-I, n = 45 + B” + 2B’,

the lower arrow indicates the B bivalent, the two upper arrows point at two additional

chromatine elements, Turkey, Hakkari province, Dez Valley, 1500-1700 m, 23.VII.1999,

leg. À. Olivier (VLCA).

almost equal in number: one group consists of comparatively large

bivalents which show a gradually decreasing series in size, the

other group of medium-sized bivalents that are almost equal in size.

The largest bivalents are situated more or less close to the center of

the metaphase plate, the medium ones usually surrounding these.

Besides the large and medium-sized bivalents, one small bivalent is

present, which is about half of the size of the smallest medium

sızed bivalent and which occupies about 1/5 of the largest bivalent

area. The two largest bivalents, as well as the smallest one, are dis-

tinguishable and hence can serve as the markers for the karyotype.

The largest bivalent is rounded, heteropycnotic and usually situat-

ed in the center of the metaphase plate. The C-heterochromatin

parts are in the centre of the bivalent, the euchromatin parts at the

sides. The second large bivalent is dumb-bell shaped and, contrary

to the first one, shows more C-heterochromatin at the edges and

much less in the central part of the bivalent. The smallest bivalent

in two metaphase plates showed a high degree of contraction and

was sharply contrasting in adhesion of dye. It was situated apart

from the rest of the bivalents, almost at the edge of the metaphase

plate. Both plates observed were in the early metaphase stage. In

the rest of the metaphases the smallest bivalent did not show such

a dense contraction, although it was usually attached to one large

bivalent by telomeric parts. The association of those two bivalents

resulted ın a V or ırregular U shape. It seems lıkely that the small-

est bivalent is the sex bivalent, but in order to establish that with

certainty, the karyotype of the female P. buzulmavi needs to be

studied. The remaining bivalents are rounded or elongated and usu-

ally isopycnotic.

Special attention was paid to B-chromosomes, that were found in

6 metaphase plates. Usually, there were two small additional

euchromatic univalents attached to A-bivalents. Due to that, A-

bivalents had a parachute-like shape. However, two metaphase

plates showed another pattern of B-chromosomes. One small,

rounded B-bivalent was found, which consists almost entirely of C-

heterochromatin and was situated at the edge of the metaphase

plate (Fig. 2). Those two plates also had two B-univalents, which

were Situated rather close to each other. The chromosome number

of those two metaphase plates is n = 45 + B” + 2 B’. One

metaphase plate showed an even more complicated pattern. At a

short distance from the metaphase plate, two small microelements

of C-heterochromatin of bivalent shape and approximately 5 tiny

univalent elements were situated: these could be additional B-chro-

mosomes as well (Fig. 3). However, their location rather apart from

the main metaphase plate keeps us from including these in the kary-

otype description of this species. Despite varying numbers of B-

chromosomes or chromatin elements, the number of A-bivalents

remained constantly at n = 45.

Discussion

Interestingly, the chromosome number of P. buzulmavi is very

different from that of P. icarus, P. juno Hemming, 1933, P. eros

eros (Ochsenheimer, 1808), P. eros yildizae Kocak, 1977 and

P. menelaos Brown, 1976, that all have a haploid chromosome

number of n = 23 (Robinson, 1971, 1990 and references therein;

Hesselbarth er al., 1995 and references therein; Larsen, 1975;

Brown, 1976). It is worth emphasizing that n = 23 (-24) appears

to be the modal number for the Lycaenidae (including

Polyommatines, Lycaenines and “Theclines”), thus conforming

to the ancestral condition (Lorkovié, 1990). Hence P. buzulmavi

represents a derived character state wıth regard to chromosome

number. Moreover, the approximate doubling of the chromosome

number in one of two evidently closely related species is sugges-

tive of chromosome fissions. Further karyological studies on

Central Asian Polyommatus s. str. species-group taxa appear high-

ly desirable.

Acknowledgements

The first author wishes to thank Dr. Luc De Bruyn

(Rijksuniversitair Centrum Antwerpen) for the logistic support and

constant encouragement, and Dr. Seppo Nokkala (Laboratory of

Genetics, University of Turku) for very valuable consultations in

staining techniques. The second author is indebted to Mr. Martin

Wiemers (Zoologisches Forschungsinstitut und Museum

Alexander Koenig, Bonn) for his company during the fieldwork

and for the German summary. We further like to thank Prof. Dr.

Konrad Fiedler (Lehrstuhl Tierökologie, Universität Bayreuth) and

Dr. Yuri P. Nekrutenko (Schmalhausen Institute of Zoology, Kiev)

for their most useful comments on the manuscript before it went to

press.

References

Brown, J., 1976. Notes regarding previously undescribed European taxa of the

genera Agrodiaetus Hübner, 1822 and Polyommatus Kluk, 1801 (Lep.,

Lycaenidae). — Entomologist’s Gaz. 27(2): 77-84, 10 text figs, 2 tabs.

CARBONELL, F., [1992]. Contribution à la connaissance du genre Polyommatus en

Turquie orientale (Lepidoptera: Lycaenidae). — Linn.belg. 13(4) (1991):

221-235, 2.pls. coul., 3 figs, 1 carte.

DE Prins, W. & VAN DER POORTEN, D., 1993. Grypocera and Rhopalocera of

Turkey 11. Description of the females of two little known lycaenid butterflies

from southeast Turkey: Polyommatus (Lysandra) dezinus (De Freina & Witt,

1983) and Polyommatus buzulmavi Carbonell, 1992 (Lepidoptera:

Lycaenidae). — Phegea 21(1): 11-16, 2 col.pls.

HESSELBARTH, G., VAN OORSCHOT, H. & WAGENER, S., 1995. Die Tagfalter der

Türkei unter Berücksichtigung der angrenzenden Lander. — Selbstverlag

Sigbert Wagener, Bocholt, 1354 S., 21 Tab. 75 Abb., 2 Farbkarten, 36 Farbtaf.

(mit 306 Abb.) (Bd. 1 & 2) + 847 S., 128 Farbtaf., 13 Taf., IV + 342

Verbreitungskarten (Bd. 3).

LARSEN, T. B., 1975. Chromosome Numbers and Notes on Testicular

Morphology of some Lebanese Rhopalocera (Insecta: Lepidoptera). —

Ent.scand. 6: 253-260.

Lorkovié, Z., 1990. The butterfly chromosomes and their application in sys-

tematics and phylogeny. /n: Kudrna, O. (ed.), Butterflies of Europe. Vol. 2:

Introduction to Lepidopterology, pp. 332-396, 36 text figs, 2 diagr., 2 col. pls.

— Aula Verlag, Wiesbaden. 559 pp.

OLIVIER, A., PUPLESIENE, J., VAN DER POORTEN, D., DE PRINS, W. & WIEMERS, M.,

1999. Revision of some taxa of the Polyommatus (A grodiaetus) transcaspicus

group with description of a new species from Central Anatolia (Lepidoptera:

Lycaenidae). — Phegea 27(1): 1-24, 2 col.pls., 2 tabs., 7 figs.

RoBINSON, R., 1971. Lepidoptera Genetics. — Pergamon Press, Oxford, New

York, Toronto, Sydney, Braunschweig, 687 p., 16 text figs, 63 tabs.

RoBINSON, R., 1990. Genetics of European butterflies. Jn: Kudrna, O. (ed.),

Butterflies of Europe. Vol. 2: Introduction to Lepidopterology, pp. 234-306,

6 text figs, 3 tabs. — Aula Verlag, Wiesbaden. 559 pp.

ve)

Nota lepid. 23 (1): 78-80; OL.IV.2000 ISSN 0342-7536

Axel Scholz

(11.6.1957-30.6.1998)

Während einer Exkursion in der Steiermark verunglückte Axel

Scholz am 30.6.1998 am Gulsenberg bei Preg a. d. Mur tödlıch.

Mit ıhm verliert die Entomologie einen versierten Kenner der

Mikrolepidopteren, der gerade erst begann, seine vielfältigen

Kenntnisse zur Systematik und vor allen Dingen zur Biologie der

europäischen Kleinschmetterlinge der Fachwelt mitzuteilen.

Axel Scholz wurde am 11. Juni 1957 in Ulm geboren. Nach dem

Besuch der Volksschule und des Gymnasiums in Laupheim legte er

1977 das Abitur ab und begann 1979, nach 2-jährigem Dienst bei

der Bundeswehr in Mittenwald, mit den Biologie- und

Sportstudium an der TH Darmstadt. Im Jahre 1984 erfolgte ein

Wechsel zur Deutschen Sporthochschule in Köln (Studiengang

Diplomsportlehrer), verbunden mit mehreren Urlaubssemestern

zur Finanzierung des Studiums. Wegen der schlechten Aussichten

auf eine Anstellung unterbrach er 1991 das Studium und begann

eine Tätigkeit als Angestellter in einer Arzneimittelfirma im

Bereich Analytik, die er bis zu seinem Tode ausführte. Parallel dazu

begann er 1993 an der Universität Augsburg einen Studiengang

Hauptschule in den Fächern Biologie, Sport und Deutsch, für den

im Frühjahr 1999 der Abschluß geplant war. Leider war es ihm

nicht mehr vergönnt, in dem von ihm gewählten Beruf tatig zu sein.

Die Liebe zur Natur wurde beı Axel Scholz schon ım Kindesalter

im Elternhaus geweckt, mit seinem leider schon 1965 verstorbenen

Vater wurden erste Sammelversuche unternommen. Sehr früh

eignete er sich sehr gute botanische Kenntnisse an, so daß er sich

bevorzugt mit der Biologie der Lepidopteren befaßte, mit einem

Gebiet, auf dem noch so Vieles Neue zu entdecken war. Auch seine

letzte Exkursion war den Minierern gewidmet, die Suche nach den

Minen von Lunakia alyssella am typischen Fundort war erfolg-

reich, wie die Blattminen an Alyssum montanum presseckeri in

seinem Exkursionsgepäck belegen. Außerordentlich förderlich für

seine Interessen wurde der seit 1988 währende Briefwechsel mit

Dr. J. Klimesch (1902-1997). Durch ihn erhielt er zahlreiche

Anregungen zur Beschäftigung mit den minierenden Vertretern der

Mikrolepidopteren, vor allem mit den Nepticuliden. Ein weiteres

wichtiges Ereignis in seiner entomologischen Entwicklung war die

Bekanntschaft mit E. Jäckh (1902-1993). Durch ihn erhielt er

neben zahlreichen Hinweisen aus dem reichen Erfahrungsschatz

als unerläßliche Hilfsmittel eine Reihe wichtiger entomologischer

Literatur, die Grundlage seiner Bibliothek bildete. Herr Jäckh war

es auch, der ıhn mit den Alucitidae vertraut machte, mit einer

Familie, über die dieser schon viele Jahre gearbeitet hatte. Herrn

Scholz ıst es zu danken, daß dieser Kenntnisschatz nicht ver-

lorenging, in der 1994 erschienenen gemeinsamen Revision wur-

den die Ergebnisse der Fachwelt vorgestellt.

Sein großes handwerkliches Geschick sowie seine ihm eigene

Exaktheit befähigten ihn, das auf Exkursionen oder durch Zuchten

erlangte Faltermaterial in beinahe unnachahmlicher Art und Weise

zu präparieren. Seine Sammlung ist nicht nur eine Quelle wertvol-

ler wıssenschaftlicher Informationen sondern auch ein ästhetischer

Genuß. Diese Exaktheit und Präzision der Beobachtung war es

auch, die ıhn auf Dinge aufmerksam werden ließ, die Andere bis-

lang übersehen hatten. Ein typisches Beispiel hierfür ist seine

Entdeckung, daß sich unter dem Namen Epermenia illigerella eine

weitere Art verbirgt, - Epermenia falciformis. Durch die Arbeit an

own classification’. After a brief Introduction, in which some problems with cur-

rent classification systems are briefly reviewed and species totals for superfami-

lies of Lepidoptera are presented on a table, the evolution of the Lepidoptera is

briefly outlined, with a diagram depicting a hypothetical ancestral moth. A

schematic phylogeny of lepidopteran superfamilies and several schemes for con-

stituent families are then presented. Then follows a brief historical review of

Lepidoptera classifications, after which the author presents his own classification

of the Lepidoptera. Subsequently, the author presents an Index to Families and

Higher Categories, an Index to Common Names of Families and various keys to

Lepidoptera Families, interspersed with various drawings of exo- and endomor-

phological characters. The book ends with a References section and an Index.

The overall impression it leaves, however, is that of a courageous but rather ama-

teuristic enterprise. In any case, its impact will painfully be overshadowed by the

recently published “Lepidoptera, Moths and Butterflies. Volume 1: Evolution,

Systematics, and Biogeography’, edited by N. P. Kristensen in the series

“Handbuch der Zoologie’ (reviewed in Nota lepid. 22(4): 317-322).

Alain OLIVIER

HALL, Jason P. W.: A contribution to Riodinid systematics (Lepidoptera:

Riodinidae).

21.5 x 28 cm, 42 pp., Tropical Lepidoptera 9(Suppl. 1) Published by Association

for Tropical Lepidoptera, Inc. Tel. (352) 392-5894 — fax: (352) 392-0479; e-mail:

jbhatl@aol.com; homepage: http://www.troplep.org, 30.X1.1998. ISSN 1048-

8138. To be ordered from the publisher.

This special issue of Tropical Lepidoptera consists of four papers (co-)authored

by Jason P. W. Hall. The first one, ‘A Review of the Genus Sarota (Riodinidae)’

includes, along with a brief characterization of the genus, a brief overview of the

biology, biogeography and taxonomic history of its component species, a key to

the identification of adults, illustrations of all known species and subspecies

(including many type specimens) with accompanying taxonomic notes and the

descriptions of five new species and two new subspecies. Twenty species are

recognised. Neotypes are designated for two species, one tribe is synonymised

with another tribe and a cladogram based on a small character matrix is used to

illustrate generic relationships within the more inclusive, newly conceived tribe.

The second paper, ‘Three new species of Riodinini from the cloud forests of

Ecuador (Lepidoptera: Riodinidae)’, further includes the illustrations of some

related taxa, two new combinations and one new synonym. The titles of the third

— ‘Nine new species and one new subspecies of Euselasia from Ecuador

(Lepidoptera: Riodinidae)’ — and fourth paper, ‘A new genus of Riodinid, with

a new species from Panama (Lepidoptera: Riodinidae: Nymphidiini’ summarize

well their content.

Alain OLIVIER

Davis, Donald R.: Lepidopterorum Catalogus (New Series) Volume 5.

Yponomeutoidea. Fascicle 48. Ochsenheimeriidae.

21.5 x 28 cm, VIII + 12 pp. Edited by J. B. Heppner. Published by Scientific

Publishers, Inc., P. ©. Box 15718, Gainesville, FL 32604 USA. 1998. ISBN 0-

945417-55-1 (Fasc. 48). — ISBN 0-945417-50-0 (Series). To be ordered from

the publisher.

In a Preface, the family is briefly introduced. After amap showing the world dis-

tribution of the overwhelmingly Palaearctic family of the Ochsenheimeriidae,

and a generic synopsis, a catalog of the 17 currently recognised species, placed

in two genera, is presented and six species are depicted on one black-and-white

plate. Brief notes are added for three of these. Then comes a References list (102

entries), a host plant index, an index to species and an index to genera.

Alain OLIVIER

HEPPNER, John B.: Lepidopterorum Catalogus (New Series) Volume 8.

Pterophoroidea. Fascicle 61. Tineodidae.

21.5 x 28 cm, VIII + 8 pp. Edited by J. B. Heppner. Published by Scientific

Publishers, Inc., P. O. Box 15718, Gainesville, FL 32604 USA. 1998. ISBN 0-

945417-58-6 (Fasc. 48). — ISBN 0-945417-50-0 (Series). To be ordered from

the publisher.

In a Preface, the family is briefly introduced. After amap showing the world dis-

tribution of the exclusively Australian family of the Tineodidae, and a generic

synopsis, a catalog of the 10 currently recognised species, placed in seven gen-

era, is presented and four species are depicted on one black-and-white plate.

Then comes a References list (23 entries), a host plant index, an index to species

and an index to genera.

Alain OLIVIER

SCOBLE, Malcolm J.: Lepidopterorum Catalogus (New Series) Volume 96.

Geometroidea. Fascicle 93. Hedylidae.

21.5 x 28 cm, VIII + 8 pp. Edited by J. B. Heppner. Published by Scientific

Publishers, Inc., P. O. Box 15718, Gainesville, FL 32604 USA. 1998. ISBN 0-

94541 7-66-7 (Fasc. 93). — ISBN 0-945417-50-0 (Series). To be ordered from

the publisher.

In a Preface, the family is briefly introduced. After amap showing the world dis-

tribution of the exclusively Neotropical family of the Hedylidae, and a generic

synopsis (only one genus, Macrosoma Hübner, 1818, is recognized), a catalog of

the 35 currently recognised species is presented and eleven representative

species are depicted on one black-and-white plate. Then comes a References list

(47 entries), a host plant index, an index to species and an index to genera.

Alain OLIVIER

HALL, Jason P. W.: A Revision of the Genus Theope, its Systematics and

Biology (Lepidoptera: Riodinidae: Nymphidiini).

21.5 x 28 cm, VIII + 127 pp., 274 figs. (incl. 9 plates with figs 10-77), 7 tables.

Paperback. Published by Scientific Publishers, Inc., P. O. Box 15718,

Gainesville, FL 32604 USA. 1999. ISBN 0-945417-95-0. To be ordered from the

publisher.

An excellent revision of the speciose and morphologically diverse Neotropical

nymphidiine riodinid genus Theope Doubleday, 1847, is presented. The taxo-

nomic history of the genus, its morphological variation and characteristics, bio-

geography, early stage biology and adult ecology, particularly male perching

behaviour, are discussed. Detailed notes are given for each species on identifica-

tion and taxonomy, habitats and ecology, and distribution, and the localities list-

ed and mapped for all examined specimens in the nearly twenty collections vis-

ited in Europe and in the Americas, as well as those recorded in the literature. A

dichotomous key and colour illustrations of adults for all species are given, and

drawings of the male abdominal and genitalic morphology for all but one

species, and those of the female genitalia for the majority of species, are pre-

sented with a list of all dissections examined. Two genera are synonymised with

Theope and five species are newly brought into combination with this genus.

Further, two new species and two new subspecies are described. À number of

taxonomic changes are also made at the species level and below, resulting in 68

species being recognised in the genus; 15 lectotypes are designated.

Alain OLIVIER

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3 a néon. L. G. 1950. A descriptive catalogue of the Palaearctic Euphydryas (Lepidoptera: Rhopalocera). —

à VE: Trans.R.ent.Soc.Lond. 101: 435-489, figs. 1-44, 7 maps.

_ Hicains, L. G. & RILEY, N. D., 1980. A field guide to the butterflies of Britain and Europe. 4th ed. — Collins,

> London. 384 p., 63 pls.

STAUDINGER, O., 1901. Famil. Papilionidae - Hepialidae. Jn: STAUDINGER, O. & REBEL, H. Catalog der Lepidopteren

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A journal of the Societas Europaea Lepidopterologica

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Wolk 23 No. 2 Basel, 01.07.2000 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

Assistant Editors: Dr. Roger L. H. Dennis (Wilmslow, GB), Prof. Dr. Konrad

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Contents @ Inhalt e Sommaire

OLIVIER, A. Discovery of the types of Polyommatus (Agrodiaetus) actis

Anais axonomic consequences (Lycaenidae) ............................... 86

GARCIA-BARROS, E. Comparative data on the adult biology, ecology and

behaviour of species belonging to the genera Hipparchia, Chazara and

Kanetisa in central Spain (Nymphalidae: Satyrinae) ...................... 119

ZOLOTUHIN, V. V. & Kostyuk, I. Yu. Phantosoma witti gen. et sp. nov., a

new autumn lasiocampid moth from Turkmenistan (Lasiocampidae) ..... 141

PFEIFER, M. A., ANDRICK, U. R., FREY, W. & SETTELE, J. On the ethology

and ecology of a small and isolated population of the Dusky Large Blue

Butterfly Glaucopsyche (Maculinea) nausithous (Lycaenidae) ............. 147

CuPEDo, F. Die geographische Variabilität von Erebia flavofasciata, nebst

Beschreibung einer neuen Unterart (Nymphalidae: Satyrinae) ............. 173

Dapporto, L. & FABIANO, F. Notes on some interesting Geometridae col-

ere) Tim TUS CEUTA May) 2... 185

Nota lepid. 23 (2): 86-118; 01.V11.2000 ISSN 0342-7536

Discovery of the types of Polyommatus

(Agrodiaetus) actis and its taxonomic

consequences (Lycaenidae)

Alain OLIVIER

Luitenant Lippenslaan 43 B 14, B-2140 Antwerpen, Belgium

e-mail: alain.olivier@village.uunet.be

Summary. The nomenclatural and taxonomic history of the nominal taxon Lycaena actis

Herrich-Schäffer, 1851 is outlined, and the taxonomic interpretations and quotations of

the names actis Herrich-Schäffer, 1851, atys Gerhard, 1851 and athis Freyer, 1851 in the

literature are reviewed. It thus appears that the three names apply to a taxon, which is

treated as Polyommatus (Agrodiaetus) firdussii (Forster, 1956) pro parte by Hesselbarth et

al. (1995) and subsequent authors. The discovery of two syntypes (d, 2) of Lycaena actis

confirms this statement. The male specimen is designated as lectotype and the type locali-

ty corrected as Turkey, Tokat province, env. Tokat. The name actis is considered as the

valid name for this taxon, thus promoting stability of nomenclature (cf. ICZN: Preamble,

Arts. 23.2 and 23.9). Current taxonomic problems in part of the actis group are discussed.

Under the name actis, probably more than one species is involved, but the picture is not

quite clear. The intensity of the blue colour on the male upperside varies from vivid violet

blue to a lighter blue, while the variation in colour does not always coincide with the

pronounced differences in chromosome numbers and karyotype that are being observed

(de Lesse, 1960, 1962; Lukhtanov er al., 1998). A provisional arrangement is proposed

for the nominal taxa Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851), P. (A.)

firdussii (Forster, 1956), P. (A.) pseudactis (Forster, 1960), species incertae sedis, P. (A.)

sertavulensis (Koçak, 1979), species incertae sedis, P. (A.) ernesti Eckweiler, 1989,

species incertae sedis, P. (A.) artvinensis (Carbonell, 1997) and P. (A.) sigberti Olivier,

van der Poorten, Puplesiene & De Prins, 2000 (= P. (A.) actis sensu Hesselbarth et al.

(1995) and subsequent authors).

Zusammenfassung. Die nomenklatorische und taxonomische Geschichte des nominellen

Taxons Lycaena actis Herrich-Schäffer, 1851 wird beschrieben, in Verbindung mit einer

Literaturübersicht über taxonomische Interpretationen und Zitate der Namen actis

Herrich-Schaffer, 1851, atys Gerhard, 1851 und athis Freyer, 1851. Alle drei Namen

scheinen sich auf dasselbe Taxon zu beziehen, das von Hesselbarth et al. (1995) und nach-

folgenden Autoren pro parte als zu Polyommatus (Agrodiaetus) firdussii (Forster, 1956)

gehörig aufgefaßt wurde. Die Entdeckung zweier Syntypen (3, 2) von Lycaena actis

bestätigt diese Einschätzung. Das männliche Exemplar wird hier als Lectotypus designiert

und seine Typuslokalität korrigiert (Türkei, Provinz Tokat, Umgebung von Tokat). Der

Name actis wird im Sinne der Stabilität der Nomenklatur (ICZN: Präambel, Art. 23.2 und

23.9) als valider Name des betreffenden Taxons angesehen. Gegenwärtige taxonomische

Probleme in der Gruppe um actis werden diskutiert. Unter dem Namen actis im Sinne der

vorliegenden Arbeit verbergen sich möglicherweise mehrere Arten, aber das gegenwärtige

Bild ist noch unklar. Die Intensität der blauen Flügeloberseitenfärbung der Männchen

variiert von lebhaftem Violettblau zu hellerem Blau, Unterschiede in der Flügelfärbung

fallen nicht immer mit den beachtlichen Unterschieden in Chromosomenzahlen und

Karyotypen zusammen (de Lesse, 1960, 1962; Lukhtanov er al., 1998). Für die nomi-

nellen Taxa Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851), P. (A.) firdussii

(Forster, 1956), P. (A.) pseudactis (Forster, 1960), species incertae sedis, P. (A.) sertavu-

lensis (Kocak, 1979), species incertae sedis, P. (A.) ernesti Eckweiler, 1989, species incer-

tae sedis, P. (A.) artvinensis (Carbonell, 1997) und P. (A.) sigberti Olivier, van der

Poorten, Puplesiene & De Prins, 2000 (= P. (A.) actis sensu Hesselbarth er al. (1995) und

nachfolgender Autoren) wird eine vorläufige systematische Anordnung vorgeschlagen.

Résumé. L’histoire de la nomenclature et de la taxinomie du taxon nominal Lycaena actis

Herrich-Schäffer, 1851 est esquissée, ainsi que celle des diverses mentions et interpréta-

tions des noms actis Herrich-Schäffer, 1851, atys Gerhard, 1851 et athis Freyer, 1851

dans la littérature. Il apparaît ainsi que ces trois noms s’appliquent a un taxon, qui est

traité comme étant Polyommatus (Agrodiaetus) firdussii (Forster, 1956) pro parte par

Hesselbarth er al. (1995) et les auteurs plus récents. La découverte de deux syntypes (à,

2) de Lycaena actis confirme ce point de vue. Le spécimen mâle est désigné comme lec-

totype est la localité type est corrigée comme étant Turquie, province de Tokat, env. de

Tokat. Le nom actis est considéré comme étant le nom valide pour ce taxon, promouvant

ainsi la stabilité de la nomenclature (Code International de Nomenclature Zoologique:

Préambule, Arts. 23.2 et 23.9). Les problèmes taxinomiques actuels relatifs à une partie

du groupe d’actis sont discutés. Probablement, plus d’une espéce est comprise sous le

nom actis, mais la situation est peu claire. L’intensité de la couleur bleue à la face

supérieure du mâle varie d’un bleu-violet intense à un bleu plus clair, tandis que la varia-

tion du teint ne coincide pas toujours avec les différences importantes du nombre de

chromosomes et du karyotype qui peuvent étre observées (de Lesse, 1960, 1962;

Lukhtanov et al., 1998). Une classification provisoire est proposée pour les taxons nomi-

naux Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851), P. (A.) firdussii (Forster,

1956), P. (A.) pseudactis (Forster, 1960), species incertae sedis, P. (A.) sertavulensis

(Kogak, 1979), species incertae sedis, P. (A.) ernesti Eckweiler, 1989, species incertae

sedis, P. (A.) artvinensis (Carbonell, 1997) et P. (A.) sigberti Olivier, van der Poorten,

Puplesiene & De Prins, 2000 (= P. (A.) actis sensu Hesselbarth et al. (1995) et auteurs

suivants).

Key words: Lycaenidae, Polyommatus (Agrodiaetus), actis, atys, athis, firdussii,

pseudactis, artvinensis, sertavulensis, ernesti, taxonomy, nomenclature, lectotype, chro-

mosome number, karyotype, Turkey, Iran, Armenia.

Introduction

The nomenclatural and taxonomic history of the nominal taxon

Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851) is a

complicated one. In the sixth volume of his “Systematische

Bearbeitung der Schmetterlinge von Europa, zugleich als Text,

Revision und Supplement zu Jakob Hübner’s Sammlung europäi-

scher Schmetterlinge” (Nachtrag zum Ersten Bande, Heft 48, p.

24), Herrich-Schäffer ([1851]) gives the following short descrip-

tion: “Actis Kad. — Sppl. 496-499. Kleiner wie Alexis; dasselbe

Blau, Vorderfliigel stumpfer, die Unterseite so grau wie bei

Donzelii, die Augen sehr scharf. — Aus Kleinasien.” and, in Heft 49,

in his first volume on plate 104, Figs 496-499 (also [1851]),

depicts respectively one d on the upperside and in resting position

on the underside, and one © in the same positions, again as “Actis”

(Fig. 1 of the present study). Subsequently, in the “Systema Lepi-

dopterorum Europae. Systematisches Verzeichniss der Europäi-

schen Schmetterlinge”, published in the sixth volume of his

“Systematische Bearbeitung”, he lists on p. 4 (Heft 55, [1852]):

“23. Lycaena. ... 235. Actis.”, while in the Index of his first volume

(Heft 65, p. [1] - [1854]), he mentions “actis (Lycaena) Kad.

pp 113 [sic! on that page in Vol. 1, Acis is quoted as synonym of

[Lycaena] Argiolus L.] HS. 466-499. Athis Fr. [the latter name thus

being quoted as a synonym]”’. Finally, in Vol. 6, on p. 172 (Heft 69

— 1856) he writes: “Actis dazu Atys Gerh. 19.3. a-d.”, thus also con-

sidering the latter name as a synonym of the former. In all, very lit-

tle information is contained in all data communicated by Herrich-

Schaffer. Carl Gotthelf Kaden (1786-1867) apparently named this

butterfly in litteris and communicated it as such to Herrich-

Schaffer. The obituary published by Staudinger (1868) gives no

evidence that Kaden ever travelled to Asia Minor, or perhaps even

abroad, hence he must have obtained his material from that coun-

try through exchange or — much more probably — purchase. The

figures of actis given by Herrich-Schäffer show a rather robust but-

terfly, of which the male wing upperside is blue with a very narrow

black marginal line and black underlining of the veins in their dis-

tal fourth, but without any trace of black suffusion in the distal half

of the hindwings. The dating of the various instalments (“Hefte”)

referred to above follows Hemming (1937) and is given on the

wrapper to each Heft. Unfortunately, a survey of contemporaneous

literature did not allow the present author to date those instalments

more precisely, i.e. as to the month of publication, hence they all

should be dated 31.XII. of the year concerned (ICZN, Art. 21.3.2).

Gerhard ([1851]) describes on p. 11 (Heft 5 — 31.V.1851, cf.

Olivier, 1999) a blue as follows: “[Lycaena] Atys. Kind. Türkei.

Female Schaeft. Platte 19. Fig. 3. a, b, 6, c, d, 2. Diese von

Herrn Kindermann entdeckte seltene Art nannte Herr Dr. Herrich-

Schaeffer in Regensburg “Actis”, da der Name Arys schon unter den

Exoten sich befindet.” The quality of the figures on plate 19 is not

very good, but one can again discern a butterfly with a very narrow

black marginal line, and absolutely no traces of black dusting in the

apical area of the hindwings (Fig. 2 of the present study). His refer-

ring to Herrich-Schäffer could be interpreted as indirect evidence

for Herrich-Schäffer’s name being published before his own atys,

but this is not necessarily so. The name actis may have circulated

on sales lists or may have become known to Gerhard vıa Lederer,

before Herrich-Schäffer actually published it, as ıs established for

the names asabinus and ochimus (Olivier, 1999: 129). The most

interesting information contained ıs perhaps, that both Herrich-

Schäffer’s actis and Gerhard’s atys were named after material col-

lected by Albert Kindermann Jr.

Finally, Freyer (1851), in his sıxth volume (Heft 96), describes

on p. 147 “Lycaena Pap.[ilio] Athis” as follows: “Der Mann von

dieser Lycaena gleicht auf der Oberseite gewöhnlich dem

P. Alexis.’ He further writes (pp. 147-148): “Eine sehr schöne

Abart desselben, die dunkler Blau und von der Außenseite gegen

die Flügelmitte, in eine schwarze Mischung auf der Oberseite ver-

läuft, zeigt meine Fig. 2. [Fig. 3 of the present study]. Man könnte

diese Abart wohl für verschieden vom gewöhnlichen Manne halten,

wenn erwiesen wäre, daß solche öfters vorkömmt, worüber mir

Herr Kindermann keine Nachrichten ertheilt hat. Das Weibchen ist

auf der Oberseite einfach braun wie das von P Acis, Die Unterseite

hat d und © ganz übereinstimmend mit P Damon, so daß sich

beide Arten von solchem nur durch die geringere Größe unter-

scheiden. Es zeigen sich wie bei Damon, auf den Vorderflügeln das

schwarze Mittelauge und die aus großen Augen bestehende weiß

eingefaßte Augenbinde, und die Hinterflügel führen die kleine

Augenreihe mit den weißen Splitterstrich durch die Flügelmitte

vom Wurzelfelde aus. Fühler wie bei den vorhergehenden Arten.

Heimath: Amasia.”. Beside the interesting information that the type

locality of athis is Amasia and that Kindermann again is the pro-

(94

pr, me IH Pr < a Res h

Lars ders... a

OOM? FIG Fels (SOLE HH Corte FIR arc Tl DAR

Fig. 1. Plate 104 from G. A. W. Herrich-Schäffer’s “Systematische Bearbeitung der

Schmetterlinge von Europa, zugleich als Text, Revision und Supplement zu Jakob

Hübner’s Sammlung europäischer Schmetterlinge”, Vol. 1, Heft 49 [31.XII.1851].

Hal IR

4 42

ZH A Le i z AH ue LL 2 a

1. y 1 . . ee loi / a 2

Fig. 2. Plate 19 from P. B. Gerhard’s “Versuch einer Monographie der europäischen

Schmetterlingsarten: Thecla, Polyomattus [sic], Lycaena, Nemeobius. Als Beitrag zur

Schmetterlingskunde”, Heft 5 [31.V.1851].

vider of the material, the description consists of two parts: first, the

‘nominotypical’ taxon looks on the upperside like “P alexis” (a

junior primary homonym of Papilio alexis Poda, 1761 [currently

Glaucopsyche alexis (Poda, 1761)] and a senior subjective syn-

onym of Papilio icarus Rottemburg, 1775 [currently Polyommatus

icarus (Rottemburg, 1775)]) and hence again as a blue with a very

narrow black marginal line; second, an “Abart” [to be interpreted as

form rather than as geographical race or, in present-day terminolo-

gy, subspecies] is described but not named (!), the d of which has

a darker blue wing upperside with extensive black dusting of the

apical area, thus suggesting that a distinct taxon is actually

involved. It ıs the latter form that is figured by Freyer. The dating

of Heft 96 was established from the printed date on the wrapper

(Tremewan, 1988), while a search through contemporaneous liter-

ature could not reveal more precise data (Olivier, in press).

At this stage, it is worthwile to report on the travelling activities

of Albert Kindermann Jr. near Amasia, in order to establish the pre-

cise origin of the material that served to describe actis, atys and

athis. In March 1848, Kindermann left Ofen [Budapest], along

with Stefan Nogell, for a planned journey to Russia (Nogell would

collect in ‘Grusien’, i.e. Georgia), but the Russian authorities

refused to let him go there and so they were forced to travel to

Anatolia. They arrived in Samsun on 3rd May and stayed there for

three weeks. As the vegetation was not well out yet and as very few

butterflies were on the wing, Nogell decided to move to Amasia,

where Kindermann joined him at the beginning of July. From here,

Kindermann explored snowy mountain ranges at six hours walk to

the north of town, at altitudes up to 6,000 feet (2,000 m), where he

collected very well, but to the end of the year he travelled through

Triest back to Ofen. In 1849 he went back to Amasia, this time col-

lecting near Tokat, that is surrounded by even higher mountains,

but added only few new species. He returned back again in autumn,

sending his material to his colleagues when in Ofen. In May 1850

he collected near Diabekir (now Diarbakir), in June near Sivas and

in July near Tokat. His assistant (Nogell) concentrated his activities

in the mountains near Tokat in 1848 and 1849 and returned back to

Ofen only in September of the latter year (Lederer, 1855; [Lederer],

1860; Staudinger, 1878). In his “Verzeichniss der von Herrn Albert

Kindermann 1848-1850 um Samsun, Amasia, Tokat, Siwas und

Diarbekir gesammelten Schmetterlinge” (pp. 235-254), Lederer

CSS 257) lists “[ycaena| Actis H.-Sch. ... Auf den

Tokateralpen”. It is quite possible, however, that Kindermann also

collected material elsewhere, as Staudinger (1878: 181) quotes:

“Auch konnte Lederer nicht alle von Kindermann gefundene Arten

aufführen, da derselbe, wenigstens an Gruner bestimmt, direkte

Sendungen machte die Arten enthielten, welche er an Lederer nicht

sandte”. In the systematic part of his publication however,

Staudinger (loc. cit.: 254-255) reports that it was found by

Kindermann “in den Tokat-Alpen”. He further mentions: “Drei alte

36 (aus Lederer’s, Weissenborn’s und Sommer’s Sammlung),

welche ziemlich sicher von Kindermann bei Tokat gefangen wur-

den, sind auf den Hinterflügeln mehr schwarz als blau”.

Interestingly, Staudinger (1899: 147) explicitely states: “Freyer

giebt zwar Amasıa als Fundort seiner Arhis an, aber fast zweifellos

stammt sein d von Tokat, da weder ich, noch irgend ein anderer

Sammler (soweit mir bekannt) L. Athis bei Amasia fand”. It thus

appears that both “forms”, 1.e. “typical actis” and the “Aberration

... Abart” with blackish suffusion in the distal part of the hind-

wings, originate from the surroundings of Tokat, rather than from

Amasia.

Taxonomic interpretations and quotations of the names actis,

athis and atys in literature

Heydenreich (1851: 13) was not yet in the possession of the crit-

ical instalments of Gerhard, Herrich-Schaffer and Freyer when

writing his catalogue (cf. Olivier, 1999, in press and in prep.) and

published under nr. 337 the name “[Lycaena] Atys Kind. i. 1.”,

which is a nomen nudum, as it is not accompanied by any descrip-

tion or reference to another work. Keferstein (1851: 305) only had

Gerhard up to Heft 5 and hence quoted under nr. 183 “[Lycaena]

Damon Var. d Atys Gerhard tab. 19. f. 3. Amasia” and apparently

the same applies to Meyer-Dür (1852: 94) who also listed the name

“[Lycaena] Atys Kinderm.” among the “Varietäten” of “Lycaena

Damon O.”.

gO Tab: 575

2.3. hs 4 Gerypiluus.

ft mpyrea

Fig. 3. Plate 573 from C. F. Freyer’s “Neuere Beiträge zur Schmetterlingskunde mit

Abbildungen nach der Natur”, Vol. 6, Heft 96 [31.XII.1851].

8 9

Fig. 4. Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851), lectotype 4 (upper-

side), [Turkey, Tokat province], “Amasia” [recte Tokat], [VII.1849], leg. Albert

Kindermann, in coll. Museum für Naturkunde der Humboldt-Universität zu Berlin.

Fig. 5. Same as Fig. 4 (underside).

Fig. 6. Same as Fig. 4 (labels, the one containing the lectotype designation not included).

Fig. 7. Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851), paralectotype 9

(upperside), [ Turkey, Tokat province], [“Amasia”, recte Tokat], [VII.1849], [leg. Albert

Kindermann], in coll. Museum für Naturkunde der Humboldt-Universität zu Berlin.

Fig. 8. Same as Fig. 7 (underside).

Fig. 9. Same as Fig. 7 (label, the one containing the lectotype designation not included).

Lederer (1852: 21) quotes “Actis H.-Sch. 496-499”, with “Athis

Freyer 575” as synonym, while later (Lederer, 1869: 81-82) he

considers actis as only a variety of damon. It is clear that, for him,

typical actis is violet blue and “der schwarze Rand fehlt (Actis H.

Sch.)”, while he further also mentions “unter Actis eine Varietat mit

fast ganz schwarzen Hinterflügeln”. He further reports actis from

Astrabad (now Iran, province of Mazandaran, Gorgan), the Taurus

Mts. (own observations), as well as from Achalzich (now Georgia,

Akhaltsikhe), Hankynda (now Nagorno-Karabach (disputed:

Armenia vs. Azerbaijan), Xankändi; known also as Stepanakert)

and several mountainous areas of Transcaucasia (leg. Haberhauer;

See also Wedenenmulon Us):

In the first edition of his catalogue, Staudinger (1861: 6), under

nr. 149, writes: “[Lycaena] Actis HS. 496-9. (1851); Athis Frr. 537,

2. 3. (Febr. 1852); Atys Gerh. T. 19, 3 a-d.”, thus also considering

actis as the valid name for this taxon. In the second edition of his

“Catalog”, Staudinger (1871: 13) considers actis again as a “vari-

etas” of L. damon, stating under entry 172g: “v. Actis HS. 496-9

(1851), VI p. 24; Ld. Hor. |. c. Athis Frr. 573, 2 3° (Gebmeiaas)):

Athys Gerh. PI. 19, 3a—d; ? Iphigenia Gerh. 20. 2 (minor, violaceo-

caerul., 4 interdum nigricans)” and reporting it from “As.min;

Arm. et Hyrc.; Mont. et Alp.”. Staudinger (1878: 251) lists

“[Lycaena] Actis HS.; Ld.; Athis Frr.; Athys Gerh. Carmon Ld. pro

parte. — Pont. alp. (Tokat); Lyd. m. alp.; var: Hyrc. et Arm.”. It is

clear that, for Staudinger (loc.cit.: 254—255) as well, typical actis

“hat ... einen weit schmäleren schwarzen Aussenrand der Flügel”.

He further notices that “Die Actis aus Armenien und Nord-Persien

sind nicht so tiefblau wie die kleinasiatischen, indessen sind sie

doch stets rein blau, etwa wie bei /carus, nie so grünblau wie

Iphigenia, haben auch einen schmalen schwarzen Randsaum, wie

die typischen Actis” and, finally, apparently considers actis as

being a distinct species again. Staudinger (1886: 214-215)

describes the nominal taxon “Lyc.[aena] Actis var. Actinides Stgr.

... aus dem südlichen Alai [now Tajikistan: Transalai Mts.]”.

Staudinger (1899: 146-147) repeats by and large what he said in

1878, adding “aus verschiedenen Orten des Taurus, aus Malatia

(westliches Kurdistan) und von Gümüschane (nordöstliches

Pontus-Gebiet)” to the known localities, but now applies the name

Athis to the “form” with extensive black suffusion: “Dahingegen

scheinen die von Kindermann bei Tokat gefangenen Actis 6 4

durch stärkeres Auftreten von Schwarz besonders auf den

Hinterflügeln (wo es bei 2 meiner 3 dd das Blau fast ganz ver-

drängt hat), vielleicht vorwiegend verschieden gewesen zu sein, so

dass sie in diesem Falle als Lokalform, jedenfalls aber als auffall-

ende Aberration, den Namen Afhis Freyer (unter welchem Freyer,

Taf. 573 Fig. 2, ein solches d abbildet) führen können”. He further

elevates actinides to species rank, which is currently still the taxo-

nomic status of this butterfly. Staudinger & Rebel (1901: 88, nr.

626) list “[Lycaena] Actis HS. 496-9 (1851) VI p. 24; Ld. Hor. VI

p- 81; Stgr. Hor. XIV p. 254; Iris XII p. 146; Athys Gerh. t. 19, f. 3

a-d.” from “Pont; Arm; Hyrc; Kurd. oc; Taur.” and, as a separate

entry, “a) v. (an ab?) Athis Frr. 573, 2 & (3 2) (& plus minusve

praesertim in al. post. nigro-mixtus)” from “Pont. (Tokat); As. min.

c. (Konia)”.

Bienert ([1869]: 46) lists “Lycaena Actis” from Amasia and

Tokat, while Kirby (1871: 370) writes: “237. c. C. Actis, Herr.

Schäff. (Lyc. A.) Schmett. Enr. (sic!) I. f. 496-499. (1851); Led.

Hor. Soc. Ent. Ross. VI. p. 81 (1869); Lyc. Athis, Frey. Neuere

Beitz [sic!]. VI. t. 537. f. 2, 3. (1852); Lyc. Damon, var. Atys, Gerh.

Mon. Lyc. t. 19. f. 3 a-d. (1853). ib. Asia Minor.” Christoph (1873:

26) mentions “Lycaena Damon ... var. Actis HS.” as “Nicht eben

selten bei Hadschyabad [now Iran, province of Mazandaran,

Hajıbad] und Schahkuh [now Iran, province of Mazandaran,

Shahkuh]”. Romanoff (1879: 487) reports Lycaena Damon Schiff.

v. Actis HS. from the “Haut-Plateau Arménien”: “Très abondante

sur le mont Ben-gel et Zchniss-Kala” (now in southwestern

Armenia and northeastern Turkey), while Bramson (1890: 42), in

his key to the genus Lycaena, lists among the characters of “Actis

H. S.”: “Der dunkle Saum der Obs. sehr schmal, Rippen am Saume

nicht schwarz. d oben tief hochblau, bisweilen schwärzlich

bestäubt, besonders auf den Htfl.”, thus suggesting that he also con-

siders typical actis to lack any extension of black suffusion. He fur-

ther lists as synonyms: “Syn. Afhis Frr. Athys Gerh. Carmon Ld.

pro parte. Damon var. (Stgr. Cat. 1871 No. 172 g)” and gives as

range: “Oestlicher Caucasus, Transcaucasien”. Groum-Grshimaïlo

(1890: 409-412), dealing with “Lycaena Actis HS. var. Actinides

Stgr.”, states a. o. (p. 411): “les d 4 sont d’une coloration plus lilas

que p. ex. les Actis de Kasikoparan, moins brillants et à bordure

noire plus large”, thus also treating the latter taxon as one lacking

any black suffusion on the male wing upperside. Rühl (1893: 290)

writes: “L. Actis H. S., Athis Frr, Athys Gerh. Wenig kleiner

(Exemplare eben so gross wie Kindermanni kommen jedoch vor).

Vorderflügel und Hinterflügel mit verändertem, in’s Violette

ziehenden Blau, Schatten am Aussenrand schmäler und schwächer,

Rippen nicht schwarz einmiindend. 4 sehr selten schwärzlich,

besonders an den Hinterflügeln bestäubt, ... Fluggebiet: in

Transkaukasien, bei Hadschyabad und Schahkuh (Persien). Heyne

(1895: 765) adds: “S. 290. L. Actis H. S. Weitere Fundorte:

Oestlicher Kaukasus, Kleinasien, Armenien”. It is thus clear that,

for Rühl also, typical actis is the blue with a very narrow black

marginal line. Holtz (1897: 47) writes about “L. actis H.-S.”: “Von

Lederer und Haberhauer an denselben Ortlichkeiten gefangen wie

die vorıge Art [“In höheren Teilen des Taurus von Lederer und

Haberhauer gefunden”]”. Radde (1899: 420), in his list of the

Lepidoptera collection housed in the Museum Caucasicum (which.

is virtually a part of Romanoff’s collection), states “[Lycaena] actis

H. S. Kasik[o]p.[oran]”.

In a faunistical paper on the butterflies of Crimea, Melioransky

(1897: 220) lists “Lycaena actis H. S. — a single specimen col-

lected in July 1893” — but without doubt he misidentified a differ-

ent species (cf. Forster, 1960: 107; Nekrutenko, 1985: 139). The

report of “Lycaena damon var. actis H.-S.” from the Rilo Dagh in

Bulgaria by Bachmetjew (1902: 404) is also based on a misinter-

pretation (Forster, loc.cit.: 107, footnote).

Seitz (1909: 318) lists “L. actis A.-Schäff. (= athys Gerh.)

(82a).”, also considering the totally blue butterfly as typical, while

“Bei der Form athis Frr. (82a) ist der Rand der Vflgl und die ganze

Aussenhälfte der Hflgl von der blauen Bestäubung freigelassen und

erscheinen tiefschwarz”. Courvoisier (1910: 192) mentions “Actis

H. S. mit Var. Athis Frr.’ as a distinct species, while Courvoisier

(1914: 152) lists actis H Sch., with “atys (Kindermann) Ghd. 1853”

as a synonym and “athis Freyer” as a “Nebenform”. Le Cerf (1913:

69) merely lists “L. actis H.-Sch.” in his account on the butterflies

of Persia.

Wagner (1929: 62) reports “Lycaena actis H. S. und var. athis

Frr.” from the Sultan Dagh (now Sultan Daÿlari, in the provinces

of Afyon, Isparta and Konya) north of Aksehir, in the western

Taurus. Osthelder & Pfeiffer (1932: 48) list “[Lycaena] actis H.

Sch. ssp.” in their account of the butterflies from Marash, also quot-

ing in their brief description of the butterfly: “Das wunderbare Blau

besonders leuchtend und ohne jede Saumlinie”. Schwingenschuss

(1935: 132) publishes a report of butterflies collected in the neigh-

bourhood of Bolu (June) and in Aksehir and on the nearby Sultan

Daglari (July). From the latter area, he reports “Lycaena athis Frr.”

and “athis nov. ab. lunulata Schw.”. As the name refers to an infra-

subspecific form, it is unavailable, the more it is not accompanied

by any description. Subsequently, Schwingenschuss (1938: 146)

quotes “Lycaena actis H. S. ssp. athis Frr.”, thus for the first time

unambiguously using this name at subspecies level, and applying it

to the taxon with blackish apical suffusion, as he further writes:

“Kommt nur in der ssp. athis mit der ganzen Variationsbreite vor

(PF). Ich habe auch nur athis gefunden, darunter Stiicke mit

prächtig blauen Monden auf den Hinterflügeln, für die ich den

Namen lunulata in Vorschlag brachte (Verh. d. z. b. G. 1935, S.

132) (ScH)”. His name /unulata does not become available by this

action, as it stays infrasubspecific. Schwingenschuss (1939: 88)

lists “L. actis H. S.” from the central Elburs Mts. in northern Iran,

reporting it “Im Tarseegebiet (Pf., Sch., W.) und bei Pelur (Sch.) in

ganz wenigen, meist schlechten d 4 erbeutet”, thus also consider-

ing the name actis to apply to a blue without any black suffusion on

the upperside.

Forster (1936: 59; 1938: 116) merely lists “actis H.-Sch.” but, in

his famous “Bausteine”, for the first time since Staudinger (1899),

deals with the subgenus Agrodiaetus in detail, undertaking a series

of taxonomic actions which are of great importance for a further

understanding of this story. Under “Agrodiaetus damone Ev.”,

Forster (1956: 58-59, Tafs 8-9, Figs 16-17) describes the nominal

taxon “A. damone firdussii ssp. nov.” from Shahkth in the eastern

Elburs (northern Iran), listing in its synonymy “actis pro parte

Staudinger, Horae Soc. Ent. Ross. 14, 1878, p. 255” and “actis

Schwingenschuss, Entom. Zeitschr. 53, 1939, p. 87”. Later, Forster

(1960: 106-107, Tafs. 10-11, Figs 1-3) deals with “A. actis actis

H. Sch.”, listing “athys Gerhard”, “damon var.’ sensu Staudinger

(1871: 13) and both “f. athis Freyer” and “f. lunulata Schw.”

among its synonyms. He considers his d material to consist of 20

actis and 70 athis, the latter being a mutation that is a little smal-

ler, a deeper violet blue and, especially on the hindwing, has an

extensive black suffusion, and believes that “Ende des 19.

Jahrhunderts war die f. athis Frr. offensichtlich noch wesentlich

seltener, wie aus der Literatur und dem untersuchten alten Material

zu entnehmen ist”. As type locality (“Loc. class.”) he mentions

“Kleinasien”. He further describes “A. actis pseudactis ssp. nov.”

(Forster, loc.cit.: 107-109, Tafs 10-11, Figs 4-5) from Armenia,

that is a more bright blue, including among its paratypes material

originating from localities now in the northeast of Turkey, 1.e. in the

Aras Valley (Kagysman, Kulp, Kazıkoparan, Agrı Dagh,

Geliagadzha (now resp. Kagizman, Igdır, Kazıkoparan, mountains

south of Kazıkoparan, untraceable locality near Kagızman, in the

provinces of Kars and [&dir)), as well as material collected by de

Lesse in 1956 (“Col du Kop Dagh entre Bayburt et Erzerum”

(Kopdagi Geçidi, at the border of the provinces of Bayburt and

Erzurum), “Mirgemir Dagh entre Erzurum et Agri” (Tahir Geçidi,

province of Agri), “Col routier entre Erzerum et Agri’), and further

13, 1? from Ordubad (now in Naxcivan). Interestingly, Forster

(loc.cit.: 109) further lists material from’ “these buns,

‘Mesopotamia’, “Kurdistan” (incl. vic. Malatya), “Zeitun” (now

Süleymanlı in the province of Maras) and the Taurus (both Lydia

and the Cilician Taurus!) as belonging only conditionally to

pseudactis, and possibly representing an undescribed subspecies. It

is worth mentioning, that Wiltshire (1957: 30) reports

“Polyommatus actis H.-S. ssp. pseudactis Forst.’ from Iraq “At

great heights in the mountains’, three years before Forster’s

description! As no definition of the taxon is given, Wiltshire’s name

is anomen nudum.

In the late 1950’s and early 1960’s, de Lesse carried out exten-

sive karyological studies on Agrodiaetus taxa from Turkey and

Iran, his results still forming the basis of much of our current views

on taxonomy in this difficult group. In his thesis, de Lesse (1960:

186-187), on karyological evidence, transfers the nominal taxon

firdussii to the rank of subspecies of “A grodiaetus actis”: from his

100

discussion, it further appears that, for him, actis is also a blue

without any black suffusion in the apical part of the wings. On his

table on p. 207, and on fig. 219 on the next page, he lists “A. actis

pseudactis Forst. in litt.” from “Mirgemir D.” and “Kop Dagh” and

“A. actis firdussi Forst.’ from the Demavend (Elburs Mts.). In a

separate paper, de Lesse (1962) presents new data on the chromo-

somal variation in “Agrodiaetus actis”, including results for one

population from Biiriicek in the Taurus Mts. (province of Icel,

Tekir) that, according to him (p. 67), may belong to a new sub-

species, “ainsi que l’a récemment indiqué Forster”. This population

is likely to belong to the newly described P (A.) sigberti Olivier,

van der Poorten, Puplesiene & De Prins, 2000 (cf. Oliver ef al.,

2000).

Higgins (1966: 213), in his checklist of Turkish butterflies, lists

under nr. 118: “Agrodiaetus actis Staudinger. “Taurus”; 28 as

lunulata Schwingenschuss; 43 as athis Freyer (42)”. His publica-

tion being based on a survey of the literature, one cannot tell which

butterfly Higgins considers to be actis. Goossens & Cromphout

(1978) and Goossens (1979), in two accounts on successive col-

lecting trıps in Turkey, include “Agrodiaetus actis Staudinger”

from the province of Icel in a list of butterfly taxa, following the

concept of Higins’s (1966) paper. Again, one cannot tell what they

understand under actis.

Kocak (1979: 323-324) describes “A grodiaetus (s. str.) pseudac-

tis sertavulensis n. subsp.” from the Sertavul Gecidi in the province

of Konya in Turkey, placing it in the “actis-pseudactis group” but

rather closer to pseudactis, according to wing shape and coloration.

He further considers that the ranges of both actis and pseudactis

overlap partly in the Taurus Mts. and consequently proposes

pseudactis to be a distinct species from actis. Subsequently, Kocak

(1980: 19), as a result of a study on the publication dates of

Gerhard’s “Versuch einer Monographie der europäischen Schmet-

terlingsarten ... (1850-1853)”, concludes that [Lycaena] actis

Herrich-Schaffer, [1851] (December) is a synonym of Lycaena atys

Gerhard, [1851] (May). In his “Critical check-list of European

Papilionoidea”, Kocak (1983: 33) lists “[Agrodiaetus] atys

Gerhard, 1851” (with actis Herrich-Schäffer, 1852 as synonym)

and “[Agrodiaetus] pseuactis Forster, 1960” [sic!], quite in agree-

ment with the conclusions arrived at in his two previous papers.

101

Eckweiler & Hofmann (1980: 20) list two subspecies of

“Agrodiaetus actis Herrich-Schäffer, 1851” in their “Checklist of

Iranian butterflies”, 1. e. ssp. pseudactis Forster, 1960 (provinces of

[West- and East-] Azerbeijan) and ssp. firdussii Forster, 1960

(Elburs), while Eckweiler & Görgner (1981: 105) report

“Agrodiaetus actis Herrich-Schäffer, 1851 ssp. pseudactis Forster,

1960” from the Palandöken Mts. in the province of Erzurum, in

northeastern Turkey. In an important taxonomic paper, Eckweiler

(1989: 97) considers, in full agreement with Kocak (1979), that

pseudactis and actis are distinct species as, according to him, both

appear to be syntopic and synchronous ın the Taurus range.

Eckweiler (pers. comm. 29.1V.2000) applied the name actis to the

taxon we now know as P. (A.) sigberti. He further underlines the

conspecificity of firdussii Forster, 1956 and pseudactis Forster,

1960 and establishes the following new combinations:

Polyommatus (Agrodiaetus) firdussii firdussii (Forster, 1956)

(Elburs), P. (A.) firdussii pseudactis (Forster, 1960) (East Anatolia)

and P. (A.) firdussii sertavulensis (Koçak, 1979) (Taurus). He

describes a new taxon, “Polyommatus (Agrodiaetus) firdussii

ernesti subspec. nov.”, from near Elmalı in the Kohu Daÿlar in the

Turkish province of Antalya, in the western Taurus, while admitting

that specimens transitional to sertavulensis (according to underside

markings) occur on the Palaz Dagi, in the eastern part of the

province of Antalya. He finally figures (Abb. 43-46) “Poly-

ommatus firdussii trans. ad sertavulensis Kocak” from the Turkish

province of Nigde (Camardı, Cukurbag).

De Prins et al. (1991: 146 and plates 1 & 2, figs 11 & 14) list

material from the province of Van, in southeastern Turkey, as

“Polyommatus (Agrodiaetus) firdussii (Forster, 1956)” and so do

Kocak & Seven (1994: 6, as “Agrodiaetus firdusii Forst.” [sic!]),

while Avcı & Ozbek (1996: 84) also name “Polyommatus firdussii

Forster, 1956” material from the province of Erzurum. Seven

(1996: 28-29) reports this taxon among material collected “in

Central and North Turkey”. Lukhtanov et al. (1998) ascribe mater-

ial from different localities in the southeastern Turkish provinces of

Bitlis and Van to “P. (A.) firdussii pseudactis (Forster, 1960)”.

In their monumental work on the butterflies of Turkey,

Hesselbarth et al. (1995) apply the species group name actis

102

Herrich-Schäffer to the smaller species, the d of which is more

violet blue, often with a more or less extensive blackish suffusion

in the distal half of the hindwing (cf. Tafel 123, Figs 1-21), that had

been ascribed to athis Freyer by several earlier authors, from the

second half of the 19th century up to Forster (1960). In their inter-

pretation, they consider both atys Gerhard and athis Freyer as sub-

jective synonyms of actis Herrich-Schäffer and further give as

chromosome number “n = 27, Bürücek (Icel) (de Lesse, 1962a:

68)”. Their interpretation (and that of Eckweiler, vide supra) is

quite understandable, when one knows of the arrangement of

Forster’s material in coll. Zoologische Staatssammlung München:

under “actis actis H.S.”,46¢, 79 of P. (A.) sigberti (now labelled

as paratypes) are included at present, while among “actis pseudac-

tis Forst.”, 14 of P. (A.) actis from “Kleinasien Amasia Korb” and

96 from various localities in the Taurus Mts. (all violet blue, quite

different from sertavulensis) are included along with P. (A.) “wag-

neri” (Forster, 1956) from “Konia” (23) and “Lydia Cilicie

Taurus”. The larger species without any blackish suffusion is

named P. (A.) firdussii (Forster, 1956) and pseudactis (Forster,

1960) is sunk in synonymy (cf. Tafel 122, figs 37-72).

Nevertheless, the authors leave open the possibility that “firdussii”

consists of more than one taxon on karyological evidence. P. (A.)

sertavulensis (Koçak, 1979) is raised to species rank on morpho-

logical-chorological data, as on Sertavul Gecidi they found both

species (sertavulensis and wagneri — not firdussii! — cf. locality

lists on pp. 728-733) syntopic and synchronous; they further syn-

onymize ernesti under sertavulensis.

Hauser & Eckweiler (1997) list alphabetically all the species

group taxa dealt with so far in the present overview, with their

original combination, type locality and data to their primary

sources (literature reference as well as current depository of extant

types (when existing), while Eckweiler & Hauser (1997), in a sys-

tematic checklist, arrange the species group taxa into different

unnamed species groups. All taxa relevant to the present study are

placed in a very inclusive, large “A. damon group”, and the nomi-

nal taxa actis (sensu Hesselbarth et al., 1995!), firdussii and ernesti

are figured in colour. The names ernesti, sertavulensis, pseudactis

— but also maraschi Forster, 1956 — are associated as subspecies

to firdussii. The latter taxon was described by Forster (1956:

103

57-58) as a subspecies of “A. damone”, but is currently considered

to represent either a junior synonym or a subspecies of P. (A.) wag-

neri (Forster, 1956) (Hesselbarth ef al., 1995; Carbonell, 1998). In

the Zoologische Staatssammlung München, I found 46 of actis

from “Syr. sept. Taurus Marasch”, that clearly differ from the holo-

type and 2d paratypes of maraschi, that were also collected in the

same locality, in being darker violet blue, with the spots and the

submarginal lunules on underside hindwing much more apparent.

The holotypes of wagneri and maraschi are also very different and

it thus appears to the present author that maraschi is a taxon quite

distinct from both actis and topotypical wagneri.

Balint & Johnson (1997: 19) list actis and firdussii without any

further subdivision, while Balint (1999: 31) lists 10 4, 2 2

paratypes of firdussi [sic], deposited in The Natural History

Museum, London.

Carbonell (1997) presents an historical overview of the “com-

plexe actis-firdussii-pseudactis”, also applying the names actis and

firdussii in the sense of Hesselbarth et al. (1995) and describes “A.

actis artvinensis n. ssp.” from the extreme northeast of Turkey

(provinces of Erzurum and Artvin), that is characterized by the

even darker violet blue colour with traces of a discoidal spot, as

well as the extreme development of the black suffusion on both

fore- and hindwings on the upperside, and by the dark grey-brown

ground-colour with sharply contrasting white streak on the under-

side. He further states about the specimens of “actis ssp?” from

Biiriicek, of which de Lesse (1962) studied the karyotype: “la

majeure partie des d d concernés ont effectivement les nervures

noircies sur plusieurs mm, mais sans aucune trace de suffusion sub-

marginale noire”. Carbonell (1998) reports the sympatry of

“pseudactis” and sertavulensis on the Sertavul Gegidi. He lists

pseudactis as a subspecies of firdussii and, in accordance with

Hesselbarth et al. (1995), considers sertavulensis as a distinct

species, but ernesti as a valid subspecies of the latter. Carbonell &

Naderi (2000: 219) again quote “A. actis artvinensis Carbonell,

1997” in the heading on the differential diagnosis of their new

taxon Agrodiaetus arasbani. Reinhard & Eitschberger (1999: 350),

following Häuser & Eckweiler (loc.cit.) and Eckweiler & Häuser

(loc.cit.), present the same arrangement and further add artvinensis

as a subspecies of actis (sensu Hesselbarth er al., 1995).

104

Lately, Kocak (1996: 22-23) has listed “Polyommatus (Agro-

diaetus) atys” and P. (A.) firdussii as two different species, the lat-

ter with four subspecies (firdussii, pseudactis, sertavulensis and

ernesti). It is not stated however if Kocak (loc.cit.) and Kocak &

Seven (1996) interpret “atys” as “actis sensu Hesselbarth er al.

(1995)” or in a different way, but Kocak & Seven (1998) probably

do, as they link artvinensis (Carbonell, 1997) as a subspecies to

“atys (Gerhard, [1851])”. They also list maraschi as a subspecies of

firdussii (presumably following Eckweiler & Häuser, 1997).

Lectotype designation

After Herrich-Schäffer died in 1874, his private Lepidoptera col-

lection was split up via Staudinger & Bang-Haas and part of it went

via the private collection of Otto Staudinger to the Zoological

Museum of Berlin [now Museum für Naturkunde der Humboldt-

Universität zu Berlin]; many Lepidoptera described from the

Kaden collection by Herrich-Schäffer became isolated in various

collections (Horn ef al., 1990).

While visting the Museum für Naturkunde der Humboldt-

Universität zu Berlin in late November 1998, I was shown a box by

Dr. Yuri P. Nekrutenko, that included two apparent syntypes (4, 2)

of [Lycaena] Actis. In their primary position in the collection, both

specimens were placed behind a bottom label attached to the first

specimen (G ); this specimen also bears a printed numerical label

“4723” to refer it to the handwritten inventory book entitled

Catalogus Generalis Musei Zoologici Berolinensis. Vol. 1 (Nos.

1-3753) Vol. 2 (Nos. 3754-10098), that is deposited in the

Lepidoptera section library; each entry (row) of the Catalogus con-

sists of five columns: number (shown on the printed label) | name

and author | number of specimens | locality | collector or provider.

In the “Catalogus”, under “4723” is written: “Lycaena Actis HSch.

| 2 | Amasia | Kindermann”.

I herewith designate the d as lectotype (Figs 4-6) and the © as

paralectotype (Figs 7-9):

Lectotype d with labels: handwritten “Actis | HSch. f. 496-971

Atys Kind.[ermann] ms. Gerh.[ard] | t. 19. f. 3. | Amasia <recte

105

Tokat — vide supra> Kind[ermann]” (recto yellow, in printed box,

verso white); printed “4723” (on white paper), “actis Herrich-

Schäffer, [1851] | SYNTYPUS | Y. Nekrutenko det. 30.11.1998”

(on yellow paper); designated now with printed label “[Lycaena]

Actis | Herrich-Schäffer, 1851 | LECTOTYPUS à | design. Olivier,

2000” (on red paper).

Paralectotype ® with printed labels “actis Herrich-Schäffer,

[1851] ISYNTYPUS | Y. Nekrutenko det. 30.11.1998” (on yellow

paper); designated now with printed label “[Lycaena] Actis |

Herrich-Schäffer, 1851 | PARALECTOTYPUS © | design. Olivier,

2000” (on red paper).

Type locality. Since label and stated type locality (“Amasia”)

are found to be erroneous (see Introduction above), it is corrected

here according to the available data as Turkey, Tokat province, env.

Tokat (ICZN, Art. 76.2, Recommendations 74E, 76A.2).

Taxonomic consequences

Nomenclature. Koçak (1980: 19) proposed “Agrodiaetus atys

(Gerhard, [1851)” as the valid name for this species, as it was

published in May 1851, thus taking precedence over [Lycaena]

actis Herrich-Schäffer, [1851] (December), the latter name further-

more having been published uninominally. Except for Heydenreich

(1851), Keferstein (1851) and Meyer-Dür (1852), who did not pos-

sess Hefte 48 and 49 of Herrich-Schäffer’s work, all authors except

Kocak, from the second half of the 19th century up to 1995, used

the name actis for this taxon. Indeed, the name actis just looks like

published without combination with a generic name, but it is obvi-

ous that Herrich-Schaffer consistently applied the Principle of

Binominal Nomenclature and ascribed it to the genus Lycaena,

likewise all the “blues” in volume | of his work are placed in this

genus (ICZN Art. 11.4). It is therefore agreed here with Hesselbarth

et al. (1995: 732) that the name actis should be considered as the

valid name for this taxon, thus promoting stability of nomenclature

(see also ICZN, Preamble and Arts. 23.2 and 23.9). The more, there

is now also a lectotype available for actis, while Gerhard’s collec-

tion (including the type specimens of atys) was probably entirely

destroyed (Olivier, 1999: 129-130).

106

Correct attribution of the name. The type specimens of P. (A.)

actis are rather robust and they leave no doubt as to their specific

distinctness from the taxon referred to under this name by

Hesselbarth ef al. (1995), thus confirming what was already appar-

ent from an attentive study of the original description and illustra-

tion by Herrich-Schäffer, as well as from that of L. atys Gerhard,

[1851] and of L. athis Freyer, [1851]. It appears that the taxon

denoted as “Polyommatus (Agrodiaetus) actis (Herrich-Schäffer,

[1851])” by Hesselbarth er al. (1995) and subsequent authors, or at

least its Taurus populations, was unnamed yet. Freyer’s athis

indeed applies to a blue butterfly “usually looking like alexis [P.

icarus] on the upperside’, while the butterfly with black suffusion

in the distal part of the hindwings is considered as “a very nice

form” but not named. The application of the name athis by several

subsequent authors to denote this form is thus probably unjustified,

although it must be said that specimens of the smaller and darker

taxon without any black suffusion exist as well. As no extant types

of athis are known however, and in the interest of nomenclatural

stability, it is best to follow current use in considering it as a sub-

jective synonym of actis. The name /unulata, attributed to material

from the Sultan Daglari, applies to an aberration and is thus

unavailable. Carbonell’s (1997) name artvinensis applies to popu-

lations from the provinces of Erzurum and Artvin in northeastern

Turkey, that look quite different from populations of “actis sensu

Hesselbarth ef al. (1995) et sqq.” and that have a different haploid

chromosome number.

Current problems and provisional arrangement. True actis, the

taxon firdussii [pro parte!] of Hesselbarth er al. (1995), most prob-

ably consists of more than one species, but the picture is not quite

clear. The intensity of the blue colour (even leaving sertavulensis

and ernesti aside) varies from vivid violet blue to a lighter blue

(especially in firdussii from the Elburs), and the variation in colour

does not always coincide with that in chromosome numbers (de

Lesse, 1960: 186-197, 1962: 66-70). The following haploid chro-

mosome numbers (CN), determined in metaphase I (I) and in

metaphase II (II), have been found in populations ascribed to actis,

firdussii or pseudactis (after de Lesse, 1960, 1962; Hesselbarth et

al., 1995; Lukhtanov ef al., 1998; Lukhtanov, unpublished):

107

Locality

E Bürücek

(Icel, Tekir, TR)

Number and

stage of cells

examined

a(t); (ED)

Name used in original

publication

A. actis ssp.

27-28

Ivrız, Pozantı

(Konya &

Adana, TR)

| firdussit

Aladasßları,

15 km SE

Camardı

(Nigde, TR)

25 km N

Erzincan (TR)

1 (1)

2().1(LID

Literary source

de Lesse (1962)

Hesselbarth ef al.

(1995)

Lukhtanov

(unpubl.)

A. actis pseudactis

Forst.

de Lesse (1962)

Kopdagi Gecidi

(Erzurum/

Bayburt, TR)

1 (D)

SW)

AE)

À. actis pseudactis

Forst.

de Lesse (1962)

(27-29)

Mirgemir Dagh

[Tahir Gecidi,

Asrı, TR) 1 dD

1; 1.

Kuzgunkiran

Gecidi (Bitlis,

TR)

1 (L ID

82 (EE)

A. actis pseudactis

Forst.

firdussii

de Lesse (1962)

Hesselbarth et al.

(1995: 705, Tab.

20)

25-26

25+m

(Forster, 1956)

8 (1)

3 (1)

P. (A.) firdussii

(1995:733)

P. (A.) firdussii

pseudactis (Forster,

1960)

(25-26)

In Catak

(Van, TR)

32 km NNE

Catak (Van, TR)

Güzeldere Gecidi

(Van, TR)

Pass E. Tabrız

(Azarbayjan-e

Sharshı, Iran)

Damavand

(Tehran, Iran)

“Abe Ali

(Elbourz)” (Iran)

firdussii

P. (A.) firdussii

pseudactis (Forster,

1960)

P. (A.) firdussii

(Forster, 1956)

P. (A.) firdussii

pseudactis (Forster,

| 1960)

À. actis SSP.

LO?20 1)

1@: dm

WESD;

(33-34: II)

A. actis Firdussi Forst.

1 (1), 1 (D)

A. actis Firdussi Forst.

Hesselbarth ef al.

Lukhtanov et al.

(1998)

Hesselbarth er al.

(1995)

Lukhtanov et al.

(1998)

Hesselbarth er al.

(1995: 733)

Lukhtanov et al.

(1998)

de Lesse (1962)

It thus appears that not only variability in the number of biva-

lents is encountered, but also the presence or absence of a small

extra element (m), while variability in specimens from different

108

localities (often at close range, compare e.g. material from Van and

Catak, both in the province of Van in SE Turkey) is significant

(Lukhtanov er al., 1998: 3, 6). Matters are further complicated by

notable differences in karyotype, even between allopatric popula-

tions with a similar chromosome number (e.g. Bürücek, Mirgemir

Dagh [Tahir Geçidi] and Tabriz), suggesting a possibly different

organisation of the hereditary material with a resulting degree of

intersterility (de Lesse, 1962: 70).

In order to ever be able to ascribe any population unambiguous-

ly to any given taxon, it is necessary to study the karyotype and

chromosome number of topotypical actis, firdussii, pseudactis, ser-

tavulensis and ernesti (there are no data at hand for the two last-

named nominal taxa), and to combine these data to those obtained

after a study of the phenotype and karyotype of any given popula-

tion. Cross-breeding studies between several populations would

further deepen our insight into this interesting case.

For the time being, the following arrangement seems to offer a

fair provisional solution (only the holotype or lectotype of each

nominal taxon is listed):

Polyommatus (Agrodiaetus) actis (Herrich-Schäffer, 1851)

“[Lycaena] Actis Kad.” Herrich-Schäffer, G. A. W., [31.XII.1851]. Systematische Bear-

beitung der Schmetterlinge von Europa, zugleich als Text, Revision und Supplement

zu Jakob Hübner’s Sammlung europäischer Schmetterlinge, Vol. 6(48), 24; “[Lycaena]

Actis”, Vol. 1(49), Tab. 104, [Figs] 496-499 [31.XII.1851]; “Lycaena Actis”, Vol.

6(55), Systema Lepidopterorum Europae. Systematisches Verzeichniss der Euro-

päischen Schmetterlinge 4 [31.X1.1852]; “actis (Lycaena) Kad. HS. 466-469. athis

Fr”, Vol. 1(65), Index 1 [31.XII.1854]; “Actis dazu Atys Gerh.’ Vol. 6(69), 167

[31.XIL.1856]; “actis Lycaena” Vol. 6(69), Index Universalis Macrolepidoptera 1

[31.XIL.1856]. Type locality: Turkey, Tokat province, env. Tokat (corrected here).

Type material: lectotype d, Turkey, “Amasia” (recte Tokat), VII.1849, leg. Albert

Kindermann, in coll. Museum fiir Naturkunde der Humboldt-Universitat zu Berlin (see

above).

“[Lycaena] Atys. Kind. Actis. Herr.-Schaeff.” Gerhard, [P.] B., [31.V.1851]. Versuch einer

Monographie der europäischen Schmetterlingsarten: Thecla, Polyomattus [sic],

Lycaena, Nemeobius. Als Beitrag zur Schmetterlingskunde, (5), 11, Platte 19, 3 a,b à,

c,d 2. Type locality: Turkey, Tokat province, env. Tokat (corrected here). Type

material: presumably lost. Subjective synonym of [Lycaena] actis Herrich-Schäffer,

1851 (Herrich-Schäffer, [1856], 6(69): 6).

“[Lycaena] Atys Kind. 1. 1’ Heydenreich, [G. H.], [31.VI1.]1851. Lepidopterorum

Europæorum Catalogus methodicus. Systematisches Verzeichniss der europaeischen

Schmetterlinge, 3. Aufl., 13. — nomen nudum.

109

“Lycaena Pap.lilio] Athis.” Freyer, C. F., [31.XIL1 1851]. Neuere Beiträge zur

Schmetterlingskunde mit Abbildungen nach der Natur, Vol. 6(96), 147. Type local-

ity: Turkey, Tokat province, env. Tokat (corrected here). Type material: not found.

Subjective synonym of [Lycaena] actis Herrich-Schäffer, 1851 (Herrich-Schäffer,

[1854], 1(65): Index 1).

Polyommatus (Agrodiaetus) firdussii (Forster, 1956)

“A.[grodiaetus] damone firdussii ssp. nov.” Forster, W., 1956. Bausteine zur Kenntnis der

Gattung Agrodiaetus Scudd. (Lep. Lycaen.) I. — Z.wien.ent.Ges. 41: 58-59, Taf. 8-9,

Figs 16, 17. Type locality: “Nordiran, Schahkuh, Westabhang, Geröllzone,

1800-2000 m”. Type material: holotype d, Iran, Mazandaran province, Shahkuh,

leg. Exp. Wernicke, in coll. Zoologische Staatssammlung Miinchen.

Polyommatus (Agrodiaetus) pseudactis (Forster, 1960), species

incertae sedis

“A.[grodiaetus] actis pseudactis ssp. nov.” Forster, W., 1960. Bausteine zur Kenntnis der

Gattung Agrodiaetus Scudd. (Lep. Lycaen.) II. Z.wien.ent.Ges. 45: 107-109,

Taf. 10-11, Figs 4, 5. Type locality: “Armenia, mts. Daralagez, pag. Martiros, ca.

2000 m alt”. Type material: holotype 6, Armenia, Daralagez Mts., Martiros, leg.

Leo Sheljuzhko & Nina Pavlitzkaja, in coll. Zoologische Staatssammlung München.

“Polyommatus actis H.-S. ssp. pseudactis Forst.” Wiltshire, E. P., 1957. The Lepidoptera

of Iraq, p. 30. — nomen nudum.

Polyommatus (Agrodiaetus) sertavulensis (Koçak, 1979), species

incertae sedis

“Agrodiaetus (s. str.) pseudactis sertavulensis n. subsp.” Kocak, A. O., 1979. Studies on

the family Lycaenidae II. New taxa and records from Turkey. Lycaeniden-Studie II.

Neue Taxa und Meldungen aus der Tiirkei (Lep. Lycaenidae). — Atalanta (Wiirzburg)

10(4a): 323-324, Fig. 31. Type locality: “South Turkey, Konya Prov. Sertavul Paß

1600 m”. Type material: holotype d, Turkey, Karaman province, Sertavul Gecidi,

leg. Ahmet Omer Kocak, in Department of Zoology, University of Ankara.

Polyommatus (Agrodiaetus) ernesti Eckweiler, 1989, species incer-

tae sedis

“Polyommatus (Agrodiaetus) firdussii ernesti subspec. nov.” Eckweiler, W., 1989. Drei

neue Lycaeniden aus der Türkei (Lepidoptera, Lycaenidae). — Nachrent.Ver Apollo,

N. F. 10(2): 97-101, Abb. 35-40. Type locality: “Türkei, Prov. Antalya, Elmalı,

Kohu Dasßları, Dokuz Gölü, 1700-2000 m”. Type material: holotype d, Turkey,

Antalya province, Elmali, Kohu Daßları, Dokuz Gölü, leg. et coll. Wolfgang

Eckweiler.

110

Polyommatus (Agrodiaetus) artvinensis (Carbonell, 1997)

“Agrodiaetus actis artvinensis n. ssp.” Carbonell, F., 1997. Contribution à la connaissance

du genre Agrodiaetus Hübner (1822), Agrodiaetus actis artvinensis n. ssp. en Turquie

orientale, (Lepidoptera: Lycaenidae). — Linn.belg. 16(4): 140-141, pl. coul., figs 1-3.

Type locality: “gorge après Tortum, 1400 m, prov. Erzurum”. Type material:

holotype ¢, Turkey, Erzurum province, gorge near Tortum, leg. J. Salmon, in coll.

Muséum National d’ Histoire Naturelle, Paris.

Polyommatus (Agrodiaetus) sigberti Olivier, van der Poorten,

Puplesiene & De Prins, 2000

“Polyommatus (Agrodiaetus) sigberti sp. nov.” Olivier, A., van der Poorten, D., Puple-

siene, J. & De Prins, W., 2000. Polyommatus (Agrodiaetus) artvinensis stat. nov. and

P. (A.) sigberti sp. nov., two vicariant species known so far only from Turkey (Lepi-

doptera: Lycaenidae). — Phegea 28(2): 61-63, 67-73, col. pls 1 & 2, figs 17-32. Type

locality: “Turkey, Kayseri province, Aladaglari E. side, 34 km S. Yahyali,

2800-2900 m, St. 2384”. Type material: holotype d, Turkey, Kayseri province,

Aladaÿlari E. side, 34 km S. Yahyali, 2800-2900 m, St. 2384, leg. W. De Prins, A.

Olivier & D. van der Poorten, in coll. Instituut voor Systematiek en Populatiebiologie,

Zoölogisch Museum Amsterdam.

“Lycaena athis nov. ab. lunulata Schw.” Schwingenschuss, L., 1935. [Versammlungen der

Sektion für Lepidopterologie. 5. April 1935. Lepidopteren-Ausbeute aus Bithynien

und Anatolien]. — Verh.zool.-bot.Ges.Wien 85: 132. — unavailable name.

“Lycaena actis H. S. ssp. athis Frr. lunulata” Schwingenschuss, L., 1938. Sechster Beitrag

zur Lepidopterenfauna Inner-Anatoliens. — Ent.Rdsch. 55: 146. — unavailable

name.

Acknowledgements

It is my pleasure to thank the following persons for their help:

Dr. Wolfram Mey (Museum für Naturkunde der Humboldt-Univer-

sıtät zu Berlin) for his kind invitation to study a.o. the relevant

Polyommatus (Agrodiaetus) material in the collection under his

care and also for the colour photographs of the types of P. (A.) actis,

reproduced on Figs 4—9 in the present study; Dr. Yuri P. Nekrutenko

(Schmalhausen Institute of Zoology, Kiev) for bringing both actis

specimens to my notice during our common stay in Berlin in

November 1998; Dr. Vladimir Lukhtanov (Dept. of Entomology,

Faculty of Biology, University of St.-Petersburg) and Mr. Harry van

Oorschot (Instituut voor Systematiek en Populatiebiologie,

Zoodlogisch Museum Amsterdam) for making available the so far

unpublished karyological data presented in the table in the present

111

study; Mr. Hans Henderickx (Mol, Belgium) for the colour pho-

tographs reproduced here on Figs 1-3; Mr. W. O. De Prins

(Antwerpen) for allowing me, as always, to make extensive use of

his library; Mr. Dirk van der Poorten (Antwerpen) and two anony-

mous referees for their constructive comments on the manuscript;

Prof. Dr. Konrad Fiedler (Lehrstuhl Tierökologie, Universität

Bayreuth) for the German summary.

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S. 1-164, Index 1-24, Taf. Papilionides 1-134, Taf. Hesperides 1-7

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1-23, a-h, k, S. 1-184, Index [1]-34, Taf. Geometrides 1-91 (1843-1856);

4(Hefte 27-30, 33-34, 39-40, 44, 46, 48, 53, 55, 57, 60, 63, 68): Sign. 1-36,

a-m, S. 1-288, Index [1]-48, Taf. Pyralides 1-23, Taf. Tortricides 1-59

(1847-1855); 5(Hefte 22, 30, 34, 39, 42, 44, 46, 48, 53, 55, 57-68): Sign.

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114

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Nota lepid. 23 (2): 119-140; 01.VII.2000 ISSN 0342-7536

Comparative data on the adult biology, ecology

and behaviour of species belonging to the

senera Hipparchia, Chazara and Kanetisa

in central Spain (Nymphalidae: Satyrinae)

Enrique GARCIA-BARROS

Department of Biology (Zoology), Universidad Autönoma de Madrid, E-28049

Madrid, Spain. e-mail: garcia.barros@uam.es

Summary. The potential longevity, fecundity, mating frequencies, behaviour, and sea-

sonal reproductive biology were studied in several satyrine butterflies belonging to the

genera Hipparchia, Chazara and Kanetisa, in an area located in central Spain. All the

species studied appear to be potentially long-lived, and a relatively long period of pre-

Oviposition is shown to occur in C. briseis and K. circe. Potential fecundity varies between

250 and 800 eggs depending on the species (with maxima exceeding 1300 eggs in

K. circe). The results are discussed in terms of the possible ecological relationships

between adult ecological traits and the species abundance, and the possibility of a marked

geographic variation between species, that might be of interest in relation to specific

management and conservation.

Zusammenfassung. Fiir mehrere Vertreter der Gattungen Hipparchia, Chazara und

Kanetisa (Satyrinae) wurden in einem Gebiet in Zentralspanien potentielle Lebensdauer,

potentielle Fekundität, Paarungshäufigkeiten im Freiland und saisonaler Verlauf der

Reproduktionstätigkeit untersucht. Alle untersuchten Arten sind potentiell langlebig, eine

relativ lange Präovipositionsperiode tritt bei C. briseis und K. circe auf. Die potentielle

Fekundität variiert je nach Art zwischen 250 und 800 Eiern (mit einem Maximum von

über 1300 Eiern bei K. circe). Die Befunde werden diskutiert im Hinblick auf mögliche

Zusammenhänge zwischen ökologischen Charakteristika der Imagines und der Abundanz

der einzelnen Arten, sowie unter dem Gesichtspunkt, daß ausgeprägte geographische

Variabilität in den ökologischen Eigenschaften auch für spezifische Management- und

Naturschutzplanungen bedeutsam sein kann.

Résumé. Le potentiel de longévité, la fécondité, la fréquence d’accouplement, le com-

portement et la reproduction saisonnière ont été étudiés chez plusieurs espèces appar-

tenant aux genres Hipparchia, Chazara et Kanetisa (Satyrinae), dans une région localisée

au centre de l'Espagne. Toutes les espèces étudiées paraissent avoir potentiellement une

longue vie, C. briseis et K. circe montrant une période de préoviposition relativement

longue. La fécondité potentielle varie entre 250 et 800 œufs selon l’espèce (avec un maxi-

mum de plus de 1300 œufs pour K. circe). Les résultats sont discutés en termes de rela-

tons écologiques possibles entre les exigences écologiques des adultes et l’abondance des

espèces, et la possibilité de variations géographiques marquées entre espèces, ce qui peut

être intéressant à relier à des mesures spécifiques d'aménagement et de protection.

Key words: Lepidoptera, Nymphalidae, Satyrinae, fecundity, life history, longevity,

mating frequency, seasonality, Spain.

Introduction

Some adult life history traits such as potential longevity and

fecundity may have a relevant bearing on other ecological features

of practical interest, like the ability for dispersal, the capability for

colonisation, or the potential for population growth (e.g. Young,

1972; Spitzer ef al., 1984; Bink & Siepel, 1986). In spite of the fact

that the members of the superfamily Papilionoidea have been thor-

oughly studied in the Western Palaearctic, detailed data on their

reproductive biology are largely confined to a relatively small num-

ber of target species.

This work intends to present some basic features of the adult

reproductive biology of Hipparchia alcyone ([Denis & Schiffer-

miller], 1775), A. statilinus (Hufnagel, 1766), A. fidia (Linnaeus,

1767), Kanetisa circe (Fabricius, 1775) and Chazara briseis

(Linnaeus, 1764), with comparative data on H. semele (Linnaeus,

1758) in central Spain. The main adult life history traits involved

are potential longevity, female potential fecundity, temporal distri-

bution of oviposition in the wild, mating frequencies as estimated

by spermatophore counts, and a general description of the activity

of the insects in the field. The results are largely based on data col-

lected more than ten years ago for a comparative study on Spanish

Hipparchia (Garcia-Barros, 1988a), to which new field and rearing

data were added.

All the species involved are strictly univoltine across their range,

and hibernate as small larvae (usually Ist or 2nd instar larvae,

depending on species and locality — Bink, 1992a and references

below). Life history data from H. statilinus and H. fidia in central

Iberia can be found elsewhere (Garcia-Barros, 1989a, 1989b; see

also Jutzeler, 1993; Jutzeler et al., 1998). Available information on

local larval host plants of the species involved is summarised by

Munguira et al. (1997). The female reproductive biology of

H. semele in the study area has been dealt with in detail elsewhere

(Garcia-Barros, 1988b, 1989a, 1992), and thus only some un-

120

published details on the behaviour of this species are included here.

This grayling is characteristic with a long flight period much alike

that of K. circe and C. briseis described below; the females under-

go delayed ovarian maturation throughout the driest part of the

Summer, and are plugged during copulation so that female

monogamy is obligatory. The adults of K. circe and C. briseis occur

all along the summer in the study area, from the beginning of July

until September (early males can be seen in June, or even late May

at especially warm locations). H. alcyone has a shorter flight peri-

od (July to late August or early September). The adults of H. fidia

and H. statilinus are the latest to emerge (mid July, with fresh

females still appearing | to 15 August) and remain on the wing

until mid or late September. Except for H. alcyone, some adults

may Stay alive as late as early October when early autumn is sunny

and dry.

Methods

Data from captive insects were derived from small numbers of

adults obtained from larvae, either reared in the laboratory or

collected in the study area described below. Once mated, the

females were kept in individual voliaries and the eggs laid were

counted daily (as described in Garcia-Barros, 1988b). Potential

fecundity was calculated as the sum of all the eggs laid by one

female (realised fecundity) plus all chorionated eggs contained in

the abdomen of a dead insect, counted after dissection. The males

were maintained in small groups (4—10 individuals) in larger cages

(1 m’). The rearing cages were placed outdoors in a grassy outer

yard at the Universidad Autonoma (Madrid) in a sunny exposure, in

a way such that a shadowy shelter was available within the voliary

allowing the insects to avoid overheating. Environmental condi-

tions were not controlled or measured. Weather was usually fine

to hot during the season when adults were available (July and

August), so that oviposition normally proceeded in a more or less

continuous manner during a female’s life period.

Field work was carried out at 15 sites within a 40 x 30 km area

surrounding the artificial lake of Entrepeñas, in the province of

Guadalajara (central Spain), at elevations between 700 and 900 m.

Preliminary field work was directed to obtain estimates of the

121

species’ abundance and site specificity. First, the study area was

divided in 48 squares of 5 x 5 km. An area with reasonably well

preserved vegetation (usually consisting of succession stages of

Quercus ballota or ©. faginea woodlands) was selected within each

square, and assessed four times (once per 15 days during July and

August 1984) for the presence of each species. Second, 15 sites that

proved particularly suitable for at least one of the species studied,

were selected for regular sampling. Transect surveys (500 x 5 m)

were carrıed out at these locations every ten days during two sea-

sons (June to October, 1985-1986), using a procedure basically

sımilar to that popularised by Pollard er al. (1975), but with explic-

it reference to a known area extension. The data from the main

flight period (July and August) were then used to calculate the aver-

age number of butterflies per area unit.

Further field observations were conducted in order (1) to deter-

mine the reproductive condition of wild females, (2) to estimate the

change in wing wear condition along a season, and (3) to quantify

the patterns of activity displayed by the two sexes of each species.

Field observations on C. briseis were restricted to step 1 since,

although spread across the study area, this hermit was compara-

tively scarce in most of the sites prospected (see Table 1).

Table 1. Species abundance in the study area, as derived from counts of adult

insects and presence/absence data: Mean number of adults per hectare (mean and

standard deviation in 15 sites), percentage of sites where the species was col-

lected (15 sites — 100%), and percentage of 5 km squares (48 squares — 100%)

in the study area where each species was observed. For comparative purposes,

data from taxonomically allied species (Satyrus series) that occur in the area are

given. These are, in addition to the species dealt with in this study: Hipparchia

semele (L., 1758), Arethusana arethusa ({D. & Schiff.], 1775), and Satyrus

actaea (Esp., 1780).

Species Mean population Area coverage

density

% sites %5x5km

squares

H. alcyone

H. semele

H. fidia

H. statilinus

C. briseis

K. circe

A. arethusa

S. actaea

Se ee ee

WAOCNH CoO

NR BN \0 Un \O 00

SII Wd 0 Lo

—

122

(1) The reproductive status of adult females was estimated from

subsamples of female insects (up to 10 individuals per site and

date) that were transported to the laboratory in a portable fridge

within 6 hours after collection, and then dissected or frozen for fur-

ther dissection. The specimens were checked for the presence of

mature eggs within the common oviduct (indicating that the female

was ovipositing). The diameter of the largest eggs within the

ovaries was measured to the nearest 0.03 mm. The total number of

eggs and number of spermatophores within the mating pouch (bur-

sa copulatrix) were counted (e.g. Burns, 1968). These results are

based on samples taken during two subsequent years. Care was

taken to obtain samples of comparable size at least once every ten

days each year, to avoid misinterpretations caused by between-year

variation.

(2) Wing wear state. The transect surveys described above were

repeated over the full flight season (June to October). In each tran-

sect, all the butterflies were collected to assess their wing state

according to the scale 1—4 (where 1 — young to 4 — old).

(3) Adult activity. The activity displayed by the adults was

recorded along the transect surveys (for obvious reasons this had to

be done before assessment of wing states). Apparently resting indi-

viduals that were suspected to have been disturbed by the

researcher were observed for one to five minutes, and then classi-

fied as resting unless other activity was displayed in that period.

This proved necessary to prevent inflated records of “resting” indi-

viduals that might actually represent cryptic behaviour as a

response to the observer’s activities. All surveys were conducted

between 11.00 and 17.00 hours, in sunny weather and under ambi-

ent temperature above 25°C.

Results

Butterfly counts and species frequencies. The estimates obtained

are shown in Table 1. The number of adults counted per hectare

reached 50, but varied widely depending on the species and

location. On average, none of the species studied could be defined

as very abundant, with average estimated densities reaching

30 adults/hectare at most.

123

Number of eggs laid

a oO

10 20 30 Days

Fig. 1. Average patterns of daily egg production by captive females of (upper to lower):

H. alcyone, H. fidia, H. statilinus, C. briseis and K. circe. Thick line in each graph indi-

cates daily means, and thin lines the value of standard deviation above and below the

mean. Sample sizes (number of females alive on date d) are given by the bars in the lower

part of the plots (scale in lower right angle). In all instances, date 1 is the first day of

Oviposition.

124

100

Sayed) paeyew Jo abeyussulad

Egg diameter (mm)

H. statilinus

0.1

Jul. Aug. Sept. Oct.

Fig. 2. Reproductive status of wild females of H. alcyone (upper), H. fidia (centre) and A.

statilinus (lower). Dots represent the diameter of the widest eggs contained in the

abdomen (in mm, left scale in all plots). The crosses indicate the percentage of mated

females plotted on 15-days intervals (right scale; the continuous line is simply a smoothed

line connecting the successive percentages). The number of individuals dissected per date

is not proportional to the actual abundance of the species on that date.

125

Beiuss1ad

Egg diameter (mm)

söjeuls, psjeui Jo ©

K. circe

Jul. Aug. Sept. Oct.

Fig. 3. Reproductive status of wild females of C. briseis (above) and K. circe (below).

Details as in Fig. 2.

Longevity and reproductive performance. The results concerning

laboratory observations, as well as those from dissection of females

from the wild, are summarised in Table 2. The temporal patterns of

oviposition of captive females are shown in Fig. 1, and the re-

productive status of wild females is represented ın Figs 2 and 3. It

was evident that the females of all the species required no less than

four days to reach ovarian maturation. The dissection of small

series of adult females of H. alcyone and A. fidia aged 1 to 10 days

126

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different age: A — 24 hours after emergence, and B — a 10 days old female on its first day

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128

confirmed that the absence of oviposition during early adult life

was due to the fact that females were reproductively immature

upon emergence (Fig. 4), and was not an artefact caused by rearing

procedures.

Wing state distribution and seasonal variation. Since wing state

was correlated to observation date (as expected, see Fig. 5), an

analysis of the variance with observation date (date 1 — June 1) as

the covariate was used to compare male and female average wing

wear controlling for the effect of capture date. The effect of the

covariate (date) was always significant (F > 594 in all instances,

P < 0.0001, 1.d.f.). Between-sex differences were significant for

the four species tested (in all instances P < 0.001, 1.d.f.): A. alcy-

one(F = 857.8), H. fidia (F = 63.33), H. statilinus (F = 902.4),

and K. circe (F = 70.0). In summary, the degree of wing wear dif-

fers between sexes within each species, ıf dates are kept constant.

In order to detect between-sex differences ın the seasonal pro-

gression of wing status, the slopes of the regression lines of male

and female individuals within each species were subsequently

compared. The interaction between the varıables “sex” and “date”

was examined in an analysis of the variance of “wing state”, where

“sex” was set as the classification factor and “date” as a covariate

(GLM procedure). Evidence for heterogeneity of the slopes arose

ion waninus (E = 9.39: P = 0.002, df : = 1) and K. circe (F

= 0.037,.d.1. = 1), but not for A. alcyone (F = 0.23;

Be = Non: fidia (Fk = 1:56; P. = 0.213, df::=

1) (see Fig. 5).

Fecundity, species abundance, and site specificity. The estimates

of fecundity obtained in the laboratory were compared with the

data in Table 1 for Hipparchia spp., C. briseis and K. circe.

Fecundity was not correlated with the species mean densities,

although it was related to the dispersion across the study area (mea-

sured as the number of study sites where a species was recorded,

see Table | (Fig. 6). No other obvious relationships were obtained

(results not given in detail). Such comparisons are probably mean-

ingless because of the small number of species included. In any

case, it is interesting to note that there is some degree of corre-

spondence between range size and potential fecundity across the

study sites.

129

H. alcyone

Kveince

Wing status

VI NT A ER X VI VIB. SVT ZERZ X

Date (months)

Fig. 5. Relationship between wing state and date in males (solid line, filled dots) and

females (dashed lines, circles) of A. alcyone, H. fidia, H. statilinus and K. circe. The dots

represent average values for each 15-days period (vertical bars — 2 x s.d.). Regression

lines for males and females in each species are (WS — wing state, D — date where | — June

1): A. alcyone — WShaies = —8.52 + 0.049(D), WSremates = —9.43 + 0.050(D); H. fidia —

WS nates = —10.0 + 0.053(D), WSvemates = —9.37 + 0.048(D); HA. statilinus — WS aa = —8.90

+ 0.048(D), WStemates = —7.12 + 0.039(D); K. circe — WSwaes = —7.25 + 0.041(D), WStemates

= —6.24 + 0.03(D). Correlations (r) are above 0.76 and significant (P < 0.0001) in all

instances. Sample sizes are those given in Table 3.

Activity and behavioural notes. The summary of adult activities

provides a general description that may well fit to intuitive descrip-

tions of the behaviour of the species involved. The males of

Chazara and Hipparchia spp. displayed typical perching behavi-

our, often fighting for territories. The males of K. circe showed a

mixed strategy, where patrolling for females (often close to

emergence sites in grassy areas) was combined with perching

activity. Overall, the group of species under study can be described

130

as conservative in relation to flight activity. As a general rule, less

than 20% of the observations concern what could be described as

erratic flisht, while a large proportion of records consisted of

apparently inactive, or thermoregulating butterflies. Although

males and females show sharply different behaviour patterns, the

sexes appear not to be very far apart when behaviour ıs classified

into four main categories (as in Table 3). In short, males and

females devote broadly similar periods of time to activities related

to reproduction, feeding, and displacement. However, these activi-

ty schedules involve a significantly higher investment in flight for

the male sex, as shown in Table 4. Finally, shifts in activity patterns

are obvious for the female sex in all species, while male activity

appears to be more constant over the season (K. circe males may be

an exception: see Table 5).

Incidentally, some differences between the kind of substrates

used as perching sites by the males of Hipparchia spp. was detect-

ed (Table 6). The males of A. alcyone and H. semele perched more

often on tree trunks or low branches, while those of A. fidia and A.

statilinus preferred rocks, stones or decaying wood pieces upon the

ground surface. This dichotomy is to some extent parallel to that ın

the adult wing colour patterns (upperside brown with light band, or

uniformly greyish). Although interesting, this may simply reflect

the preference of the former two species for woody places in the

study area.

As noted for A. statilinus, H. fidia and H. semele (Garcia-Barros,

1989a, 1989b and references therein), the eggs of C. briseis and A.

alcyone were laid singly, most frequently on dry grass stems or

other decaying plant material on the ground and more rarely

attached to the base of grass tufts. K. circe females drop their eggs

while settling close to the ground (e.g. Geiger, 1987, and personal

observations). This was frequently observed during September.

Ovipositing females concentrated at wood margins, road verges,

and especially at relatively moist sites around small water courses.

The females of this satyrine were observed all along the summer,

but they were scarcer and more concentrated in wooded areas dur-

ing the driest period (mostly August). The adults of this species

have a complex biology that probably implies relatively important

movements between different areas. Captive females of K. circe

131

have been reported to stridulate, probably as a part of their behav-

iour to reject courting males (Garcia-Barros, 1986), but no further

data on this behaviour were obtained in the field.

200 300 400 500 600 700

r=0.30 (P=0.57)

Mean population density (adults/ha)

Fecundity (mean number of eggs)

r=0.98 (P<0.001)

Percentage of study sites (x 100)

200 300 400 500 600 700

Fig. 6. Interspecific relations between average fecundity and the relative abundance and

range size of the species in the study area. Data from Table 1 (data on egg production by

H. semele from Garcia-Barros, 1988b, 1992).

182

N © &

Mean daily egg production

r=0.87 (P=0.03)

Average preoviposition time (days)

800

600

400

Potential fecundity

r=0.52 (P=0.28)

0) 10 20 30 40

Fig. 7. Interspecific relationships between the average time required to lay the first egg

(preoviposition), and both potential fecundity (number of eggs) and mean daily egg pro-

duction in laboratory reared females. Based on the data from the species included in the

study plus Hipparchia semele (from Garcia-Barros, 1988b, 1992).

Table 3. Frequency distribution of main activity types in males (m) and females

(f) within each species, expressed as a percentage of the total number of records

(n — sample size). B — resting or basking, R — reproduction and related activities

(including courtship, territorial defence, mating and oviposition), F — flying

(erratic flight), N — feeding. The differences tested are between sexes within each

species (x°, 3 d.f.).

Species

H. alcyone

H. semele

H. fidia

H. statilinus

K. circe

ee =

Table 4. Comparison between the proportion of males (m) and females (f) in

flight (including territorial fights between males) or settled on the ground or

other substrates (either active, basking or resting), within each species (n — sam-

ple size). The value and statistic significance of Chi-square tests between sexes

of each species is given (X, 1 d.f.). Sample sizes as in Table 3.

Species

Activity classes

Flying Settled

H. alcyone

H. semele

H. fidia

HA. statilinus

K. circe

134

39.41 60.59

15.89 84.11

38.98 61.02

1832 81.68

40.46 59.54

18.89 81.11

46.89 Ss

15415 84.85

64.76 35.24

TS 22/2p)

Table 5. Evidences for seasonal shifts in activity types along the season for adults

of Hipparchia spp. and K. circe. The variable “date” was tested by means of one-

way analyses of variance using “activity type” as classification factor (indepen-

dent ANOVAs for each sex and species). The value of the F statistic is given

together with the level of significance (3 d.f.; *** — P < 0.001, ** — P < 0.01, *

— P<0.05, n.s. — P > 0.05). Sample sizes as in Table 3.

Species

H. alcyone

H. semele

H. fidia

H. statilinus

K. circe

J'EN DB Er MERE

Table 6. Comparison between the substrates used by perching males of the four

species of Hipparchia (proportion of males on tree trunks or branches, vs. stones

or other substrates at ground level): x’ value and level of significance (1 d.f.: ***

— P < 0.001, ** — P< 0.01, n.s. — P > 0.10). Sample sizes as in Table 3.

H. semele H. fidia H. statilinus

H alcyone 1.016 67.12*** 112.603***

H. semele 3750 — 10.496**

H. fidia 3.093 »*

Discussion

Adult butterflies are often regarded as ephemeral organisms,

with a short life and relatively fast reproduction. This is by no

means a general pattern. The temporal distribution of female repro-

ductive effort of butterflies may in fact be ordered along a continu-

um (Boggs, 1986; Tammaru & Haukioja, 1996). Female Hippar-

chia, Chazara and Kanetisa have a potentially long adult life span

(in terms of an insect), and at least a few days are required before

the eggs mature and are ready to be laid. Ovarian maturation is

delayed for no less than two weeks in K. circe and C. briseis (as in

H. semele: Garcia-Barros, 1988b). Whether or not long delayed

Ovarian maturation is a geographically variable trait within species

135

is central to the interpretation of the observed patterns in terms of

intraspecific variation (i.e., at a population or regional level, e.g.

Garcia-Barros, 1992 for A. semele), or interspecific variation. For

instance, the flight period of C. briseis in the southern and northern

boundaries of its distribution range 1s apparently uniform (basical-

ly, July to September: Seufert & Grosser, 1996), but no evidence

for any kind of aestivating behaviour has been recorded from non-

Mediterranean populations. According to available evidence,

delayed oviposition in this and closely related species with wide

geographic distribution has to be interpreted in terms of adaptation

to local climate conditions that imply a dry summer period when

adequate food plants (grasses) are not suitable as larval food. The

species considered show remarkable coincidence in their period of

Oviposition in the wild, irrespective of their pattern of adult emer-

gence. It could be argued that species where the females undergo a

long pre-reproductive life would have comparatively less chances

to reach fecundity comparable to those of related species with a

more or less immediate oviposition, unless pre-reproductive female

mortality was compensated for. This may have important conse-

quences related to adult mortality schedules, and it is thus interest-

ing that there is some coincidence between long pre-reproductive

female life and high daily egg production rates among the species

studied here which, however, does not result in a correlation

between long preoviposition and high potential fecundity (Fig. 7).

Female fecundity varies widely across species, and may reach

remarkably high figures in K. circe (up to 1300 eggs). The present

estimates are well above those presented by Bink (1992a) for the

same species (for instance, 140 versus 598 in A. statilinus, 250 ver-

sus 666 in K. circe, 100 vs. 369 in H. alcyone). These differences

could probably be attributed to the more favourable temperatures

experienced by the insects at Madrid (in comparison with The

Netherlands), or to geographic variation in the life history traits

involved (sharp differences in fecundity and female pre-reproduc-

tive life span have been found after comparison of geographically

distant populations of H. semele — Bink, 1992a; Garcia-Barros,

1992: see also Wiklund & Karlsson, 1988). However, there is the

possibility that some of the data on butterfly fecundity by Bink

(1992a) have to be critically reviewed on the light of new evidence.

136

While counts of ripe eggs contained in the abdomen of female

butterflies have occasionally been proposed as estimates of poten-

tial fecundity (e.g. Ehrlich & Ehrlich, 1978), such procedure is

obviously inaccurate in species where egg maturation proceeds

along the female’s life. Most females in this study died without

having laid even all the mature eggs contained in their abdomens.

Such counts might, however, represent reasonable approximations

to daily egg production under favourable conditions (e.g. compare

Table 2 and Fig. 4 for H. alcyone). Follicle counts (as used by

Braby & New, 1988) may produce less conservative estimates, but

still the contents of an individual ovariole can represent a continu-

um between ca. 0.05 mm and a mature egg (up to | mm, depend-

ing on species). The results of dissection of wild females show that,

while female monogamy is the rule, repeated copulation does

sometimes occur (unlike H. semele from southern Europe, where

multiple mating is usually prevented by a sphragis secreted by the

male: Garcia-Barros, 1989a). C. briseis females appear to be strict-

ly monogamous, but this might be due to small sample size.

Male and female activity patterns differ not as much in the time

spent in activities intended to get food resources or reproductive

success, but in the effort required: males invest more energy in

flight-related activities. Interestingly, there 1s evidence for season-

al shifts in activity in the female sex, probably highlighting the

importance of seasonal distribution of female activity. This might

result from the seasonally uneven distribution of adult food

resources, that may be crucial to complete ovarian maturation (cf.

Wheeler, 1996). The results show that the females mate relatively

soon after emergence (alternatively, the catchability of virgin

females is very low). Young mated females probably remain mod-

erately active until the onset of oviposition. The observed pattern

would lead to predict different patterns of wing state change along

the summer in the female sex. The fact that this is corroborated in

only two species (H. statilinus and K. circe) is difficult to explain,

but little else can be said until more accurate methods are applied.

Seasonal variation in male behaviour has been found in other

satyrines (Wickman & Wiklund, 1983; Wickman, 1992; Van Dyck

& Matthysen, 1998), and seasonal changes in behaviour and habi-

tat selection have been reported for satyrines from tropical or

137

subtropical areas where rainfall is strongly seasonal (e.g. Braby,

1995). What is of interest is that West-Palaearctic satyrine

nymphalids are predominantly univoltine (that is, they complete

one generation per year), and that this pattern applies as well to the

Mediterranean species or populations. Hence, seasonal variation in

activity patterns in Hipparchia, Kanetisa or Chazara represents

environmental variation that is faced by the individual adults (not

by the different individuals belonging to successive generations).

The amount of time spent settling on the ground by the adults of

both sexes must be partly attributable to thermoregulating behav-

iour (since overheating must be a serious risk for several hours a

day during the central part of the summer), as well as to the domi-

nating perching strategy used by the males to locate mates.

In summary, that insects such as butterflies have varied and com-

plex life styles is not a surprise. A more detailed knowledge of

some key biological traits may help in our understanding of the

species responses to environmental changes. As suggested by Bink

(1992b), some ecological characters of the adults might prove of

predictive value in relation to changes in abundance. More esti-

mates of such traits are required, however, to get an adequate con-

trast of their reliability and of their possible geographic variation

(which in turn constitutes an evidence of genetic diversity). To the

extent that such variation occurs, the extrapolation of local results

(e.g. for purposes of population management for conservation) may

prove inadequate.

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140

Nota lepid. 23 (2): 141-146; O1.VIL.2000 ISSN 0342-7536

Phantosoma witti gen. et sp. nov., a new autumn

lasiocampid moth from Turkmenistan

(Lasiocampidae)

Vadim V. ZOLOTUHIN* & Igor Yu. Kostsuk**

* Ablukova Str. 13-12, RUS-432005 Uljanovsk, Russian Federation

** Zoological Museum, National Shevchenko University, Vladimirskaja Str. 60,

UA-01033 Kiev, Ukraine

Summary. A new monotypic genus Phantosoma gen. nov. is established for a new

species Phantosoma witti sp. nov. from the Kopet-Dagh Mts., S. Turkmenistan. The sys-

tematic position of the taxon within the family Lasiocampidae is still uncertain because

only a single female is known.

Zusammenfassung. Eine neue monotypische Gattung Phantosoma gen. nov. wird fiir

eine neue Art, Phantosoma witti sp. nov. aus Siid-Turkmenistan (Kopet-Dagh-Gebirge),

errichtet. Die systematische Position des Taxon in der Familie Lasiocampidae ist unklar,

weil nur ein einziges Weibchen bekannt ist.

Résumé. Un nouveau genre monotypique Phantosoma gen. nov. est établi pour la nou-

velle espèce Phantosoma witti sp. nov. des monts du Kopet-Dagh, au sud du

Turkménistan. La position systématique du taxon au sein de la famille des Lasiocampidae

reste incertaine à l’heure actuelle, vu qu’une seule femelle est connue jusqu’à présent.

Key words: Lasiocampidae, Phantosoma gen. nov., witti Sp. nov., Turkmenistan.

A single female of an unknown lasiocampid moth species was

collected in November 1991 in the Kopet-Dagh Mts (south

Turkmenistan) by our Hungarian colleagues, Gabor Ronkay and

Marton Hreblay. To our surprise, examination of this specimen

revealed the necessity to establish a new genus for it.

Phantosoma gen. nov.

Type-species: Phantosoma witti sp. nov.

Derivation: phantasma (Gk. appearance, phantom) + soma (Gk. body); gender: neuter.

Diagnosis (female). Lasiocampid moth of small size. Head

small; eyes small. Antenna bipectinate, flagellar processes long.

141

Labial palps (Fig. 2) short, porrect, 3-segmented. Proboscis

reduced. Front with longitudinal, sclerotized, wrinkled keel.

Thorax and abdomen strong, covered with long scales. Legs long

and slender, without epiphysis (Fig. 3). Abdomen with apical tuft

of modified scales forming a pillow at its tip.

Female with developed wings (Fig. 4). Frenulum absent; costal

margin of the hindwing slightly extended. Forewing with 5 branch-

es of R: R, free; R; and R; on a long stem, both running to the costal

wing margin; R, and R; on a short stem, both running to outer wing

margin; origins of R.+R; and R,+R; stalked; M, free but its origin

common with radial branch; origins of M, and M, in close proxim-

ity. Origins of Cu, and Cu, moved apart, only one A present. In

hindwing, Sc forming an anastomose with Rs, both forming a long

and slender Sc-R cell; the common branch Sc+Rs connected with

M, by a short transversal vein; origins of M, and M; in close prox-

imity; both Cu and two A present. Discal vein slender in both

wings.

Wing pattern almost completely reduced, forewing with post-

median line hardly visible. Scale cover thin.

Female genitalia (Fig. 5). Anterior apophyses vestigial; both

vaginal plates present; antrum and ductus short; bursa copulatrix

bag-shaped, without signum.

Male. Unknown.

So far the genus includes only a single species.

Phantosoma witti sp. nov.

Holotype 2: “USSR, Turkmenia, Kopet-Dagh Mts., 15 km SE of Nochur, 1300-1400 m,

57°09’E, 38°21’N, 13.-14.XI 1991. N° L49, leg. M.Hreblay & G.Ronkay”. Deposited in

Museum Witt (Munich).

Description. Female (Fig. 1) with characteristics as given under

generic diagnosis above. Wingspan 26 mm, forewing length

14.5 mm. Outer margin of the wings rounded. Forewing olive-

green with greenish-yellow spot at R, tip; hindwing yellowish-

grey, darker towards external area; cilia dark grey on both wings.

Body covered with long ash-grey scales sprinkled with less numer-

ous black ones. Anal tuft consisting of dark elongated scales,

widened distally and slightly curved. Fore tibia without apophysis

(Fig. 3).

142

US®", Türkmenia, Kopet-Dagh

Mis., 15 km SE of oom,

1300-1 400 m, 57709°E, 38°21°N,

Fig. 1. Phantosoma witti gen. et sp. nov. holotype © : a — upperside, b — underside.

Female genitalia (Fig. 5). Papillae anales with short setae;

apophyses anteriores forked, 7-7.5 times shorter than apophyses

posteriores. Antevaginal plate membranous but distinct and pre-

sented as a wrinkled lobe; postvaginal plate transverse, strongly

sclerotized, U-shaped. Ostium membranous; antrum and ductus

short, indistinct, membranous: practically, the ostium is opened

immediately into bursa copulatrix; signum absent.

143

Biology. Late autumn species, admittedly hibernating at the egg

stage; the eggs apparently laid in clusters and covered with scales

from the female abdominal tuft. The following information about

the biology and habitat of this species was given by Gäbor Ronkay:

“The new species was found in an uplifted area of the Karayalchi

valley, inside the nature reserve, at an elevation of 1300-1400 m

above sea level. The relief is rather complex, consisting of rocky

slopes, deep gorges and large limestone cliffs. The slopes and the

lower parts of the gorges covered with shrubby deciduous forest

and smaller grassy patches with low Cytisus shrubs.

Two individuals were observed during the day in strong sunshine

flying rapidly over the surface, looking like a small yellowish bul-

let. These specimens were probably males, resembling in flight

Chondrosoma fiduciarium Anker, 1854 (Geometridae) but even

stronger, faster. Due to the surface relief conditions and rapid flight

the moths cannot be overtaken at a run.

The single female specimen was picked up at a frosty dawn sit-

ting on grass under a Cytisus shrub, having the wings folded over

its back like a resting individual of some Chondrostega species.

The collecting period was the very late autumn, possibly the last

aspect of the year. Dasypolia, Agrochola, Pachyagrotis species

were observed during that night appearing around the light and the

portable light traps”.

Figs. 2-3

144

(1Q

Figs 2-5. Phantosoma witti gen. et sp. nov. holotype 2: 2 — labial palp, 3 - foreleg (tar-

sus removed), 4 — venation, 5 — female genitalia (ventral and lateral view).

145

Discussion. The new species is distinct from other lasiocampid

moths by several unique characteristics. The presence of the frontal

keel (a plesiomorphic character) suggests pupation in a stiff, parch-

ment-like, bag-shaped cocoon, typical for the primitive

Lasiocampidae (e.g. Chondrostega, Trichiura, Poecilocampa,

Eriogaster). At the same time, the moth has no specific teeth or

spurs on the fore tibia, suggesting pupation on the ground or on the

host plant. Very long flagellar processes on the female antennae are

absolutely uncommon for the Lasiocampidae. Venation as a whole

is of a primitive type as well as the lack of humeral veins in the

costal hindwing area. Conversely, a large Sc-Rs cell is quite untyp-

ical for a lasiocampid complex with the venation characteristics

pointed above.

When the female genitalia structure is taken into consideration,

it becomes possible to suppose the male genitalia to be of a slight-

ly modified type. Among all known Lasiocampidae no genus has

forked apophyses in the female genitalia as in this new species.

Thus, the systematic position of this interesting genus within the

family Lasiocampidae is not clear. It can be defined more precise-

ly only when males and immature stages will become available for

examination. Now this genus is provisionally placed between the

so-called “gastropachoid” lasiocampid branch and the subfamily

Poecilocampinae.

Acknowledgements

We would like to express our sincere thanks to all persons who

helped us in this study. Gabor Ronkay and Marton Hreblay kindly

gave this specimen for examination and provided us with the infor-

mation about its biology. Dr Dieter Stiining (Zoologisches

Forschungsinstitut und Museum Alexander Koenig, Bonn) made

photographs of the holotype and assisted us in many other ways.

Thomas Witt (Munich) kindly provided an access to his rich col-

lection enabling a comparative analysis; the new species is named

in his honour.

146

Nota lepid. 23 (2): 147-172; O1.VIL.2000 ISSN 0342-7536

On the ethology and ecology of a small and

isolated population of the Dusky Large Blue

Butterfly Glaucopsyche (Maculinea) nausithous

(Lycaenidae)

Manfred Alban PFEIFER', Ulf Robert Anprick’, Wolfgang FREY’ &

Josef SETTELE*

! Bahnhofplatz 5, D-67240 Bobenheim-Roxheim, Germany

? Mannheimer Str. 12, D-67665 Kaiserslautern, Germany

> Universität Kaiserslautern, Fachgebiet Wasserbau und Wasserwirtschaft, D-

67663 Kaiserslautern, Germany

*UFZ - Centre for Environmental Research Leipzig-Halle, Department of

Conservation Biology and Natural Resources, Permoserstraße 15, D-04318

Leipzig, Germany (corresponding author; e-mail: settele@pro.ufz.de)

Summary. During a study of a small, relatively isolated population of Glaucopsyche

(Maculinea) nausithous (Bergsträsser, 1779), nearly every new adult entering the popula-

tion was individually marked in summer 1990. Marked individuals were regularly recap-

tured. The population assessment was nearly complete, as shown by a comparison with

estimates based on the Jolly-Seber-model. The average adult residence time was estimat-

ed at 2.3 days. Individual observations were made of the diurnal distribution of the fol-

lowing activities: resting, flight, nectaring, oviposition, copulation, and [other] move-

ments. Our results show sex and age specific differences. On the day of marking, which

was normally the day of eclosion, all activities were comparatively low. Older individu-

als of both sexes were generally active for the same amount of time per day, but differed

with regard to activity profiles. Males undertook longer flights to search for females with-

in the habitat, while the females flew more often but over much shorter distances, result-

ing in a shorter total flight time. Instead, females stayed on the flower buds of

Sanguisorba officinalis — the only larval host plant and main adult nectar source — for

much longer, and laid eggs. Females copulate directly after hatching and then immedi-

ately start to oviposit.

Zusammenfassung. Eine kleine, isolierte Population von Glaucopsyche (Maculinea)

nausithous (Bergsträsser, 1779) wurde im Sommer 1990 nahezu vollständig erfaßt.

Aufgrund der intensiv angewendeten Fang-Wiederfang-Methode wurden im Habitat Tiere

mit hoher Wahrscheinlichkeit am Tag des Schlüpfens markiert. Immigration konnte weit-

gehend ausgeschlossen werden. Dadurch kennen wir mit großer Sicherheit das Alter der

Tiere, an denen ethologische Messungen durchgeführt wurden. Von Individuen dieser

Population liegen zeitliche Messungen der Aktivitäten Ruhen, Fliegen, Saugen, Eierlegen,

Kopulieren und Bewegen (Sammelgruppe für weitere Aktivitäten) im Tagesgang vor.

147

Unsere Ergebnisse zeigen, wie sich die Aktivitäten geschlechtsspezifisch unterscheiden

und wie sie sich im Verlauf des Falterlebens verändern. Am Tag der Markierung, der in

den allermeisten Fällen mit dem Tag des Schlüpfens zusammenfiel, waren die Aktivitäten

vergleichsweise gering. Beide Geschlechter verbrachten ungefähr gleich viel Zeit aktiv,

allerdings mit deutlich verschiedenen Schwerpunkten. Während die Männchen längere

Suchflüge nach Weibchen im Habitat unternahmen, waren die Flüge der Weibchen zwar

zahlenmäßig häufiger, insgesamt verbrachten sie aber deutlich weniger Zeit fliegend.

Dafür bewegten sich die Weibchen länger auf den Blütenköpfen der Saug- und

Eiablagepflanze Sanguisorba officinalis und legten Eier. Weibchen kopulierten unmittel-

bar nach dem Schlüpfen und begannen direkt danach mit der Eiablage.

Resume. À l’occasion de l’étude, l’été 1990, d’une petite population relativement isolée

de Glaucopsyche (Maculinea) nausithous (Bergsträsser, 1779), presque chaque individu

nouveau entrant dans la population a été marqué individuellement, puis régulièrement

recapturé. L'évaluation de la population était presque complete, comme l’a montré la

comparaison avec le modèle de Jolly-Seber. La moyenne de la durée de séjour des adultes

est de 2.3 jours. L'observation des individus est basée sur la répartition des activités

diurnes: repos, vol, alimentation, copulation, oviposition et autres. Nos résultats montrent

des différences selon le sexe et l’âge. Le jour du marquage, qui est normalement celui de

l’emergence, toutes les activités sont comparativement réduites. Les vieux individus des ©

deux sexes sont généralement actifs pour la même durée chaque jour, mais diffèrent par

leurs profils d’activités. Les mâles passent de longues périodes en vol à la recherche des

femelles dans le milieu, alors que les femelles volent moins souvent et sur des distances

plus réduites, résultant donc en une durée de vol total plus courte pour celles-ci. À la

place, les femelles se reposent sur les inflorescences de Sanguisorba officinalis — la seule

plante-hôte des chenilles et la principale source de nectar — pour des durées plus longues

et déposent leurs œufs. Les femelles copulent immédiatement après l'émergence et com-

mencent la ponte des œufs juste après.

Key words: Lepidoptera, Lycaenidae, Glaucopsyche (Maculinea) nausithous, diurnal

behaviour, age dependent behaviour, etho-chronogramme, population structure, dispersal,

residence time, MRR-study, Rhineland-Palatinate, S. Germany.

1. Introduction

The ecology of the Dusky Large Blue Glaucopsyche (Maculi-

nea) nausithous (Bergsträsser, 1779) has been quite well studied.

This is due mainly to its high level of vulnerability as expressed in

Red Lists (for Germany e.g. Pretscher, 1998) and the annexes of the

Habitat Directive of the EU and in the Bern Convention (Gruttke,

1996; van Helsdingen et al., 1997), and also to its fascinating

Maculinea-type ecology.

The present study aims to improve knowledge on the adult

ethology and population ecology of the species with special respect

148

to small and isolated populations. An example of such a population

was studied in 1990 throughout the flight period. As the individu-

als of G. nausithous fly slowly compared to other butterfly species,

they are easily followed. The species may reach high densities in its

habitats, and is closely associated with its principal nectar source

and larval host plant Sanguisorba officinalis. For a mark-release-

recapture study, it is therefore sufficient to search for patches with

flowering plants.

2. Materials and methods

2.1. The species Glaucopsyche nausithous. G. nausithous is

univoltine. Eggs are exclusively deposited on Sanguisorba offici-

nalis, where the first three larval instars develop. Hereafter the

caterpillar reaches the ground to be adopted by ants of Myrmica

rubra L., the only known host ant of the butterfly. Within the ant

nest the larvae live predaciously and later pupate. On average, 2.5

pupae are found in one nest. This and further recent information on

the ecology of the species are to be found e.g. in SBN (1991),

Elfferich (1998), Thomas ef al. (1998), Thomas & Elmes (1998)

and Wynhoff (1998).

2.2. Study sites. The study was conducted on a ca. 2000 m? fal-

low grassland within the Mooswieser Tal (Mooswieser Valley, site

M hereafter) south of the city of Kaiserslautern, between the set-

tlements Aschbacherhof and Weiherfelderhof at an elevation of 300

m above sea level. The site was characterised by a high density

stand of Sanguisorba officinalis, unlike the meadows located both

to the East and to the West of the site. The northern and southern

limits of the site are dense pine forests of the Pfalzerwald (Palatine

Forest). Thus, the area of suitable habitat was relatively isolated.

The only possible immigration of G. nausithous from its immedi-

ate surroundings was from the margins of meadows lying west of

the study site, where some S. officinalis plants flowered during the

flight period and where some individuals of G. nausithous were

found. No host plant was found to the East of the site. This rela-

tively isolated site was chosen for the study because it seemed pos-

sible to mark a high percentage of all individuals at the day of

emergence with the manpower available and to identify their age

for the behavioural analysis.

149

The site was divided into two parts. The northern sub-area of the

Aschbach (a little creek) was drier with Sanguisorba officinalis

dominating the vegetation, while the wetter sub-area south of the

Aschbach held high densities of Lythrum salicaria L. and sedges,

but only occasional exposed S. officinalis plants.

An additional site Hagelgrund (in the Eselsbachtal, site H here-

after), north of the city of Kaiserslautern, was also studied. This

was used mainly to test our method of using small portable com-

puters (see 2.5). In some cases we later used data obtained from

Hagelgrund to enlarge the basis of our study (see Tab. 1 below).

There the first G. nausithous could be observed on 13 July, one

day before the first male on site M. As the next observations at site

M have only been made from 17 July onwards, we made some

more studies at site H until that day.

2.3. Marking. Adult butterflies were caught at rest with our fin-

gers. A waterproof pen was then used to write numbers on the

underside of both hindwings so that the individual could be easily

recognised while following it. The site in the Mooswieser Tal was

visited almost daily during the flight period (see Fig. 1), and all

unmarked individuals encountered were marked and previously

marked specimens were recorded.

The S. officinalis stands of the more western meadow margins

were visited less often. All newly encountered individuals were

also marked and we searched for marked adults which might have

emigrated from our main study site.

2.4. Estimation of population size. One way to quantify the

size of a population is to sum the minimum number alive (MNA).

As the name implies, this gives the minimum number of animals

that are present in the population at one particular time. To reach

that number, the number of marked individuals registered (or

marked) in the particular time step (e.g. day) and the number of

individuals marked before that time step and recaptured again after

the time step, but not registered at the time step itself, are added.

Individuals that have left the population in the meantime, as well as

those which were overlooked at the time step and were not re-sight-

ed later (because they have died, emigrated or been overlooked

repeatedly) do not contribute to the MNA of a particular time step.

The total number of individuals present (for a particular time step,

150

like one day or the whole time of a species activity, e.g. 5 weeks)

must therefore be at least as high as this minimal number alive, but

normally is much higher. As the number directly depends upon the

mark recapture efforts invested and the size of the population, its

application is more appropriate in small populations of animals

with a comparatively high recapture rate, but it cannot be reliably

used to compare different populations (Settele et al., 2000).

In order to compare population sizes, the Jolly-Seber method

might be suitable to get a first rough idea on daily as well as total

population sizes (see e.g. Seber, 1982, and Pollock er al., 1990, for

details). For the latter one may e.g. sum up the B-values (i.e. the

individuals added to the population between two population esti-

mates) resulting from the Jolly-Seber model (see Settele ef al.,

2000 for an example and further details with respect to butterflies).

However, one has to keep in mind, that Jolly-Seber estimates are

not very robust with respect to population size.

Table 1. Origin of additional data for the diurnal etho-chronogrammes (Figs 3,

4), which served as a basis for the computing of the age dependent etho-chrono-

gramme (Fig. 5) (M — Mooswieser Tal (Aschbachtal), H — Hagelgrund (Esels-

bachtal)).

Completed classes Origin of data

Age class | Period CS) Ss

before 10:50 | &d 1-2, before 10:50,M | 2 9 1-2, before 10:50, M

10:50-11:40 | 36 0, 10:50-11:40, M+H | 2 2 0, 10:50-11:40, M+H

11:40-12:30 | dd 0, 11:40-12:30, M+H |

15:50-16:40 36 0, 15:50-16:40, M+H |

15:50-16:40 199

1-2, 15:50-16:40, M

16:40-17:30 CADET UM

1-2

after 17:30 2 2 1-2, after 17:30, M

Strictly spoken, only for the age class 1-2 days a complete diurnal etho-chronogramme

could be constructed. Thus, in some cases additional data have been used from the site

Hagelgrund to construct an age dependent etho-chronogramme out of the diurnal etho-

chronogramme. Data of the class 1-2 day old butterflies have been used, if still the pooled

data from both sites (M and H) have not been sufficient. There have been no data of both

sexes of age class 0 for the daily time before 10:50 h. Therefore data of 1-2 day old but-

terflies have been used. For the period 10:50 h — 11:40 h of the O day old butterflies of

both sexes the data of M and H had to be combined to achieve enough observation time.

For the males it also has been necessary to use data from H for the other age classes. For

the 3-10 day old females data of 1-2 day old females have completely been taken for the

last 3 time classes in order not to miss the age dependent etho-chronogramme.

151

individuals

te)

date (1990)

Fig. 1. Minimum daily population sizes (minimal number alive; for days without obser-

vations no entries have been made).

DOSSIERS =

90%

—t— males

80% —{— females

70%

60%

50%

40%

30%

20%

percentage of butterflies (individuals)

10%

0 1 2 3 4 5 6 7 8 9 10

residence time (days)

132

2.5. Ethological observations.

2.5.1. Definitions. The activities flight, moving, nectaring,

oviposition, copulation, and resting were recorded separately for

each individual. Registration was performed with small portable

computers, using a programme especially written for our study.

The observer randomly chose a butterfly and entered its number

into the computer. For each activity an abbreviation, consisting of

one letter, was used. At the beginning of each activity the observer

pressed the relevant letter, which was saved together with the time.

The duration of one activity was obtained from the difference

between the start of the present activity and the start of the next

one.

The activity moving consisted of diverse activities, which were

registered separately in the field, but have been lumped here. These

activities were: walking, turning (e.g. females turning on the flower

heads to look for suitable egg laying niches), cleaning, fluttering

while sitting, and the characteristic lycaenid behaviour of parallel

up and down movement of the closed wings. Thus, turning when

looking for a suitable egg laying niche was not counted as egg lay-

ing behaviour, as it could not be clearly separated from nectaring.

A female looking for an egg laying locality and turning therefore

on the flower, every once in a while is nectaring. Oviposition was

defined as the time while the butterfly has put its abdominal tip

between individual flowers of a flower head. Sometimes females

rapidly changed between different single flowers, performing

abdominal bends and touching the surface of the flower head with

the abdominal tip. Each of these events was regarded as one ovipo-

sition behaviour (sensu Figurny & Woyciechowski, 1998).

Fig. 2. Relative observed residence time and residence time according to the Jolly-Seber-

method. There are hardly any differences between males and females. The residence prob-

ability within the population is 74% for each age, according to Jolly-Seber. This means

that 74% of all individuals reside for 1 day, 55% for 2 days, ... 5% for 10 days. This expo-

nential dependence only is valid if residence probability is age independent. As the

observed number of residing individuals decreases exponentially as well, age indepen-

dence can be assumed. The figure proves the intensive assessment of the population,

because otherwise the Jolly-Seber-curve would be clearly above the observed values.

153

Nectaring was scored as the time when the butterfly put its pro-

boscis into the flowers or when it rapidly moved from one flower

to the next looking for the best nectar source.

About 150 hours of observations were registered on our portable

computers, consisting of about 11,000 events.

2.5.2. Analysis of behavioural observations

2.5.2.1. Procedure and classification. The data have been

analysed so that on the one hand the diurnal distribution of the sin-

gle activities can be described (diurnal etho-chronogramme), and,

on the other to quantify how the sexes differ in their behaviour and

how behaviour changes with age (age dependent etho-chrono-

gramme).

The data were therefore classified as follows:

e sex: male, female

e age: 0 days, 1-2 days, 3-10 days

e time intervals: 10:00-10:50, 10:50-11:40 ... 17:30-18:20

e activities: flight, moving, nectaring, egg laying, copulation,

resting

The classes were kept as small as possible. Even so, the activity

copulation caused problems in analysis, as there have been rela-

tively few events lasting for relatively long time.

This procedure resulted in every day consisting of 10 classes of

50 minutes each. The total observation time within each class (1.e.

the total of all single events) had a length of at least 3,000 seconds

(= 50 min.). However, because few activities occurred early and

late in the day, the first class summarised all results before 10:50

Central European Summer Time (= GMT or 9:20 local time), start-

ing With the first observations at around 9:30; the last class sum-

marised results after 17:30 (GMT), with last observations around

19:30. Most data of the first class are indeed from 10:00 — 10:50

and of the last class from 17:30 — 18:20.

Sexes were treated separately and three age classes formed (see

also results) with an approximately even distribution of total obser-

vation time. This resulted in the age classes 0, 1-2, and 3-10 days.

2.5.2.2. Diurnal Etho-Chronogrammes (Figs 3, 4). Observa-

tion time (t) of one activity was summarised within one class (e.g.:

males, age: 1-2 days; time: 10:50-11:40):

154

> Ciight, 1-2days, 10:50-11:40, all males = Cight, 1-2days. 10:50-11:40. male 1 DE Cight, 1-2days, 10:50-11:40, male 2- =P cee

=> (an AS CIO sac ( i )

Then the relative proportion (T) of one activity compared to the

total duration of all activities was calculated. This was done as in

the following example for flight of the 1-2 day old males for the

period 10:50-11:40:

T flieht, 1-2 days, 10:50-11:40, all males — > Cight, 1-2 days. 10:50-11:40, all en (2 Cight, 1-2 days, 10:50-11:40, all males

ar > Cros 1-2 days, 10:50-11:40, all males oF > regain 1-2 days, 10:50-11:40. all males

+ > on 1-2 days, 10:50-11:40, all males ap > sms 1-2 days, 10:50-1 1:40, all ae) (2)

Figs 3 and 4 show the diurnal etho-chronogrammes of 1-2 day

old males and females. Within the other age classes, data were

insufficient to provide a complete diurnal distribution. Never-

theless, diurnal etho-chronogrammes were computed as these were

needed in the age dependent etho-chronogrammes.

2.5.2.3. Age dependent Etho-Chronogrammes (Fig. 5). The

age dependent etho-chronogrammes show how the distribution of

activities changes with increasing age and how the sexes differ.

Absolute values (in minutes) were calculated from the relative pro-

portions of the time classes of the diurnal etho-chronogrammes. In

other words, we calculated how long a virtual butterfly of a certain

age class performed each activity during a whole day. The compu-

tation is shown for the total flight time of 1—2 day old male butter-

flies:

> Cight, 1-2days, all males = (Taight 1-2 days, before 10:50, al males = Thisht, 1-2 days, 10:50-11:40, all males 2e

* -

ME 2 Thisht, 1-2 days, after 17:30, all Nas) 50 min (3)

In some classes the data were insufficient (observation time in

the 50 minute class was less than 3,000 seconds) to reach a bal-

anced distribution of activity. In these cases, additional data were

included from a nearby locality (H — Hagelgrund, see chapter 2.2).

Even so, data for the last three time classes of 3-10 day old females

were still too few (see Tab. 1). Since the values for the first six time

classes (10:50-15:50) of 3-10 day-old animals were very similar to

those for the diurnal etho-chronogrammes of 1-2 day old insects,

we assumed that the values of the three remaining classes were

only slightly different from the values of 1—2 day old females.

Although this is not entirely correct, for the sake of obtaining a

155

A 0:21 -

- O€-Z1

0r:91

- 05:91

Sits 00:5

ANNEE

ZO ©

WSS ZEN cv:

= OL:VE

ge lee

‘EL 073741

- Or:

- 09:01

‘Ol - 00:01

AuAN9e 18101 Jo 9bejuo91od

time of day

Fig. 3. Diurnal distribution of activities (diurnal etho-chronogramme) of 1-2 days old males (for computation compare chapter 2.5.1).

fi] movement

Wnectaring

El oviposition

E3 copulation

Ei resting

flight

TIMI) 02:8» - ce:2:

Fig. 4. Diurnal distribution of activities (diurnal etho-chronogramme) of 1-2 days old females (for computation compare chapter 2.5.1). In

- OV-91

- 05:S1

- 00:S1

-Ol:vl

-OgEl

- O€-c1

-OV-LL

- 05:01

- 00:01

AyAN2e 18101 Jo 9Bejus919d

time of day

both sexes (Figs 3, 4), the maximum of activity of 1-2 days old individuals is around noon time.

& =

Cre aes

a xs

A À ©

à ©

= 5 2

a Oo >

complete picture of the age dependent etho-chronogrammes we

decided to include values of 1—2 day old females for computing

values of 3-10 day old females. In the first class (before 10:50) of

the O day old insects, no data were available, as the insects had just

been marked. To achieve a complete picture, the values of the 1-2

day old butterflies were also used in the first time class of the 0 day

old ones.

For an overview of the manipulations performed for the

computation of the age dependent etho-chronogrammes see

Table 1.

2.6. Discussion of methodology

2.6.1. Handling and behaviour. Morton (1984) states that

handling in some butterfly species can affect recapture probability.

He also mentions that different species react quite differently in this

respect. However, for G. nausithous there is no indication for a

handling effect. For example, neither Geißler-Strobel (2000) nor

Binzenhöfer & Settele (2000) detected any behavioural change

after handling this species; some individuals even copulated imme-

diately after the handling and females continued to lay eggs

(Binzenhôfer & Settele, 2000).

2.6.2. Observation and behaviour. The presence of (generally)

three observers on a relatively small site might have influenced

adult behaviour, especially flight activity and emigration. However,

our observations suggest that the insects were not affected. We

observed the butterflies from an average distance of about 2 meters,

where they did not show any reaction, although one can approach

as close as few decimetres without noticeably affecting behaviour.

Although the S. officinalis plants were occasionally shaken, this

was greatly outnumbered by the number of natural disturbance

events e.g. by males performing courtship behaviour or by other

insects.

2.6.3. Limits of observation time due to methodological con-

Straints. The insects were observed for as long as possible. How-

ever, we sometimes lost track of flying adults, especially males,

leading to an underestimate of this activity.

159

Table 2: Residence times of G. nausithous on the main research site (n — total

number of butterflies marked)

Number of days

Males (n = 68)

Females (n = 57)

Table 3. Average residence time of the Maculinea species.

Species Average residence | Reference

time [days]

Glaucopsyche (Maculinea)

nausithous

. (M.) nausithous

(M.) nausithous

. (M.) nausithous

(M.) nausithous

(M.) teleius (Bergsträsser)

(M.) teleius

(M.) arion (L.)

(M.) rebeli (Hirschke)

present paper

resent paper?

eißler-Strobel (2000)°

Geißler-Strobel (2000):

Laux (1995)?

Laux (1995)°

Seiler (1991)**

Wynhoff (1998)°*

Laux (1995)°

Wynhoff (1998)

Pauler et al. (1995)*“

Kockelke et al. (1994)*«

Kockelke et al. (1994)**

+003+003+003 +005

O0, 0, 0, 0,

+0 +0 +0 +0

NER SSOS8 8 OR wUNNS

Un 00 00 PITT | COW BW OO

* Average residence time: difference between first and last sighting.

° Average residence time based on Jolly-Seber estimates.

° The authors mention one day more, as ne regard an individual at the day of first sight

as one day old already. Thus the data have been corrected by —1 for this contribution, to

make comparison of data possible.

* data from 1991 until 1996.

Table 4. Number of eggs laid per female per day, based on mean daily behavi-

oural profiles. |

Age class Number of eggs laid (estimated*)

0 day 80

1-2 days 94

3-8 days 81

“ according to our estimate; we assume that in about 95% of all oviposition behaviour

events eggs have really been laid (compare text).

160

Table 5. Average number of flights and average flight duration of 1-2 days old

males and females of Glaucopsyche nausithous during one day of observation

(10:00 to 18:20 h Central European Summer Time).

Sex events/day/individual duration/day/individual duration/event

161 71 min

223 38 min

3. Population ecological results and their discussion

3.1. Population dynamics. The flight period of the population

lasted from 13 July until 28 August 1990 (at the site Mooswieser

Tal). Weather conditions were favourable for nearly the whole peri-

od, with the exception of two rainy days and one further day with

maximum temperatures below 20C. Thus, periods of reduced

abundance were neither expected nor observed during the course of

the population curve (based on the minimal number alive: see

ESP:

The proterandric phenology of G. nausithous 1s clearly visible in

Fig. 1. For example, the date when half the males in the whole pop-

ulation had been observed (29 July) is clearly (5 days) earlier than

the date for half the females (3 August). In G. nausithous, pro-

terandry has been observed in many instances (Geißler-Strobel,

2000; Garbe, 1991; Laux, 1995) and is a common phenomenon in

other butterfly species (see e.g. Wiklund & Fagerström, 1977).

Our populatıon reached its maximum on 3 August 1990, with a

minimum number (see chapter 2.4) of 18 males and 13 females (see

Fig. 1). This was also the day of maximum male numbers, while

that of females (with 15 individuals) occurred on 7 August. As the

population curve is asymmetric (left sided), the date at which half

of the total annual population appeared is earlier (29 July for males

and 3 August for females, see above).

3.2. Population size. In total, 68 males and 57 females were

marked on the main site. A further 18 males and 20 females were

marked elsewhere, of which 28 butterflies were encountered near

the settlement Alte Schmelz, >1.5 km away. None of these marked

specimens were found entering the main site, nor was any speci-

men found outside after being marked on the main site.

161

Due to the intensive effort, nearly every adult entering the popu-

lation was marked on the day it emerged. Thus, an estimate based

on the daily estimates and a summation of the B-values (i.e. the

individuals added to the population between two population esti-

mates) of the Jolly-Seber model (see e.g. Seber, 1982, and Settele

et al., 2000) resulted in 122 specimens, three specimens less than

the 125 marked individuals.

The use of this method was not ideal for this study because the

small population sıze and short residence time of individuals result-

ed in few recaptures on some days. Thus, for a more thorough sta-

tistical analysis, only the central 18 days of field assessment could

be used (omitting the first 5 and the last 7 days). For this period the

total population size is 119, with a 90% confidence interval of +/—

16 (and a MNA of 112; resulting in a population size between 112

and 135; calculation according to Settele et al., 2000:181, Tab.

5.14a; but see “http://www.utz.de/spb/nat/settele/tagfalter.html” for

corrections).

Estimates of total population size support our notion that the

population was almost completely assessed on the main site. In

contrast, the number of individuals outside the main site was

undoubtedly larger than the marked 38 specimens (due to low

intensity of field work and thus low numbers of marked and recap-

tured specimens Jolly-Seber estimates can not be performed). But,

as no individual among those marked outside the main site was

recaptured within the site and vice versa, we assume that the num-

ber of individuals immigrating into the site must have been very

low. Additionally, none of the specimens freshly marked on the site

showed obvious signs of extensive previous flight activity (like loss

of scales or fringed wing margins), which would have suggested it

was an immigrant.

3.3. Dispersal and isolation of the population. Our assump-

tion of low emigration and immigration is consistent with the state-

ments of low mobility of the species by Bink (1992) or Weidemann

(1995). New studies however reveal that it is much less sedentary

(Settele et al., 1996). The longest dispersal distance observed so far

is 5100 m (Binzenhôfer & Settele, 2000). Detailed studies on the

species’ mobility (e.g. Geißler & Settele, 1990; Binzenhöfer &

Settele, 2000) have mainly been performed in landscapes with

numerous local populations and rather large habitat patches. There-

162

fore these distances might be due to mobility within habitats or

between habitats, using stepping stones (e.g. singular Sanguisorba

stands) in the latter case. The low density of habitats in the study

region and the isolated situation of the study site (due to a large

portion of surrounding forests) makes it less probable, that butter-

flies immigrate from other populations in large numbers. As in the

study of Laux (1995), high recapture rates indicate comparatively

sedentary behaviour in these types of environments. Some emigra-

tion as well as immigration can of course never be excluded.

3.4. Residence time. It follows from 3.3 that the vast majority

of butterflies must have hatched from the site itself. We can also be

confident that the majority of animals has been marked on the day

of hatching, so their date of eclosion is known. Consequently, it

was generally possible to identify the exact age of a butterfly at

each activity or recapture.

The average residence time was estimated based on Jolly-Seber

(Seber, 1982; Pollock et al., 1990) estimates of the population size

and residence rate (~). The average residence rate equals —(In@)'

and resulted in a value of 0.74, which is a weighted average (with

respect to the daily populations size), covering the whole flight

period. Because of the small population size, both sexes were com-

bined, which does not introduce bias so long as both sexes have

similar recapture rates, as here (61.8% for males, 56.1% for

females; compare Tab. 2, which also gives an indication for the fre-

quency of capture as assessment was made nearly every day and the

z-values for the Jolly-Seber-estimate are mostly 0).

The residence time of the different age classes (in days) can thus

be expressed as function f(age) = 0.74** (Fig. 2). The average resi-

dence time is the average time one individual spends in the popu-

lation or the time when the population is reduced by 50%, Le.

f(age) = 0.5. In our case, the average residence time was 2.3 days,

which is only slightly higher than the average value resulting from

the difference between the day of marking and the day of last

observation of all butterflies (2.0; compare Fig. 2 and Tabs 2, 3).

This is a function of the comprehensiveness of our observations.

As our population has both been studied intensively and experi-

enced low losses due to emigration, we can regard the average res-

idence time as a good approximation of the average life time. If we

163

additionally regard the average residence times of other authors

(see Tab. 3), we can state that the longevity of adult G. nausithous

is rather short compared to other butterfly species (compare e.g.

Scott, 1973). Looking at other Maculinea species (Tab. 3), this

seems to be typical for the whole taxon (but not exclusive, as shown

for many other temperate-zone Lycaenidae: Väisänen et al., 1994;

Arnold, 1983; Warren 1992).

The observed age independent survival probability of the species

also indicates that the butterflies reach their potential physiological

age only in few exceptional cases, as in all free living organisms.

On our study site we had 6 cases of spider casualties (web or crab

spiders). 2 butterflies were killed by cars on a nearby road. One

specimen just died and dropped off a Sanguisorba stem for no

obvious reason. Despite our intensive study, we thus know the rea-

sons for the death of only 8 out of 125 marked specimens. We only

can assume the high activity of potential predators (high densities

of damselflies, dragonflies and birds along the nearby creek; fur-

ther spiders with undiscovered prey, or nocturnal predators).

3.5. Availability of Sanguisorba flower heads as nectar

sources. The number of flower heads was partly counted. Stands

with counted flower heads were used to estimate further

Sanguisorba stands by eye. The number of heads increased from

ca. 400 ın the early flight days to ca. 1600 towards the end of the

emergence. The number of flower heads must have been sufficient

throughout the flight period, as it probably has only to provide the

carbohydrates needed for flight, as is the case with most short lived

butterflies, which normally acquire the resources needed to devel-

op eggs during the earlier larval stage (capital reserves, compare

Sibly & Calow, 1984, 1986).

3.6. Availability of oviposition sites and emigration. Eggs are

exclusively laid on Sanguisorba officinalis. As shown by Thomas

(1984) and intensively studied by Figurny & Woyciechowski

(1998), closed or only slightly opened flower heads are preferred.

Nearly all flower heads on the study site were in the preferred

stage. The total number of flower heads then steadily increased,

reaching a maximum of about 1600 towards the end of the flight

period, although the number of closed or only slightly opened ones

was lower than in the beginning.

164

Based on our behaviour data (see below) we tried to estimate the

average number of eggs laid per female per day. Laux (1995) stat-

ed, that in G. nausithous only an egg laying behaviour that lasts at

least 3 seconds results in an egg being laid. In most cases she reg-

istered one egg per egg laying event. In our study, in 95% of all

cases, egg laying behaviour lasted at least 3 seconds. In fact, only

in 22 of 53 egg laying events (42%) observed by Laux (1995) were

eggs actually laid. However, we were unaware of this observation

when we collected our data, nor did we interrupt the time registra-

tion when a female quickly inserted her abdomen between single

flowers, which led to the high value of 95%. On the other hand, this

presumed overestimate will be compensated for, as several egg lay-

ing behaviours sensu Laux (1995) have been regarded as only one.

This has to be kept in mind when interpreting the following rough

estimates of eggs laid.

(Precise empirical data on egg production and its dynamics are

lacking for all Maculinea species. Such data are urgently needed in

order to evaluate the usefulness of published estimates, like in Bink

(1992), as well as for parameterizing models.)

According to our estimates, just under 10,000 eggs were laid on

our study site (assuming 60 females with an average life expectan-

cy of 2 days and an egg production of 85 per day; see Tab. 4). The

number of Sanguisorba flower heads was around 1600 at the end

of the flight period, many of which were not available during the

peak of population density at the main flight period. Other flower

heads never reached the size necessary for G. nausithous females

to lay eggs on (according to Figurny & Woyciechowski, 1998, the

flower head has to have a length of at least about 6.5 mm). If we

assume that about three quarters of all heads were used for ovipo-

sition, we would have 1200 flower heads used. This would give an

average of 8 eggs per flower head. This is of the same order of den-

sity reported by Fiedler (1990), who found an average of 7.65 eggs

per head in habitats with high density of butterflies but relatively

few Sanguisorba plants (with a maximum of more than 20 per

head).

Fiedler (1990) found that more than 3 larvae seldom survived in

one flower head. Larger numbers of eggs per flower head thus have

low survival probability, which means that the number of egg-lay-

165

ing possibilities may limit population size. G. nausithous, accord-

ing to Figurny & Woyciechowski (1998), does not seem to be able

to recognise whether a flower head already contains eggs. Hence,

the increased tendency for dispersal within the second half of the

flight period, which was observed by Garbe (1991), may be stimu-

lated by decreasing number of flower heads available in a suitable

stage. As ın our study, the number of preferred non-flowering

flower heads (compare Figurny & Woyciechowski, 1998) was

(even absolutely, not only relatively) decreasing towards the end of

the phenology, ın spite of an increasing total number of flower

heads, we could also imagine such effects to be of relevance for our

population. However, due to the small number of marked butter-

flies, we have no proof for differences in residence times during the

phenology of the species. This merits further study.

4. Results and discussion of behavioural observations

4.1. Preconditions for the interpretation of behavioural

observations. As the population results show, most adults were

marked on the day of hatching. Thus we could identify the age of

the butterfly at a later recapture. We considered that there was no

lack of food, resting sites or similar resources. We assume that the

flower heads often receive more eggs than they can support, as the

females cannot identify whether a head is already occupied. Thus

the behaviour of the adults would not have been affected.

4.2. Handling. Our results show that <1 day-old individuals are

less active than older ones (Fig. 5). For the age class 3-10 days

(diagrams not shown) the diurnal etho-chronogrammes for both

sexes are very similar to those of the 1—2 days old adults (Figs 3,

4). In contrast, the results for age class 0 (diagrams not shown) are

very variable and show no clear maximum of activity around noon.

As stated, Binzenhôfer & Settele (2000) found no handling effect

for this species (see 2.6.1). Our data show that insects are less

active on the day of marking than on other days. This might be

explained as natural for the species or as a handling artefact.

However, as males and females have been affected in a very similar

way, we regard the comparisons between both sexes as justified.

4.3. Copulation and oviposition. For the 1-2 day-old males,

the diurnal distribution of activities is strongly influenced by the

166

occurrence of copulations (see Fig. 3). Although we observed only

2 copulations, these have a strong influence because of their long

duration. The females of this age class normally had no copulations

(Fig. 4). In total, three copulations were observed for their entire

length, which lasted between 24 and 34 minutes.

If observations from Eselsbachtal are included, we know the age

Of 12 females and 10 males of a total of 13 copulating couples.

Females probably copulate only once, in all but one observed case

on the day of emergence. Of the 13 copulations, 11 were in the

morning or early afternoon (until ca. 14:30 h). Only twice was cop-

ulation observed after 17:00 h. These results indicate that females

copulate as early as possible. Only in one female at Eselsbachtal

was copulation observed on the next morning after the day of mark-

ing (but then earlier than all other copulations with newly hatched

females; at 10:27 h, when the couple was sighted, copulation was

already going on).

The 10 males of known age at copulation were up to 7 days old.

The sexual activity of males seems to last for their whole lifetime.

Two of the males were observed copulating twice. According to

SBN (1991), older Maculinea males are more often involved in

copulations than younger ones, which also fits to the observed

lower activity of 0 day adults in our study. And if males mate less

often when they are young, it would not be surprising as they first

have to reach sexual maturity (sclerotization of genitalia, matura-

tion of testes), which may take longer in males of protandrous

species (own unpubl. data; Fiedler, pers. comm.).

In one case, egg laying behaviour was observed as soon as 17

seconds after the end of copulation, although we did not see

whether an egg was actually laid. In a second case, a female did not

lay for 35 minutes, and was then lost from sight; however, after 1

hour and 44 minutes, the same female was resighted laying eggs.

The copulation of females immediately after eclosion and mating

may be seen as consequence of their short residence time. Before

leaving the population (due to death or emigration) they leave as

many eggs as possible in their habitat of origin. According to

Fiedler (pers. comm.) this has to be regarded as a remarkable

exception from all other Polyommatinae lycaenids, where notice-

able egg production starts about 1—2 days after mating and reaches

its peak another 1-2 days later.

167

4.4. Flight. Flight in both sexes peaked shortly after 12:00 h

local time (i.e. 13:30 h central European summer time) when the

sun was already in its highest position, but temperature had not yet

reached ıts maximum (compare Figs 3 and 4). Diurnal changes in

activities were more pronounced in males (Fig. 3) than in females

(Fig. 4). In the males they are clearly paralleled with the time spent

for nectaring, while in females the times are similar but less clear.

During a whole day, males fly twice as long as females (Fig. 5),

although there has been an underestimation in the latter (compare

2.6.3). The number of flights per day however is greater in females

(Tab. 5), resulting in a much lower duration per flight. Thus, with-

in the habitat females mainly undertake very short flights (1.e. from

one flower head or plant to the next in search for oviposition sites).

In both sexes, some flights serve to reach nectar sources. Females

also fly to reach egg laying sites. Potential nectar and egg laying

plants can normally be reached in very short distances. Males

repeatedly undertake longer flights searching for females. The

more a male flies, the higher his probability of finding an unmated

female. On each male there is selection to be more active in flying

and thus more successful in finding unmated females than other

competing males. Thus, we can expect a higher reproductive suc-

cess in more actively flying males, which explains the observed dif-

ferences in sexes in terms of flight.

In some butterfly species there are temperature-dependent

differences in male searching behaviour for females, which might

indirectly lead to a diurnal switch. In Coenonympha pamphilus

(L.), for example, Wickman (1985) observed that males tend to

perch within their territories at lower temperatures, while at higher

temperatures they patrol. Also in G. nausithous males, the greatest

flight activity is around noon, but temperature dependent change

in behaviour (e.g. from patrolling to perching) was neither ob-

served nor expected. As females copulate directly after eclosion

and are not active flyers, it would represent a poor strategy for

males to perch rather than patrol and actively seek newly eclosed

females (see Rutowski, 1991, for a review on male mate-locating

behaviour).

Because they patrol and mate with freshly emerged females,

males have to stay within their habitat to reproduce successfully

168

rather than undertake long distance flights. The latter risk does not

exist for females, which makes them more suitable agents for the

(genetic) exchange between populations and the colonisation of

new habitats (independent of the numbers of males which also

leave the habitat). This fits to the observed higher flight distance of

females and the observations that females have, perhaps temporari-

ly, left the site more often than males.

Dispersal of males from one habitat to another, as observed by

Binzenhöfer & Settele (2000) and Geißler-Strobel (2000) cannot be

disregarded. Its relevance depends much on the distance between

habitats or the degree of isolation of the original habitat. Nearby

habitats might be reached so easily that individuals might be

regarded as belonging to the same population (compare discussions

of gradual changes from subpopulations to proper populations and

finally metapopulations; e.g. Hanski 1999, Settele 1998). In such

cases, male dispersal is expected.

4.5. Movement and nectaring. Nectaring is strongly associated

with flight, especially in males (Fig. 3). Females of all age classes

show more pronounced movement on flower heads than males

(Figs 3, 4, 5), as females have to move for nectaring as well as

between each egg laying event. In females, nectaring and egg lay-

ing may be combined as both activities often happen within a short

time on the same flower head. Males and females both spend less

time nectaring on the day of eclosion than when older (Fig. 5). The

strong decline in nectaring of 3-10 days old males is probably due

to the strong influence of the few but long lasting copulation

events. Most probably in males and females, nectaring has the same

duration, as e.g. also observed for Anthocharis cardamines (L.) by

Wiklund & Ahrberg (1978).

As shown in Fig. 5, both sexes are active for the same period of

time, although the type of activity is very different: males spend

more time on long patrolling flights while females invest more time

visiting flower heads and egg laying. It can be assumed that time of

activity is temperature limited. However, the butterflies often rest

around noon (Figs 3, 4), presumably to recover or to avoid over-

heating.

169

Acknowledgements

For the constant and patient field observations we are very much

indebted to Frank Lohner, Karin Bink, Carola Kleinschmidt, and

Mario Nenno. For organisational assistance we want to thank Uwe

Koch and Michael Werner. The computers for field data registration

have kindly been supplied by Dr. de Kramer (BASF company,

Ludwigshafen). The Fachschaft Biologie of Kaiserslautern

University gave financial support for equipment. For general sup-

port and co-operation we thank Sabine Geißler-Strobel. Critical

and very constructive comments on earlier versions of this manu-

script have been provided by Irma Wynhoff and Konrad Fiedler.

Jeremy Thomas kindly made final comments on content and lan-

guage.

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172

Nota lepid. 23 (2): 173-184; O1.V11.2000 ISSN 0342-7536

Die geographische Variabilität von Erebia

flavofasciata, nebst Beschreibung einer neuen

Unterart (Nymphalidae: Satyrinae)

Frans CUPEDO

Processieweg 2, NL-6243 BB Geulle, Niederlande. e-mail: fcupedo @tip.nl

Summary. According to the male wing-characters, the presently known populations of

Erebia flavofasciata split up in three distinct morphological-geographical groups, the dif-

ferentiation of which is a result of the isolation during the Würm glaciation: ssp. flavo-

fasciata Heyne, [1895], inhabiting the Tessin mountains, ssp. thiemei Bartel, 1904, con-

fined to the Bernina group, and ssp. juliana nov., which is found in the mountains north

of the Inn valley and east of the Ticino valley, from the San Bernardino-pass in the west

to the Samnaun-group in the east. Differentiation of ssp. warreni Verity, 1923 took place

only in postglacial time.

Zusammenfassung. Die bisher bekannten Populationen von Erebia flavofasciata lassen

sich auf Grund der Flügelmerkmale der d 4 in drei verschiedene geographisch-morpho-

logische Gruppen gliedern, deren Differenzierung auf eine Würmglaziale Isolierung

zurückzuführen ist: ssp flavofasciata Heyne, [1895] aus den Tessiner Alpen, ssp. thiemei

Bartel, 1904 aus den Bernina-Alpen, und ssp. juliana nov., deren Areal sich östlich des

Tieinotales und nördlich des Inntales vom San Bernardinopass bis in die Samnaun-

Gruppe erstreckt. Die Differenzierung der ssp. warreni Verity, 1923 ist dagegen post-

glazialen Datums.

Résumé. D’après les caractères alaires des d 6, les populations connues d’Erebia flavo-

fasciata se divisent en trois groupes morphologiquement et géographiquement distincts,

dont la différenciation a été établie suite à l’isolement pendant la période glacière du

Würm: la ssp. flavofasciata Heyne, [1895] des Alpes du Tessin, la ssp. thiemei Bartel,

1904 qui vit dans le groupe de la Bernina, et la ssp. juliana nov., qui se retrouve dans les

montagnes à l’est de la vallée du Tessin et au nord de la vallée de l’Inn, du col San

Bernardino à l’ouest, au groupe Samnaun à l’est. La différenciation de la ssp. warreni

Verity, 1923 fut, au contraire, effectuée durant l’époque postglaciaire.

Riassunto. In base al disegno delle ali dei dd, le popolazioni finora conosciute

dell’Erebia flavofasciata si dividono in tre gruppi morfologici-geografici, la cui differen-

ziatione fu effettuata durante l’isolamento nell’epoca glaciale del Würm: la ssp. flavofas-

ciata Heyne, [1895], abitante delle Alpi Ticinesi, la ssp. thiemei Bartel, 1904, indigena del

gruppo Bernina, e la ssp. juliana nov., che si ritrova diffusa nei monti a Nord della valle

d’Inn ed ad Est della Val Leventina, tra il Passo di San Bernardino ed il Gruppo Samnaun.

La differenziatione della ssp. warreni Verity, 1923 data solo dall’epoca postglaciale.

198

Key words: Nympahlidae, Satyrinae, Erebia flavofasciata, range, variation, new sub-

species, Switzerland.

Einleitung

Erebia flavofasciata Heyne, [1895] hat ein stark zersplittertes

Areal (Abb. 1). Die jetzt bekannten Flugplätze liegen in drei, von

tiefen Tälern voneinander getrennten Gebirgsstöcken:

l. Tessiner Alpen. Hier kommt die Art im oberen Val Maggia

zwischen Passo Campolungo und Passo Naret (Verity, 1953;

Bischof, 1967), im oberen Val Formazza (Bischof, 1967), und am

Alpe Veglia (Verity, 1953; Leigheb, 1976) vor. In den Tessiner

Alpen fliegt die Nominatform.

2. Bernina-Alpen. Die Art fliegt hier lokal in den das Berninatal

umschliessenden Bergketten (Bartel, 1904; Warren, 1923; Verity,

1923). Aus diesem Gebiet sind zwei Unterarten beschrieben wor-

den: die ssp. thiemei Bartel, 1904, vom Schafberg oberhalb von

Pontresina, und die ssp. warreni Verity, 1923', deren Typenlokalität

sich auf der anderen Seite des Berninatales, im oberen Val Roseg

befindet.

3. Adula-Alpen, Albula-Alpen, Silvrettagruppe, Samnaungruppe,

die sıch zu einer ununterbrochenen Gebirgskette nördlich des

oberen Inntales und östlich des Ticinotales zusammenreihen.

(Im Folgenden “Albulakette”” genannt). Die bekannten Flugplätze

liegen hier weit auseinander: San Bernardinopass und oberes

Val Calanca (Vorbrodt, 1931), Julierpass (in der Literatur noch

nicht belegt) und Fimbertal (Pfeiffer & Burmann, 1986). Es gibt

ın der Literatur keine Hinweise dafür, zu welcher Unterart

die Populationen dieses Subareals gehören. Ziel dieser Arbeit

ist es, die Angehörigkeit der Populationen der Albulakette zu

bestimmen.

' thiemei und warreni sind ursprünglich als Varietät beschrieben worden, sind jedoch

gemäß Art. 45.6 der Code (ICZN) als Unterarten zu werten.

174

Abb. 1. Verbreitung von Erebia flavofasciata: A — Tessiner Alpen, B — Albulakette, C —

Bernina-Alpen. Gestrichelte Linie: Grenze des Alpenraumes.

Material und Methoden

Herkunft des Materials. Damit die Variation zwischen den drei

Gruppen mit der Variation innerhalb jeder Gruppe verglichen wer-

den kann, wurden in jedem Teilareal Stichproben von mindestens

zwei, möglichst weit voneinander entfernt vorkommenden

Populationen gesammelt:

1. Alpe Veglia, oberhalb Varzo (Novara, Italien), 1850 m (west-

lichster Fundort der Art).

2. Alpe Pianascio, oberhalb Fusio (Tessin, Schweiz), 2000 m.

Die Fundstelle ist in der Luftlinie ein Kilometer von der Typen-

lokalität von E. flavofasciata (Campolungopass) entfernt.

3. San Bernardinopass (Graubünden, Schweiz), 2200 m.

4. Julierpass (Graubünden, Schweiz), 2400-2500 m.

5. Fimbertal (Tirol, Österreich), 2500 m (östlichster Fundort der

Art).

175

6. Schafberg, oberhalb Pontresina (Graubünden, Schweiz),

2400-2600 m, die Typenlokalität der ssp. fhiemei.

7. Val Roseg, Tschierva Gletscher (Graubünden, Schweiz),

2200-2300 m, die Typenlokalität der ssp. warreni. (Die genaue

Lage der Typenlokalität geht nicht aus Verity, 1923, sondern aus

Warren, 1923 hervor).

Gemessene Merkmale. Die bisher beschriebenen Unterarten

unterscheiden sich voneinander in der Ausdehnung der

Bindenzeichnung auf der Flügeloberseite und -unterseite (Bartel,

1904; Verity, 1923).

Submarginalbinde der Hinterflügelunterseite. Als Maß der

Ausdehnung der Binde wurde die Breite des gelben Flecks ın Zelle

Abb. 2. Der Fleck in Zelle 4 der Hinterflügelunterseite. Breite und Höhe wurden entlang

den eingetragenen Linien gemessen.

176

4 gemessen, und zwar genau durch die Mitte des schwarzen

Zellpunkts (siehe Abb. 2). Es ist der breiteste Fleck der Binde, und

deshalb am einfachsten zu messen, auch bei stark reduzierter

Bindenzeichnung. Die Messung erfolgte mit einem Okular-

mikrometer bei einer Vergrößerung von x20. Da die Falter geringe

Größenunterschiede aufweisen, wurde zudem die Zellhöhe bes-

timmt, ebenfalls durch die Mitte des Zellpunkts. Daraus wurde das

Verhältnis: Breite/Höhe berechnet.

Binde der Vorderflügeloberseite. Wegen der oft diffusen

Begrenzung der Bindenflecke ist deren ‚Breite nicht genau fest-

zustellen. Deshalb wurde der Prozentsatz der Tiere, bei denen der

Fleck in Zelle 3 völlig fehlt, bestimmt. Es ıst dieser Fleck, der als

erster dazu neigt, zu verschwinden.

Weil für eine statistische Bewertung zu wenig ?% zur

Verfügung standen bezieht sich diese Arbeit ausschließlich auf die

Merkmale der dd.

Ergebnisse

Hinterflügelunterseite. In Abb. 3 ist die Verteilung des

Verhältnisses Breite/Höhe graphisch dargestellt. In Tabelle 1 sind

die Populationsmittelwerte und die Standardabweichungen einge-

tragen. Die Unterschiede zwischen Populationen des gleichen

Teilareals sind in allen Fällen weitaus geringer als die Standard-

abweichung. Beim Vergleich von Populationen aus verschiedenen

Teilarealen sind die Unterschiede dagegen etwa zweimal so groß

wıe die Standardabweichung. Weil die Messwerte eine Normal-

verteilung aufweisen, lässt sich mit Students t-Test die Wahr-

scheinlichkeit, dass zwei Stichproben identischen Populationen

entstammen, berechnen (siehe Tabelle 2). Der paarweise Vergleich

von Populationen innerhalb eines Teilareals ergibt dabei p-Werte

zwischen 0.02 und 0.97. Vergleicht man dagegen die Gesamt-

populationen der drei Teilareale miteinander, dann ergeben sich

p-Werte von <0.0001. In Bezug auf die Bindenzeichnung der

Hinterflügelunterseite lassen sich die studierten Populationen also

in drei Gruppen gliedern. Diese entsprechen nicht den drei

beschriebenen Unterarten (flavofasciata, thiemei und warreni),

sondern den drei geographischen Teilarealen der Art (Tessiner

Alpen, Albulakette und Bernina-Alpen).

177

Prozentsatz

06 08 10 12 14 16 18 20 22 APE 6-02

— Fusio + Alpe Veglia -+ S Bernardino + Julierpass

+ Fimbertal + Schafberg = Tschierva

Abb. 3. Das Verhältnis: Breite/Höhe des Flecks in Zelle 4 der Hinterflügelunterseite der

6 Ge

Vorderflügeloberseite. In Tabelle 3 ist pro Population der

Prozentsatz der Tiere ohne Fleck in Zelle 3 der Vorderflügel-

oberseite eingetragen. Hieraus ergibt sich die gleiche Dreiteilung

der Populationen wie aus Tabelle 1. Der Prozentsatz der Tiere mit

verloschenem Fleck ist in den Populationen der Albulakette um

Faktor drei hôher als in den Tessiner Alpen, und in den Bernina-

Alpen wieder um Faktor drei höher als in der Albulakette. Mit dem

x°-Test ist die Homogenität der Populationen der einzelnen

Teilareale, sowie die von allen möglichen Kombinationen von

Teilarealen berechnet (siehe Tabelle 4). Innerhalb der einzelnen

Teilareale liegen die berechneten p-Werte zwischen 0.42 und 0.95;

Kombinationen von Teilarealen ergeben p-Werte von 0.01 und

niedriger.

178

Tabelle 1. Das Verhältnis: Breite/Höhe des Flecks in Zelle 4 der Hinter-

flügelunterseite der d d.

Fusio

A. Veglia

S.Bernardino

FRE

Julier

Fimbertal

Schafberg

Tschierva

NS IRON)

+= ND 00 00 \O

PONNON

N — Zahl der untersuchten Falter; M - Populationsmittelwerte; SD — Standard-

abweichung.

Tabelle 2. Aus Students t-Test hervorgehende p-Werte beim paarweisen

Vergleich des Verhältnisses: Breite/Höhe der untersuchten Populationen.

A. Innerhalb eines Teilareals

Fusio-A.Veglia

S.Bernardino-Julier

S.Bernardino-Fimbertal

Julier-Fimbertal

Schafberg-Tschierva

B. Zwischen Teilareale

Tessın-Albula <0.0001

Tessin-Bernina <0.0001

Albula-Bernina <0.0001

Tabelle 3. Zahl der € € mit (M) und ohne (O) Fleck in Zelle 3 der Vorder-

lügeloberseite. %O — Prozentsatz der Tiere ohne Fleck.

Fusio

A.Veglia

S.Bernardino

Julier

Fimbertal

Schafberg

Tschierva

179

Tabelle 4. Vorderflügeloberseite: Homogenität der einzelnen Teilareale und

derer Kombinationen.

Tessın

Albulakette

Bernina

Tessin+Albulakette

Tessin+Bernina

Albulakette+Bernina

Tessin+Albulakette+Bernina

Schlussfolgerungen.

In den beiden untersuchten Merkmalen liegt die gleiche mor-

phologische Gliederung in drei Gruppen vor. Diese steht mit der

geographischen Gliederung des Areals in Einklang. Dabei erweist

sich die Form der Albulakette als morphologisch intermediar zwis-

chen denen der Tessiner Alpen und der Bernina-Alpen. Aus Abb. 1

ergibt sich jedoch, dass es sich in der Albulakette nicht um

Mischpopulationen handelt, sondern um ein selbständiges Taxon:

die häufigsten Varianten der ssp. flavofasciata und von den ssp.

thiemei und warreni sind in der Albulakette stark untervertreten,

und umgekehrt.

Differenzierungsniveaus. Einer morphologischen Differen-

zierung liegt normalerweise eine langfristige Isolierung zugrunde.

In diesem Fall, beim Entstehen der drei Gruppen von E. flavofasci-

ata, geschah dies während der Würmeiszeit. Das lässt sich daraus

schlieBen, dass man bei E. flavofasciata, wie bei fast allen

Erebienarten, zwei Differenzierungsniveaus unterscheiden kann,

die auf unterschiedliche Isolierungsphasen zurückgehen:

1. Auf dem niedrigeren Niveau gibt es die Unterschiede zwis-

chen heute voneinander getrennten Populationen innerhalb des

gleichen Teilareals (Unterschiede 1. Ordnung sensu Cupedo,

1997). Zum Beispiel die Unterschiede zwischen Alpe Veglia und

Fusio; oder zwischen Schafberg und Val Roseg. Diese sind post-

glazialen Datums. Sie sind eine Folge der jetzt noch andauernden

postglazialen Isolierung der einzelnen Populationen. Bei E. flavo-

fasciata besteht wohl kaum Zweifel, dass der Genfluss zwischen

den Populationen gegenwärtig völlig unterbrochen ist, und dass

180

diese Situation schon seit etwa zehn Jahrtausenden anhält. Die

Abtrennung der ssp. warreni von der ssp. thiemei beruht also auf

Unterschieden, die erst in postglazıaler Zeit entstanden sind.

2. Auf dem höheren Niveau gibt es die Unterschiede zwischen

den Gesamtpopulationen der drei Teilareale (Unterschiede 2. Ord-

nung). Diese müssen zustande gekommen sein ın einer Periode, in

der die (jetzt voneinander isolierten) Populationen jedes Teilareals

noch eine einzige, zusammenhängende Population bildeten. Das

war vor der Wiederbesiedlung der Alpen, ın ıhren Glazialrefugien

irgendwo am Alpenrand, der Fall (für eine ausführlichere

Auseinandersetzung siehe Cupedo, 1997). Das deutet darauf hin,

dass sich E. flavofasciata während der Würm-Vereisung in drei

verschiedene Refugien am Alpenrand zurückgezogen hat.

Taxonomie. E. flavofasciata lässt sich also in drei morpholo-

gisch erkennbare geographische Gruppen gliedern, deren Differen-

zierung durch langfristige (mehr als 100.000-jährige) Isolierung in

voneinander getrennten Glazialrefugien zustande gekommen ist.

Dies ist eine natürliche Gliederung der Spezies in Subspezies. Die

ssp. flavofasciata bildet die erste Gruppe. Die zweite natürliche

Einheit wird von den ssp. thiemei und warreni gebildet. Die noch

nicht beschriebene Unterart der Albulakette nenne ich.

Erebia flavofasciata juliana ssp. nov.

Holotypus d, etikettiert: ‘Julierpass (Graubünden, CH); 2400-2500 m.; 22.V11.[19]91;

F.Cupedo les, in Coll. F. Cupedo. Paratypen: 39d, 169 in Coll. F. Cupedo, 106 in

Coll. Instituut voor Systematiek en Populatiebiologie (Zoölogisch Museum), Amsterdam.

Alle mit gleichem Etikett.

Beschreibung. Die Submarginalbinde der Hfl.Us und Vfl.Os der

dd sichtlich schmaler als bei der ssp. flavofasciata, jedoch nicht

so reduziert wie bei der ssp. thiemei (Abb. 4 und 5).

Verbreitung. Adula-Alpen und Albula-Alpen, östlich bis in die

Samnaungruppe.

Derivatio nominis. Der Name leitet sich vom Julierpass, der

Typenlokalität der Unterart, her.

Weil der Unterschied zwischen den beiden Differenzierungs-

niveaus nomenklatorisch nicht zu erfassen ist, ist in Abb. 6 der

181

Abb. 4. Erebia flavofasciata 3 3, Oberseiten: Links ssp. flavofasciata, Mitte ssp. juliana

(zweite von oben: Holotypus), rechts ssp. thiemei.

Abb. 5. Wie Abb. 4, Unterseiten.

182

natürliche Zusammenhang der Unterarten von E. flavofasciata

schematisch dargestellt.

Diskussion

E. flavofasciata ist ein Beispiel dafür, wie eine bestehende sub-

spezifische Gliederung der Einsicht in die natürliche Gliederung

einer Art im Wege stehen kann. Die Ursache davon ist, dass nicht

alle beschriebenen Unterarten gleichwertig sind. Postglazial ent-

standene Unterarten sind mit Unterarten glazialen Alters nicht zu

vergleichen. Postglaziale Differenzierung spielt sich bei den

Erebien (und natürlich nicht nur bei diesen) auf Populationsniveau

ab. Demzufolge beziehen sich Beschreibungen von postglazial dif-

ferenzierten Unterarten prinzipiell nur auf eine einzige Population.

Die Population vom San Bernardinopass könnte mit genau soviel

Recht von der ssp. juliana abgetrennt werden, wie warreni von

thiemei. Und im Grunde genommen trifft das für jede isolierte

alpine Erebia-Population zu. Unterschiede 1. Ordnung sind daher

für das Verständnis des natürlichen Zusammenhangs der

Populationen (also für die Taxonomie) ohne Bedeutung. Beim

Studium der Variabilität der einzelnen Arten ist dies zu berück-

sichtigen.

E. flavofasciata

A. flavofasciata juliana thiemei+warreni

B. thiemei warreni

Abb. 6. Der natürliche Zusammenhang der Unterarten von Erebia flavofasciata:A —

Gliederung infolge glazialer Isolierung, also auf Unterschiede 2. Ordnung gegründet; B —

Gliederung infolge postglazialer Isolierung, also auf Unterschiede 1. Ordnung gegründet.

Dankwort

Die wertvollen Bemerkungen von zwei anonymen Referenten

haben zu ansehnlichen Verbesserungen des Manuskripts geführt.

183

Besonderen Dank schuldige ich Herrn Manfred Pfeiffer (Ischgl,

Österreich), dank dessen Bemühungen ich über eine Serie von

E. flavofasciata aus der Samnaungruppe verfügen konnte, und

Herrn Marcel Prick (Heerlen, Niederlande), der den deutschen Text

korrigierte.

Literatur

BARTEL, M., 1904. Über eine neue Form von Erebia flavofasciata Heyne. —

Dt.ent.Z.Iris 17: 164-167.

BiscHor, A., 1967. Beitrag zur Kenntnis des Verbreitungsgebietes von Erebia

flavofasciata Ruehl. — Mitt.ent.Ges.Basel 17: 1-3.

CupEDo, F., 1997. Die geographische Variabilität und der taxonomische Status

der Erebia manto bubastis-Gruppe, nebst Beschreibung einer neuen Unterart

(Nymphalidae: Satyrinae). — Nota lepid. 20(1/2): 3-22.

ICZN — International Code of Zoological Nomenclature, 1999. Fourth edition,

adopted by the International Union of Biological Sciences. International Trust

for Zoological Nomenclature, c/o The Natural History Museum, London. xxx

+ 306 pp.

LEIGHEB, G., 1976. Contributo allo studio delle Evebia alpine italiane: II. Erebia

flavofasciata Heyne (Lepidoptera Satyridae): morfologia, eto-ecologia,

corologia. — Redia 59: 345-353.

PFEIFER, M. & BURMANN, K., 1986. Erebia flavofasciata Heyne, 1895.

Erstnachweis für Österreich. (Insecta: Lepidoptera, Satyridae). — Ber.nat.

med. Ver. Innsbruck 73: 155-157.

VERITY, R., 1923. On some Italian races of Erebia aethiops, Esp., and of E.

pirene, Hub. (=stygne, O.), and on E. flavofasciata, Ruhl-Heyne. —

Entomologist’s Rec.J.Var. 35: 134-136.

Verity, R., 1953. Le farfalle diurne d’Italia, vol. V. — Marzocco, Firenze:

18-354.

VORBRODT, C., 1931. Tessiner und misoxer Schmetterlinge. — Mitt.schweiz.

ent.Ges. 14: 201-395.

WARREN, B. C. S., 1923. Six weeks among the Butterflies of Eastern Switzerland.

— Entomologist’s Rec.J.Var. 35: 73-79, 96-101.

184

Nota lepid. 23 (2): 185-190; 01.VII.2000 ISSN 0342-7536

Notes on some interesting Geometridae

collected in Tuscany (Italy)

Leonardo Dapporto & Filippo FABIANO

Museo Zoologico “La Specola”, Via Romana 17, I-50125 Firenze

e-mail: lepidoptera@ www.specola.unifi.it

Summary. Eleven species of Geometridae are listed. Among these, Eupithecia orphnata

W. Petersen, 1909 is new for the Italian fauna and Stegania cararia (Hübner, 1790) is

recorded for the first time from the Italian peninsula. Furthermore, six species are new for

the fauna of Tuscany. For each species the presently known distribution in Italy is given.

Zusammenfassung. Elf Arten der Geometridae werden aufgelistet. Von diesen wird

Eupithecia orphnata W. Petersen, 1909 erstmalig fiir die Fauna Italiens nachgewiesen,

Stegania cararia (Hibner, 1790) erstmalig von der Apenninenhalbinsel. Weitere sechs

Arten sind Neufunde fiir die Fauna der Toskana. Fiir jede Art wird die derzeit bekannte

Verbreitung in Italien dargestellt.

Résumé. Onze espèces de Geometridae sont recensées. Parmi celles-ci, Eupithecia

orphnata W. Petersen, 1909 est nouvelle pour la faune italienne et Stegania cararia

(Hübner, 1790) est mentionnée pour la première fois de la péninsule italienne (déjà con-

nue du Piémont). De plus, six espèces sont nouvelles pour la faune de Toscane. Pour

chaque espéce, la répartition connue actuellement en Italie est présentée.

Key words: Lepidoptera, Geometridae, new records, Italy, Tuscany.

Tuscany is a region with a great diversity of environmental con-

ditions and, consequently, a rich Lepidopteran fauna. Our present

knowledge in this field is quite good, thanks to the efforts of many

lepidopterists who carried out specific studies in this region during

the last two centuries.

Nevertheless, the moth fauna in several areas of Tuscany still

remains poorly known and, consequently, we have focused our

attention on such areas.

The Apuan Alps is a mountainous chain that, despite its low

elevation, has a cold alpine-like climate. This is evidenced by the

presence of alpine species (e.g. Erebia spp.) that are absent in the

Italian peninsula or limited to the higher altitudes in the central

Apennines (Marche, Abruzzi, Lazio). The Farma Valley in the

185

“Colline Metallifere” (central Tuscany) is a well known refuge

area. It is located in a warm Mediterranean region, but its peculiar

geomorphology confers it meso-frigophilous characters. It is one of

the two known localities of central Italy where Archiearis notha

(Hiibner, [1803]) is found (Fabiano & Zilli, 1998).

Our research in these areas, as well as in other interesting zones

of Tuscany, led us to discover some rare or newly found

Geometridae. This paper presents an annotated list of these species,

among which Eupithecia orphnata W. Petersen, 1909 is also new

for the Italian fauna.

Unless otherwise stated, the specimens have been collected by

the authors and are being kept in their collection.

Chloroclysta miata (Linnaeus, 1758)

8 , Massa, Apuane Alps, Passo del Vestito, 1100 m, 9.X.1999.

Distribution in Italy: Trentino, Alto Adige, Veneto, Friuli (Wolfsberger, 1971),

Emilia, Marche, Lazio, Abruzzi, Sardinia (Prola & Racheli, 1979; Raineri & Zangheri,

1995). — New for Tuscany.

Triphosia sabaudiata (Duponchel, 1830)

23, Lucca, Apuane Alps, Mount Matanna, 1050 m, 17.VII.1999.

Distribution in Italy: Piedmont, Lombardy, Trentino, Alto Adige, Veneto, Friuli,

Emilia, Marche, Lazio, Abruzzi, Campania, Lucania (Parenzan, 1994). — Rare mountain

species, occurring from 1000 to 2000 m and more. First record from Tuscany.

Solitanea mariae (Stauder, 1921)

26 , Lucca, Apuane Alps, Fociomboli, 1230 m, 23.VII.1999; 9, Siena, Farma Valley, Iesa,

Carpineto, 200 m, 20.VIII.1999; 3, 9%, Arezzo, Casentino, Moggiona, 700 m,

26.V1.1997, 4.VIII.1997; 23, Arezzo, Casentino, Camaldoli, 800-1000 m, 29. VII.1997,

23.2.1997.

Distribution in Italy: Veneto, Liguria, Romagna, Tuscany, Umbria, Marche, Lazio,

Campania, Lucania, Calabria (Parenzan, 1994). — Previous records of this species in

Tuscany were scarce. It appears now to be well distributed in the region.

Eupithecia extraversaria Herrich-Schäffer, 1852

d, Siena, Farma Valley, Iesa, Carpineto, 200 m, 13.VI.1997, F. Balderi leg. et coll.

Distribution in Italy: Trentino, Alto Adige, Veneto, Romagna, Marche, Lucania

(Parenzan, 1994). — Scattered populations in the Alps. Extremely scarce in the Italian

186

peninsula where it appears to be confined to mountain areas between 1300 and 1400 m,

except for a single female collected by P. Zangheri near Forli (Fiumi & Camporesi, 1988).

First record for Tuscany.

Eupithecia gueneata Mabille, 1862

4 , Lucca, Apuane Alps, Mount Matanna, 1050 m, 17.VII.1999.

Distribution in Italy: Lombardy, Trentino, Alto Adige, Veneto, Emilia, Lazio,

Lucania, Calabria, Sicily (Parenzan, 1994). — New for Tuscany.

Eupithecia inturbata (Hübner, [1817])

4,39, Siena, Farma Valley, Iesa, Carpineto, 200 m, 8.IX.1999, 2.X.1999.

Distribution in Italy: Alto Adige (Dannehl, 1927), Emilia, Abruzzi (Flamigni, Bastia

& Dapporto, in prep.). — The first record of this very rare species from Tuscany.

Eupithecia orphnata W. Petersen, 1909

29, Lucca, Apuane Alps, Mount Matanna, 1050 m, 21.VI.1999, 17.VII.1999.

Distribution in Europe: Sweden, Finland, Poland, Czech Republic, Slovakia,

Germany, France, Spain, Switzerland, Austria, Hungary, Romania, Bulgaria, Albania

(Karsholt & Razowski, 1996). — This species, typical of cold climates, is new for the

Italian fauna. Its presence in the Apuane Alps is noteworthy, as it evidences the impor-

tance of this mountain chain as a refuge area for the northern species who colonised the

Italian peninsula during the glacial periods. Many specimens of Eupithecia subumbrata

([Denis & Schiffermüller], 1775) have been collected in the same locality and at the same

time. E. orphnata and E. subumbrata are difficult to tell apart by their wing pattern,

although the first is usually darker. Correct identification depends on examination of the

genitalia (Figs 1-2). E. subumbrata has been recorded for the first time in the northern

Apennines to Poggio della Doganaccia (Pistoia) (Flamigni, Bastia & Dapporto, in prep.).

Stegania cararia (Hübner, 1790)

53, 2, Siena, Farma Valley, Iesa, Carpineto, 200 m, 6.V.1999, 27.V.1999, 14.VII.1999,

13.VIIL.1999; à , Siena, Farma Valley, Iesa, Carpineto, 200 m, 13.VI.1997, F. Balderi leg.

et coll.

Distribution in Italy: Piedmont (Mariani, 1941), Brenta Mountains, and few other

localities in the Alps (Hellmann, 1987). — This is the first record of the species for the

Italian peninsula (Figs 3-5). The presence of this species in the Farma Valley is a further

confirmation of the refugial character of the site. Besides the above mentioned Archiearis

notha, Eupithecia extraversaria and Eupithecia inturbata, other species typical of colder

climates like Cleorodes lichenarius (Hufnagel, 1767), Gluphisia crenata (Esper, 1785),

Lamprotes c-aureum (Knoch, 1781), Catocala fraxini (Linnaeus, 1758) and Hyphoraia

testudinaria (Geoffroy, 1785) are found here.

187

Megalycinia serraria (A. Costa, 1881)

?, Siena, Farma Valley, Bagni di Petriolo, 160 m, 20.IX.1998; 29, Arezzo, Casentino,

Moggiona, 700 m, 18.1X.1995, 23.1X.1997.

Distribution in Italy: Emilia (Flamigni, 1990); Romagna, Tuscany, Marche, Umbria,

Lazio, Abruzzi, Lucania, Calabria, Sicily (Parenzan, 1994). — This Italian endemic

species has been recorded from about twenty localities of central and southern Italy. A

single old record from Camaldoli (Arezzo) (Calberla, 1890) was known hitherto in

Tuscany.

Cleorodes lichenarius (Hufnagel, 1767)

d, Siena, Farma Valley, Bagni di Petriolo, 160 m, 20.1X.1998.

Distribution in Italy: Trentino, Piedmont, Lombardy, Romagna, Tuscany, Umbria,

Lazio, Abruzzi, Apulia, Sicily, Sardinia (Parenzan, 1994), Friuli, Calabria (Parenzan er

al., 1998) Emilia (Flamigni, Bastia & Dapporto, in prep.). — This uncommon species is

usually found in mountain habitats and is extremely rare at low altitudes. It had not been

collected in Tuscany since the 19th century (Zeller, 1849; Calberla, 1890).

Yezognophos dilucidarius ([Denis & Schiffermüller], 1775)

28, Lucca, Apuane Alps, Campocatino, 10.VIII.1999.

Distribution in Italy: Trentino, Alto Adige, Veneto, Lombardy, Piedmont, Liguria

(Wolfsberger, 1971). — An old record from Emilia appears to be doubtful and probably

referable to Yezognophos dognini (Thierry-Mieg, 1910) (Flamigni, Bastia & Dapporto, in

prep.). New for Tuscany.

Acknowledgements

We are grateful to Mr Felice Balderi (Lucca), who kindly allow-

ed us to examine his collection and to publish some of his data.

References

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brata, female genitalia, Apuane Alps; 3. Stegania cararia, Farma Valley; 4. Stegania

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Boll. Inst.Ent.Univ.Bologna 34: 191-246.

RAINERI, V. & ZANGHERI, S., 1995. Lepidoptera Drepanoidea, Axioidea,

Geometroidea. Jn: Minelli, A., Ruffo, S. & La Posta, S., (eds). Checklist delle

specie della fauna italiana. 90. — Calderini, Bologna. 23 p.

WOLFSBERGER, J., 1971. Die Macrolepidopteren-Fauna des Monte Baldo in

Oberitalien. — Memorie Mus.civ.Stor.nat. Verona fuori serie 4: 1-333.

ZELLER, P. C., 1849. Verzeichniss der von Herrn Jos. Mann beobachteten

Toscanischen Microlepidoptera. — Stettin.ent.Ztg. 10: 200-220.

190

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NOTA LEPIDOPTEROLOGICA

A journal of the Societas Europaea Lepidopterologica

Published by Societas Europaea Lepidopterologica

Vol. 23 No. 3/4 Basel, 01.12.2000 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

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Contents @ Inhalt e Sommaire

NEKRUTENKO, Y. P. A catalogue of the type specimens of Palaearctic

Riodinidae and Lycaenidae (Lepidoptera, Rhopalocera) deposited in the

collection of the Museum fiir Naturkunde der Humboldt Universitat zu

a nem emery ree UNS Stalin wy bene Dal na. Jee Anal 192

Vor. 23 — 2000

Dates of publication — Publikationsdaten — Dates de publication ....... 333

New taxa described in Vol. 23 — Neue Taxa in Band 23 beschrieben —

nine aeeaxadeentsidans le Vole23) 15... 353

19]

Nota lepid. 23 (3/4): 192-352; 01.X11.2000 ISSN 0342-7536

A catalogue of the type specimens of

Palaearctic Riodinidae and Lycaenidae

(Lepidoptera, Rhopalocera) deposited in

the collection of the Museum für Naturkunde

der Humboldt Universität zu Berlin

Yuri P. NEKRUTENKO

Schmalhausen Institute of Zoology, UA-01601 Kiev 30, MSP, Ukraine

e-mail: ypnekrut @ mbat.freenet.kiev.ua

Summary. A revision of the Lepidoptera collection of the Museum fiir Naturkunde der

Humboldt Universität zu Berlin showed it to contain the type material of 3 nominal

species-group taxa of Palaearctic Riodinidae and 273 of Lycaenidae; of these 42 label

names were found to be unavailable as infrasubspecific (17), preoccupied (22; published

replacement names are cited) and/or unpublished (3); the type material of 30 species-

group taxa, thought to be deposited in the collection, was not found. Among others, the

collection contains type material of Alphéraky, A. & O. Bang-Haas, Christoph, Grum-

Grshimailo, Herrich-Schäffer, Klug, Kollar, Lederer, Püngeler, Rebel, Staudinger and

others.

Zusammenfassung. Eine Revision in der Schmetterlingssammlung des Museums fiir

Naturkunde der Humboldt Universität zu Berlin zeigte, daß das Typenmaterial nomineller

Taxa der Artengruppe von 3 paläarktischen Riodinidae sowie 273 Lycaenidae vorhanden

ist; von diesen sind 42 Namen nicht verfiigbar als infrasubspezifisch (17), präokkupiert

(22, veröffentlichte Ersatznamen werden zitiert) und/oder unveröffentlicht (3); das

Typenmaterial von 30 Taxa der Artengruppe, welches in der Sammlung vermutet wurde,

konnte nicht gefunden werden. Die Sammlung enthält unter anderem Typenmaterial von

Alphéraky, A. & O. Bang-Haas, Christoph, Grum-Grshimailo, Herrich-Schäffer, Klug,

Kollar, Lederer, Piingeler, Rebel, Staudinger u. a.

Résumé. Une révision de la collection de lépidoptéres du Museum fiir Naturkunde der

Humboldt Universität zu Berlin a démontré que celle-ci contient le matériel type de

3 taxons nominaux du niveau espéce de Riodinidae et 273 de Lycaenidae; parmi ceux-ci,

42 noms se sont révélés non disponibles, étant infrasubspécifiques (17), préoccupés (22;

les noms de remplacement publiés sont cités) et/ou non publiés (3); le matériel type de

30 taxons du niveau espèce, supposé être déposé dans la collection, n’a pas été retrouvé.

La collection comprend, entre autres, du matériel type d’Alphéraky, A. & O. Bang-Haas,

Christoph, Grum-Grshimaïlo, Herrich-Schäffer, Klug, Kollar, Lederer, Püngeler, Rebel,

Staudinger et autres.

192

Key words: Lepidoptera, Rhopalocera, Riodinidae, Lycaenidae, Museum für

Naturkunde der Humboldt Universität zu Berlin, collections, type specimens, nomencla-

ture.

...et non est creatura invisibilis in conspectu eius, omnia autem

nuda et aperta sunt oculis eius, ad quem nobis sermo.

Ad Hebraeos 4, 13

The Museum für Naturkunde der Humboldt Universität zu

Berlin is a depository of the largest and most important reference

collection of the Lepidoptera in Europe. It contains the type series

of the species-group nominal taxa established during the 19th and

20th centuries by the founders of modern lepidopterology Gottlieb

August Wilhelm Herrich-Schäffer, Julius Lederer, Otto Staudinger

and their successors Andreas and Otto Bang-Haas and many of

their contemporaries, giving the Museum an importance of the

Promised Land for anyone involved with systematics, taxonomy,

nomenclature, faunistics and zoogeography of Palaearctic and

extra-Palaearctic Lepidoptera.

This publication is one of the results of my research visit to the

Humboldt University Natural History Museum during October-

December 1998, sponsored by the Deutscher Akademischer

Austauschdienst (DAAD). At the outset of this project, the

Palaearctic Riodinidae/Lycaenıdae collection was rearranged to

assemble chaotically diffused material into an integrated body

convenient for practical use.

A first examination of the Lepidoptera collection as a whole

brought to light that Palaearctic Riodinidae/Lycaenidae material

was scattered among 5 main more or less compact collection

bodies left almost intact since their arrival to Berlin:

(1) Primary collection arranged by Klug or under Klug’s supervision (early 1820’s).

The Berlin Lepidoptera collection became renowned after 1818, when Johann

Christoph Friedrich Klug took up the post of the second director (after Illiger) of the

Zoological Collection. From this time on (tentatively from between the 1820’s and

1830’s) a documentary history of the collection begins in form of a hand-written invento-

ry book entitled Catalogus Generalis Musei Zoologici Berolinensis. The Lepidoptera sec-

tion of this document is bound in several volumes (Vol. 1: Nos 1-3753; Vol. 2:

Nos 3754-10098 enumerated in chronological order but not dated — see Appendix)

and is deposited in the Museum’s Lepidoptera Section library. It reflects the collection

inventory connected with the collection itself through the specimens’ inventory numbers

198

(first column of each Catalogus entry/row). In its original arrangement, the specimens

were placed in conventional insect drawers, each series or single specimen under a draw-

er label with species-group name and author, reference to original and subsequent

descriptions and illustrations, locality and collector (provider); the first specimen below

the drawer label was labelled with a printed inventory number referring to a Catalogus

Generalis row of the following columns: inventory (label) number | binominal or trino-

minal name | number of specimens | locality | collector/provider. All these data, along with

those contained in the original description, may provide some evidence for the origin and

thus for the nomenclatural status of the specimen(s). All Catalogus Generalis entries for

type specimens are quoted in the Appendix below.

(2) Staudinger collection (including Lederer and partly Herrich-Schäffer collections as

an integral part; received 1948 — see Horn et al., 1990: 168, 229, 377).

The Staudinger collection, in its primary state, has been placed in so-called “glass

cases”: wooden drawers with a glass lid and a glass bottom supplied with transverse

grooved wooden bars parallel to smaller side. The groove of each bar is filled with a cork-

or peat-like material in which pinned specimens could be set in one line columns far

enough from one another to accommodate specimen wingspan. To save space, the speci-

mens in a column were originally pinned tilewise overlapping (Fig. 1, bottom left), caus-

ing a tangible inconvenience in manipulations that often resulted in damage from clumsy

handling. Each species-group taxon was set under a drawer name label, with each series

beginning with a specimen having a handwritten coloured locality label; the rest of the

specimens from the same locality are usually labelled with circles punched from the same

label paper as the locality label. In addition to locality labels, most of the type specimens

also have a characteristic printed label ”Origin.[al]” (on pink paper), with the first speci-

men in the type series being supplied with a handwritten (by Staudinger) identification

label. The specimens from the Lederer collection were signed by Staudinger with printed

labels "Coll. | Led.” (on white paper). In case of non-designated types, when their type

status (as syntype or holotype by monotypy) was traced and determined now by external

unambiguous proof, they were provided with new designation labels.

(3) Bang-Haas (A. & O.) collection (received 1948 — see Horn et al., 1990: 27, 377).

(4) Püngeler collection (acquired 1928 — Horn et al., 1990: 314).

(5) New contributions from different donors (mostly after World War II) irregularly

placed into free space in any cabinets and drawers.

When I reorganised the collections, the specimens were removed

from the primary collections, pinned into plastic unit trays in their

original order (each taxon separately) and assembled in conven-

tional collection drawers to be housed in special new cabinets

(Fig. 1, top left and right). Each specimen was supplied with a

printed label to indicate the collection of origin: “ex coll. |

STAUDINGER’, “ex coll. | [A. and/or O.] BANG-HAAS”, “ex coll

| PÜNGELER” and “Zool. Mus. | Berlin” (for specimens of the pri-

mary collection and new contributions as specified respectively

under (1) and (5) above). The specimens of the type series were

194

marked with the same labels numbered with fraction numbers to

show the number of specimens in a series and a serial number of a

specimen in its original position (for instance, 2/8 means second

specimen in a series of eight). The type material has been placed in

separate unit trays clearly designated with recognisable drawer

labels.

All of Klug’s extant Rhopalocera type material is now merged in

two clearly marked drawers and is easily accessible (for a complete

review see Olivier & Nekrutenko, 2000).

In the present publication, the extant type specimens (syntypes, holotypes by mono-

typy and, when validly fixed, lectotypes, paralectotypes, neotypes with reference to pri-

mary sources) of the riodinid and lycaenid butterfly nominal species-group taxa deposit-

ed in the Lepidoptera collections of the Museum für Naturkunde der Humboldt

Universität zu Berlin are listed and commented. The species-group taxa names are

arranged in alphabetical order, separately for Riodinidae and Lycaenidae.

Each entry opens with the species-group name originally applied to the specimens,

with its author and date (as a heading line); available names are given in boldface,

unavailable ones (infrasubspecific, junior homonyms, unpublished, nomina nuda) in

lightface (as here) characters; with homonyms and unpublished names not italicized). The

heading line is followed by a paragraph containing original combination quoted from the

primary source with full reference; a section Type locality containing a quotation from the

original description along with data on the type material when stated; then, after an em-

dash (—), a synonymy, homonymy (and a replacement name when available) are given

with reference to the appropriate literary sources.

For each specimen the label text is quoted in single quotation marks in following order:

identification and locality label; accessory label(s) (origin, depository, inventory (cata-

logue) nos. etc.); designation label(s). In cases when a specimen is considered to belong

to the type series only doubtfully, it is indicated in the list with a question mark (?-d or

7-9).

In the section Types (entitled, dependent on status, as Syntype(s), Holotype,

Paratype(s), Allotype, Lectotype, Paralectotype(s), Neotype), to assist better recognition

of the specimen, the quotation of each label is provided with the characteristics of the

paper on which it is written. In run-in quotations, each line of the label text is separated

by a vertical line (I). All extensions of abbreviations used in the label text are enclosed in

square brackets ([ ]); necessary comments or remarks inserted in the label text are given

in angle brackets (< >). The square brackets are still used here to include the year of

publication established after external sources, although this is no longer recommended in

the current edition of the Code. In quotations of combined labels (handwritten on printed

forms), the handwritten text is reproduced in italics; completely handwritten and com-

pletely printed labels are quoted in a standard character face (as here); the printed labels

are quoted with their original text layout. When important and known, for handwritten

text the writer’s name is given in parentheses.

195

This Catalogue reflects the state of the collection on 31 December 1998. Subsequent

taxonomic treatments (lectotype designations etc.) will, however, briefly be referred to,

with reference to the publication.

Fig. 1. — Lycaenidae in the collection of the Museum für Naturkunde der Humboldt

Universität zu Berlin, before (bottom left) and after (top left, right) rearrangement for the

purpose of the present Catalogue.

196

RIODINIDAE

lua Grum-Grshimailo, [1891]

‘Polycaena Lua’ Grum-Grshimailo, [1891]: 454. Type locality: ‘In montibus Sinin-

Schan detecta’.

Syntypes 2d, 9:

e 3 with handwritten labels: “Thibet | Sinin Alp.[es] | [18]90

[legit] Gr. Gr. (on brown paper), small date label

‘27.V1.[18]90’ (on faded white paper, Grum-Grshimailo),

‘Polycaena | Lua | Sinin Schan’ (on white faded paper, A. Bang-

Haas);

e 3, 2 with circles of brown locality label paper.

The syntypes, received by Staudinger from Grum-Grshimailo,

are provided now with a printed label ‘/ua | Grum-Grshimailo

[1891] | SYNTYPUS | Y. Nekrutenko det. 26.11.1998’ (on yellow

paper).

tamerlana Staudinger, 1886

‘Polycaena (nov. genus) Tamerlana Stgr. Staudinger, 1886: 227. Type locality:

“...wurde 1882 von Haberhauer jun. bei Osch entdeckt... Später erhielt ich auch Stücke

vom Alai und Namangan...’

Syntypes 9 , 49:

e d with labels: handwritten (Staudinger) ‘Polycaena Stgr. |

Tamerlana Stgr.’ (on white paper), handwritten ‘Margelan |

Maur.[er] [18]84’ (on brown paper), printed ‘Origin. (on pink

paper) and a cut [from envelope or wrapper?] ‘von 17/6 bis 29/6

<year?> (on white faded paper);

e 6 with labels: handwritten ‘Osch | Hbh. [Haberhauer]’ (on

brown paper), handwritten date label ‘19/7 [1882]? (on white

paper) and printed ‘Origin.’ (on pink paper);

e 54,39 with circles of brown locality label paper and printed

‘Origin. (on pink paper), one d also with handwritten

(Staudinger) label ‘Tamerlana | Stgr.’ (on white paper);

e d with labels: handwritten ‘Namangan | Hbh. [Haberhauer]

[18]84’ (on brown paper) and printed ‘Origin. (on pink paper);

e 3, © with circles of brown locality label paper and printed

‘Origin. (on pink paper).

timur Staudinger, 1886

‘Polyc.[aena] Tamerlana var Timur Stgr. Staudinger, 1886: 230. Type locality: ‘Herr

Tancré sandte mir von seinem Sammler Rückbeil im Kuldja-District gefundene

Polycaena...’

Syntypes 64,39:

e d with labels: handwritten (Staudinger) ‘v. Timur | Stgr.’ (on

white paper), handwritten ‘Kuldja | Rkbl. [Rückbeil] [18]84’

(on brown paper), printed ‘Origin. (on pink paper);

e 53,32 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper), one male specimen also has a hand-

written (Staudinger) label ‘v. Timur | Stgr. (on white paper).

LYCAENIDAE

acamas Klug, 1834

‘Lycaena Acamas’ Klug, 1834: Decas Quarta: Signature k(iv), Tab. 40, Figs 7-9. Type

locality: ‘Habitat in Syria et Arabia felici’.

Types: not found (Catalogus Generalis: 4105, 4106 — see

Appendix); probably in The Natural History Museum in London (a

passage “The only specimens I have seen are 1 4 from Klug’s type

material, Butler’s type © of bellatrix from Suakin in the British

Museum, and a cotype of the latter in Mr. Joicey’s collection’

(Riley, 1925: 88) suggests at least one of Klug’s types to be

deposited in (or received on loan by) the British Museum).

Lectotype 2: ‘...females in figs 8 and 9 [of the Symbolae

Physicae] are of “Syrian” origin. In order to avoid upsetting esta-

blished usage it seems best to restrict ssp. acamas to Syrian popu-

lations by designating the female upperside illustrated as the actu-

al type specimen’ (Larsen, 1983: 377; ICZN Art. 74.4).

198

acaudata Staudinger, 1901

‘Thecla Lunulata Ersch. v. Acaudata Stgr. Staudinger, 1901: 70. Type locality:

‘Ferg.[ana]’.

Syntypes 646,59:

e 3 with labels: handwritten (Staudinger) ‘Lunulata var. |

Acaudata Ster. (on white paper) and handwritten ‘Margelan |

Hbhr. [Haberhauer]’ (on brown-pink paper);

e 36, 2 with circles of brown-pink locality label paper, 2 has

also handwritten date label *30/5 [18]80’ (on white paper);

e 6 with handwritten label “Taschkend | [18]91 Schott’ (on

brown paper);

e ?-¢ with handwritten label ‘Alex.[ander] Geb.[irge] | [18]86

Hbhr. [Haberhauer]’ (on brown paper);

e © with labels: handwritten ‘Osch | Hbhr. [Haberhauer] [18]82’

(on brown paper) and date label *26/6’ (on white paper);

e © with circle of brown locality label paper and handwritten

date label ‘18/6’ (on white paper);

e 6 with labels: handwritten ‘Namangan | Hbh. [Haberhauer]

[18]84’ (on brown paper) and date label ‘6/6’ (on white paper);

e © with a circle of brown locality label paper.

Likewise most of the Staudinger species-group taxa becoming

available after his death in 1900 (1. e. in Staudinger, 1901), the type

specimens were not designated with characteristic ‘Origin. labels.

The specimens are provided now with a printed label ‘acaudata

Staudinger, 1901 | SYNTYPUS | Y. Nekrutenko det. 3.11.1998’ (on

yellow paper).

actinides Staudinger, 1886

‘Lyc.[aena] Actis var. Actinides Stgr. Staudinger, 1886: 214. Type locality: ‘Diese ist

bei dem vorliegenden ganz frischen d aus dem südlichen Alai...’

Holotype & (by monotypy):

e 3 with labels: handwritten (Staudinger) ‘Actinides | Ster. (on

white paper), and ‘Alai | [18]85 Hb. [Haberhauer]’ (on brown

199

paper), small new label ‘68’ (on white paper), printed ‘Origin.

(on pink paper) — cf. Häuser & Eckweiler (1997: 59);

Eckweiler & Häuser (1997: Pl. 10).

The specimen is provided with a printed label ‘actinides

Staudinger, 1886 | HOLOTYPUS (par monotypiam) |

Y. Nekrutenko det. 4.11.1998 | cf. Häuser & Eckweiler,

Nachr.ent.Ver. Apollo Suppl. 16: 59 (1997)’ (on red paper).

actis Herrich-Schäffer, [1851]

“[Lycaena] Actis’ Herrich-Schäffer, [1851] 1(49): Tab. 104, Figs 496-499; [1851] 6(55):

24. Type locality: ‘Aus Kleinasien’.

Syntypes d, ¢:

e 3 with labels: handwritten ‘Actis | HSch. f. 496-971 Atys

Kind.[ermann] ms. Gerh.[ard] | t. 19. f. 3. | Amasia

Kind[ermann]’ (recto yellow, in printed box, verso white),

printed ‘4723’ (on white paper);

e 2 without any label.

Catalogus Generalis: 4723 — see Appendix; cf. Hauser &

Eckweiler (1997: 59).

The specimens are provided now with a printed label ‘actis

Herrich-Schäffer, [1851] | SYNTYPUS | Y. Nekrutenko det.

30.11.1998’ (on yellow paper). Subsequently, Olivier (2000a:

105-106) has designated the d specimen as lectotype and the ©

specimen as paralectotype, further restricting the type locality as

‘Turkey, Tokat province, env. Tokat’.

aedon Christoph, 1877

‘Lycaena Aedon Chr. Christoph, 1877: 236, Tab. 5, Fig. 8. Type locality: ‘Ich fing

überhaupt nur 3 4 & davon bei Schahkuh im Juli’.

Syntype d:

e 3 with labels: handwritten ‘Schahkuh | Chr.[istoph]’ (on yellow

paper) and printed ‘Origin.’ (on pink paper).

200

aegagrus Christoph, 1873

‘Lycaena Aegagrus Chr.’ Christoph, 1873: 24, Tab. 1, Figs 3, 4. Type locality: ‘Ich fand

der Schmetterling nur auf einer etwa 10-11,000° hoch gelegenen... muldenartigen

Einsenkung am Schahkuh’.

2-Syntypes d, 9:

e 3, 2 with labels: handwritten (Püngeler) ‘Hyrcania’ (on white

paper, in printed box) and handwritten (Püngeler) on printed

form ‘Cotype <underlined> | v. aegagrus Chr. 3 (®) | von Stgr.

| Püngeler’ (on pink paper). |

The specimens are placed under a handwritten (Püngeler) drawer

label “Pyrenaica B. | v. Aegagrus Chr.’ (on white paper, in printed

box).

aereus Eckweiler, 1998

‘Polyommatus (Agrodiaetus) antidolus subsp. aereus ssp. nov.’ Eckweiler, 1998: 220,

Farbtafel 1. Type locality: ‘Iran, Azarbayjan-e-Gharbi, Takab, 10 km E Takht-e-

Soleyman, 2300 m (Iran, Zagros mts.)’.

Paratype 3:

e 6 with printed labels: ‘Iran, Azarbayjan-e-Gharbi | 10 km E

Takht-e-Soleyman | 2300 m, 24.-25.V11.1998 | leg. Eckweiler

548’ (on white paper, in box) and ‘Paratypus | Polyommatus

(Agrodiaetus) | antidolus aereus subsp. nov. | ECKWEILER 1998’

(on bluish paper, in box).

aeruginosa Staudinger, 1881

‘Lycaena Cyllarus Rott. var. aeruginosa’ Staudinger, 1881: 285. Type locality: ‘Ala

Tau’ .— Junior subjective synonym of Papilio alexis alexis Poda, 1781.

Syntypes 2d, 2:

e <6 with labels: handwritten ‘Saisan | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 6 with labels: handwritten ‘Lepsa | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

e © with circle of brown locality label paper and printed ‘Origin.’

(on pink paper).

201

The specimens are placed under a handwritten drawer label

‘Aéruginosa | Stgr.’ (on white paper, in box).

aestiva Staudinger, 1871

‘Lycaena Astrarche Bgstr. v. gen. Il (m.) Aestiva’ Staudinger, 1871a: 11. Type locali-

ty: not stated. — Junior subjective synonym of Lycaena calida Bellier, 1862. —

Infrasubspecific.

Syntypes 86, 2:

e d with labels: handwritten (Staudinger) ‘Sn Ildefonso | m.[ihi]’

(on blue-green paper), printed ‘Origin.’ (on pink paper);

(on blue-green paper), small date label ‘29/6’, printed ‘Origin.’

(on pink paper);

e 3 with circle of blue-green locality label paper and printed

label ‘Origin.’ (on pink paper);

e 3 with labels: handwritten (Staudinger) ‘Granada | m.[ihi]’ (on

blue-green paper), small date label “6/8”, printed ‘Origin. (on

pink paper);

e © with labels: handwritten (Staudinger) ‘Sier. Nev. | m.[ihi]’

(on blue-green paper), printed ‘Origin. (on pink paper);

e 6 with labels: handwritten ‘Sardinia m.[ihi]’ (on pale pink

paper), printed ‘Origin.’ (on pink paper);

e d with circle of pale pink locality label paper and printed label

‘Origin.’ (on pink paper);

e 3 with labels: handwritten ‘Graecia | Kr.[ueper]’ (on yellow

paper), small square piece of green paper, printed ‘Origin.’ (on

pink paper);

e JS with labels: handwritten ‘Parnass | 24/6 [18]65’ (on white

paper), circle of yellow locality label paper above and printed

‘Origin.’ (on pink paper).

affinis Staudinger, 1892

‘Thecla Affinis Stgr. n. sp.’ Staudinger, 1892b: 148. Type locality: ‘brachte Dörries 3 gute

(franzenreine) anscheinend gefangene 2 © aus dem Sutschan-Gebiet mit...’ — Junior sub-

jective synonym of Thecla eximia Fixsen, 1887 (Staudinger, 1901: 69).

202

Syntypes 39:

e 2 with labels: handwritten ‘Sutschan | [18]90 Dörr.[ies]’ (on

blue-violet paper) and printed ‘Origin. (on pink paper):

e 2? with circles of blue-violet locality label paper, one with

printed ‘Origin.’ (on pink paper).

agnata Staudinger, 1889

‘Lycaena Agnata Stgr (Christophi var.?)’ Staudinger, 1889: 19. Type locality:

‘...sandte mir Herr Tancré 5 d und 2 2 ein, die Riickbeil in den Gebirgen südlich vom

Issyk-Kul fing....’

Syntypes 34, 2°:

e 3 with labels: handwritten (Staudinger) ‘v. Agnata Stgr. |

Rogneda Gr. Gr. (on white paper), handwritten ‘Issyk-Kul |

mer.[idionalis] occ.[identalis] | 18[88] Rckbl. [Rückbeil]’ (on

brown paper), printed ‘Origin.’ (on pink paper);

e 25,2% with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

alaica Staudinger, 1886

‘Lyc.[aena] ... Cytis var. Alaica Stgr. Staudinger, 1886: 206, 207. Type locality:

“...auf dem Alai (bei Margelan)...” — Junior subjective synonym of Lycaena cytis

panaegides Staudinger, 1886 (Staudinger, 1901: 80).

Syntypes 54,49:

e ¢ with labels: handwritten (Staudinger) ‘var. Alaica | Stgr. (on

white paper), “Alai | 85 Hbh. [Haberhauer]’ (on brown paper),

printed ‘Origin.’ (on pink paper);

e 46,49 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper) (figured in Tshikolovets, 1997a: 133,

238, pl. 40, figs 9(11); one male specimen has also labels:

handwritten (Staudinger) ‘Cytis var. Alaica | Stgr.’ with inser-

tion in pencil ‘Alai’ (on white paper), large handwritten

‘v. Panaegides | Stg.! = v. Alaica | Stg. (in pencil, on white

paper).

203

alaina Staudinger, 1887

‘Lycaena Alaina Stgr. Staudinger, 1887a: 50. Type locality: ‘...sandte mir Maurer 10

Stücke aus dem südlichen Alai (Transalai) ein, unter denen nur ein ? ist.’

Syntypes 48, 2:

3S with labels: handwritten (Staudinger) ‘Alaina | Stgr.’ (on

white paper), handwritten ‘Transalai | [18]86 Maur.[er]’ (on

brown paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow

paper), printed ‘Origin. (on pink paper), handwritten “Gen.

Prep. | N 3096 3’;

33, 2 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper) — figured in Tshikolovets (1997b: 40,

figs 12-15);

3d with labels: printed ‘Tura’ (on white paper faded yellowish,

in box), ‘Staudinger’ (on white paper faded yellowish, in box),

handwritten (Püngeler) ‘alaina d, | Cotypus <underlined> |

11/09 ex coll. | Dieckmann’ (on white paper) and handwritten

(Piingeler) on printed form ‘Cotype <underlined> | alaina Stgr.

3 | Püngeler’ (on pink paper).

The specimens are placed under handwritten (Piingeler) drawer

label ‘Alaina Stgr.’ (on white paper, in box).

albicans Gerhard, [1851]

‘Lycaena Corydon Hbr.{Hiibner] Var. Albicans Hbr.’ Gerhard, [1851] (8): 17, Pl. 31,

Fig.

3 a-d. Type locality: ‘Spanien’ ... ‘Auch diese Var. stammt aus Spanien... Den

Mann besitze ich, das Weibchen erhielt ich durch Herrn Lederer.’ — Junior primary

homonym of Lycaena dorylas albicans Gerhard, [1851].

Neotype à:

204

d with labels: handwritten ‘Granada | m.[ihi] [18]80’ (on pale

greenish paper), ‘albicans’ (on white paper, in ball pencil),

printed ‘Zool. Mus. | Berlin’ (on pale yellow paper), handwrit-

ten (Schurian) on printed lined form ‘Polyommatus | (Lysandra)

| albicans | GERHARD 1851 | det. Schurian 1988’ (on white

paper), handwritten (Schurian) on printed form ‘Gen. Praparat |

Nr. 208/1984 | Schurian’, handwritten (Schurian) on printed

form ‘Neotypus à | Lysandra albicans | H.-S. (1852) | teste

SCHURIAN 1984’ (on red paper), and ‘Neotypus 3 <written

over printed ‘Paratypus’> | Lycaena | corydon albicans | GER-

HARD 1851 | det. Schurian, 1988” (on red paper, in box) —

Schurian (1988: 134, Taf. 1, 2, Fig. 3).

albicans Herrich-Schäffer, [1851]

‘[Lycaena] Corydon var. albicans’ Herrich-Schäffer, [1851] 1(47): Tab. 103, Figs 494,

495; [Lycaena] Corydon Zell. [Lokalvarietät] Albicans [1852] 6(55): 27. Type locali-

ty: ‘Aus Spanien.’ — Junior primary homonym and junior subjective synonym of Lycaena

corydon albicans Gerhard, [1851]. The priority of Gerhard’s name over that of Herrich-

Schäffer’s [1851] is established here after Kocak, 1980c: 17 (cf. ICZN Art. 21.3.1,

232):

Types: not found.

alexandra Püngeler, 1901

“Chrysophanus athamantis Ev. var. nov. alexandra’ Püngeler, 1901: 179, Taf. 2, Fig. 6, 6a,

6b. Type locality: ‘...in Anzahl aus dem Alexandergebirge’.

Lectotype d:

e d (the end of abdomen removed, genitalia in a plastic tube)

with labels: printed, with handwritten (Püngeler) insertion ‘Asia

centr.[alis) | Alexandergebirge | Rückbeil 1899 (recto, in box) |

athamantis var. | alexandra Püng. 3 | Original <double under-

lined> Oberseite | abgebildet Iris 1901 | (v. R. Tancre 1.01)

(verso) (in white paper), printed “Type’ <double underlined> |

with handwritten (Püngeler) insertion ‘v. alexandra Püng. à |

Püngeler’ (on pink paper), printed “Zool. Mus. Berlin’ (on pale

yellow paper), handwritten (Nekrutenko on printed form

‘Figurat | par Y. Nekrutenko in: | Vestn. zool., 1984, 6: | tab. ex

adv. p. 48° (in white paper, in box), handwritten (Nekrutenko)

‘alexandra Püng. | Lectotypus 4 | Y. Nekrutenko design. |

13.X.1983’ (on yellow paper) — Nekrutenko (1984: 44, Pl. [1];

1983730, fig: 2; a, b).

Paralectotypes ¢, 29:

e 3,2% with labels: printed, with handwritten (Püngeler) inser-

tion ‘Asia centr.[alis] | Alexandergebirge | Rückbeil 1899’ (in

white paper, in box) and handwritten (Püngeler) on printed

205

form ‘Cotype <underlined> | w alexandra Püng. & (®) |

Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler)

drawer label ‘Athamantis Ev. | v. Alexandra Püng.’ (on white paper,

in box).

alpina Staudinger, 1871

‘Lycaena Medon Hufn. (Agestis S. V.) var Alpina’ Staudinger, 1871b: 52. Type locali-

ty: not stated [Parnass?] — see also Staudinger, 1901: 83 for synonymy with ‘ab. (et v.)

allous Hb..

Types: not found; apparently they have never been designated.

alticola Christoph, 1893

‘Lycaena Coelestina Ev. var. Alticola’ Christoph, 1893: 86. Type locality: ‘Urmus,

Kasikoparan. Armeniae rossicae.’

Paralectotypes 4 d :

e 6 with handwritten (Staudinger) labels ‘Coelestina var. |

Alticola | Christh.<sic>’ (on white paper) and ‘Urmus

Cauc.[asus] m.[eridionalis] | Chr.[istoph] [18]83’ (on green

paper);

e 36d with circles of green locality label paper; one specimen has

also a printed label ‘Zool. Mus. | Berlin’ (on pale yellow paper)

and handwritten (Wagener) ‘Lycaena | alticola Chr. | Urmus

Cauc. m. | =Neolysandra | pontica Courv.’ (on white paper), the

second has a handwritten (Staudinger) label ‘Urmus’ (on green

paper), the third — handwritten (Christoph?) ‘L. Coelestina v.

Alticola Chr.[istoph] | 1. 1.

Lectotype designated by Nekrutenko in Hesselbarth et al., 1995:

652 (deposited in Zoological Institute, Russian Academy of

Sciences, St.-Petersburg, Russia). Since the forementioned speci-

mens are undoubtedly an integral part of Christoph’s type series

forwarded to Staudinger, they are provided now as paralectotypes

with a label ‘coelestina alticola | Christoph, 1893 | PARALECTO-

TYPUS | Y. Nekrutenko det. 11.11.1998’ (on yellow paper) —

ICZN Art. 74.1.3, Recommendation 74F.

206

amasina Neuburger, 1900

‘Lycaena menalcas Frr. 3 ab. amasina’ Neuburger, 1900: 370. Type locality: ‘Aus der

Gegend von Amasia erhielt ich eine bedeutende Anzahl Lycaena menalcas Frr. Unter

ihnen befand sich ein d, welches abweichende Merkmale tragt.’— Infrasubspecific.

Holotype 4 (by monotypy):

e 3 with labels: handwritten (Neuburger?) ‘Lycaena menalcas |

ab. amasina | Neubgr. Amasia’ (on white paper, in printed box),

printed ‘Type’ (on pink paper), handwritten (Forster) on printed

form ‘Agrodiaetus | dolus 6 | menalcas Frr. | det W. Forster

1948’ (on white paper) — cf. Hauser & Eckweiler (1997: 62).

The specimen is placed under a handwritten drawer label

“menalcas | ab. amasina | Neubgr.’ (on white cross-lined notebook

paper).

amor Staudinger, 1886

‘Lyc.[aena] Eros Ochs. var. Amor Stgr.’ Staudinger, 1886: 211. Type locality: ‘Ich

erhielt diese Lokalform von Eros in Anzahl aus dem Alai (siidlich von Margelan gelege-

nen Gebirgen), von Osch, und von den Gebirgen bei Samarkand.’ — Junior primary

homonym and junior objective synonym of Lycaena eros amor Lang, 1884: 371.

Lectotype 3:

e JS with labels: circle of brown paper, printed ‘Origin.’ (on pink

paper), handwritten (Lukhtanov) on printed form ‘Lectotypus

3 | amor Stgr. design. V. Lukhtanov’ (on white paper, recto red,

verso white) — Lukhtanov & Lukhtanov (1994: 272).

Paralectotypes 11d, 109:

e <6 with labels: handwritten (Staudinger) ‘Eros v. Amor | Stgr.’

(on white paper), and ‘Prov. Samark.[and] | Hbhr. [Haberhauer]

[18]81’ (on brown paper), small date label ‘6/7’ (on white

paper), printed ‘Origin. (on pink paper), printed ‘Zool. Mus. |

Berlin’ (on pale yellow paper), handwritten ‘Polyommatus |

amor Stgr. | typus (recto) | Gen. Prep. N 3092 & (verso)’; (on

white paper, in printed box);

e 6 with labels: handwritten (Staudinger) ‘circa Napaea à |

GrGr’, a circle of brown locality label paper, small date label

‘6/7’ (on white paper), and printed ‘Origin.’ (on pink paper);

207

e 6 with labels: handwritten (Staudinger) ‘oben genau wie

Erigone | Gr.-Gr. | die unten auf Htfl. | deutlicheren weissen

Langsst[rich?] u. rothen Randaugen hat’ (on white lined note-

book paper), circle of brown locality label paper, small date la-

bel ‘6/7’ (on white paper), and printed ‘Origin.’ (on pink paper);

e 3 with labels: handwritten ‘Margelan | Hbhr. [Haberhauer]’,

small date label ‘2/7’ (on white paper), and printed ‘Origin. (on

pink paper);

e 39° with circles of brown locality label paper and printed label

‘Origin.’ (on pink paper);

e d with labels: handwritten (Staudinger) ‘Eros v. Amor | Stgr.’

(on white paper), handwritten label ‘Alaı | [18]85 Hb.

[Haberhauer]’, and printed label ‘Origin. (on pink paper);

e 6, 3% with circles of brown locality label paper and printed

label ‘Origin. (on pink paper); one female specimen has a

handwritten label ‘Lyc.[aena] Eros | 3000 m’;

e ¢ with labels: handwritten (Staudinger) ‘Eros v. Amor | Stgr.

(on white paper), ‘Margelan | Maur.[er] [18]84’, and printed

label ‘Origin.’ (on pink paper);

e 3, 3% with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper); the male specimen also has a

handwritten date label ‘27/6’ (on white paper) and a handwrit-

ten label ‘Kara Karek’ (on white paper);

e 3 with labels: handwritten ‘Osch | Hbhr. [Haberhauer]’ and

printed ‘Origin. (on pink paper);

e 24, 2 with circles of brown locality label paper and printed

label ‘Origin. (on pink paper); one male specimen also has a

small date label ‘19/7’ (on dirty white paper).

amurensis Staudinger, 1892

‘Polyommatus Hippothoé L. var. Amurensis Stgr. Staudinger, 1892b: 155. Type locali-

ty: ‘Die mir vorliegenden 3 Pärchen vom Uss.[uri] (1 2 von Radd.[efka]) berechtigen...’

Syntypes>o sr

e 3 with labels: handwritten (Staudinger) ‘v. Amurensis | Stgr.’

(on white paper), handwritten ‘Sutschan | [18]90 Dörr.[ies]’ (on

bluish-violet paper), printed ‘Origin.’ (on pink paper);

208

e 3, with circles of bluish-violet locality label paper and print-

ed label “Origin. (on pink paper);

e G with labels: handwritten ‘Ussuri | Dörr.[ies] [18[82]’ (on

bluish-violet paper) and printed ‘Origin.’ (on pink paper);

e © with circle of bluish-violet locality label paper and printed

label ‘Origin.’ (on pink paper);

e 2 with labels: handwritten ‘Raddefskaja | Chr.[istoph]’ (on

bluish-violet paper), small handwritten labels ‘14/7 [18]76’ and

‘11/7’ (on white paper) and printed ‘Origin.’ (on pink paper).

amurensis Staudinger, 1892

‘Lycaena Amanda Schn. var. Amurensis Stgr. Staudinger, 1892b: 162. Type locality:

“Von Radde im Bur.[eja]-G.[ebirge], von Maack am Uss.[uri] gefangen. Christoph fand

diese Art im Juli bei Radd.[efka] nicht selten; Dörries sandte sie mir von Ask.[old],

Baran.[owka], vom Sutschan und vom Uss.[uri]; Graeser fand sie bei Chab.[arowka] sel-

ten, bei Pokr.[ofka] ziemlich häufig’.

Syntypes 54, 9:

e d with labels: handwritten (Staudinger) ‘v. Amurensis | Stgr.’

(on white paper), and ‘Raddefskaja | Chr.[istoph] [18184 (on

violet paper), printed ‘Origin.’ (on pink paper);

e 6, with circles of violet locality label paper and printed label

“Origin. (on pink paper);

e d with labels: handwritten ‘Ussuri | Dörr.[ies] [18]82’ (on vio-

let paper), printed ‘Origin.’ (on pink paper);

e d with circles of violet locality label paper and printed label

‘Origin.’ (on pink paper);

e 3 with labels: handwritten ‘Baranowsky | Dörr.[ies]’ (on violet

paper), printed ‘Origin.’ (on pink paper).

Another d, 2 from ‘Sutschan | [18]90 Dorr[ies]’ and 6, © from

‘Askold | Dör.[ries]’ are not designated as ‘Origin.’ though they fit

all requirements to be syntypes.

ankara Schurian & Hofmann, 1983

‘Lysandra ankara sp. nov.’ Schurian & Hofmann, 1983: 121. Type locality (holotype):

‘Anatolia/Ankara, vic. Kizilcahamam, 5 km südlich, 1100-1400 m...’

209

Paratype à:

e d with labels: printed, with handwritten (Schurian) insertion

‘ANATOLIA/Ankara | vic. Kızılcahamam | 5km <sic>

südl./nördl. <sic> | 1100-1400 m NN | leg. Schurian |

30.VII.[19]77’ (on white paper) and handwritten (Schurian) on

printed form ‘PARATYPUS 4 | Lysandra | ankara sp. n. | det.

Schurian/Hofmann 1983’ (on red paper, in box).

antidolus Rebel, 1901

‘Lycaena Dolus Hb. var. Antidolus n. var. Rebel, 1901: 167. Type locality: ‘...am

Takaltu Dagh bei Kulp (Russ.[isch]-Südarmenien) um den 15. Juli 1901...’ ‘Die Typen

befinden sich im k. k. naturhistorischen Hofmuseum und in der Sammlung des Herrn

Bohatsch’.

Paralectotype à:

e 3 with labels: handwritten ‘Lyc.[aena] dolus Hb. | v. Antidolus

Rbl. <in black ink> | Type 1902 <in red ink>’ (in white paper),

handwritten (Püngeler) ‘Armenia | (Kasikoparan) |

Gebirgsthäler, Korb 15.7.[19]01 (recto) | v. antidolus Rbl. à |

Type <double underlined> ex coll. Bohatsch | 12/1912 M. Korb

Püngeler (verso)’ (on white paper), handwritten (Püngeler) on

printed form “Type <double underlined> | v. antidolus Rbl. 3”

(on pink paper), handwritten (Forster) on printed form

‘Agrodiaetus 3 | antidolus Rebel | det. W. Forster 1948’ (on

white paper) handwritten (Forster) on printed form ‘Para-Typus

3 | Lycaena dolus | antidolus Rebel’ (on pink paper) — Forster

(1961: 111); cf. Hauser & Eckweiler (1997: 63).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Dolus Hb. | v. Antidolus Rbl.’ (on white paper, in printed

box).

antiochena Lederer, 1861

‘Lycaena antiochena m. Lederer, 1861: 148, Taf. 1, Figs 2, 3. Type locality: ‘Bei

Antiochia. ...erhielt ich nur 4 d und 3 2’.

Syntypes 6, 9:

e d, ? with labels: handwritten ‘Antiochia’ (on white paper),

printed ‘Coll. Led.’ (on white paper) and printed ‘Origin.’ (on

pink paper).

210

The specimens are placed under a handwritten drawer label

‘Antiochena | Ld.’ (on white paper, in box).

antiqua Staudinger, 1900

‘Lyclaena] Chiron Hufn. var. (Antiqua)’ Staudinger, 1900: 334. Type locality: ‘aus

dem östlichen Thian Schan-Gebiet’.

Syntypes 94, 59:

e 3 with labels: handwritten (Staudinger) ‘v. Antiqua’ (on white

paper), and “Thian or.[ientalis] [18]96 | Hbh. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

e 3 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper);

e ?-¢ with labels: handwritten (Staudinger) ‘v. Antiqua | Stgr. in

1. | (recto) | v. Cogistana ? (verso)’ (on white paper) and ‘Usgent

| Hbh. [Haberhauer] [18]83’ (on brown paper);

e ?-G with handwritten label ‘Osch | Hbh. [Haberhauer] [18]80’

(on brown paper);

e ?-G with handwritten label ‘Alai | [18]85 Hbh. [Haberhauer]’

(on brown paper);

e ?-2¢ with circles of brown locality label paper:

e ?-2 with handwritten label ‘Namangan | 18[84] Hbhr.

[Haberhauer]’ (on brown paper).

e ?-G, 2 with labels: printed ‘Korla’ (on white paper), the male

specimen has a handwritten (Staudinger) label ‘Chiron v. |

Antiqua’ (on white paper);

e ?-¢ with labels: handwritten (Staudinger) ‘v. Antiqua | Ster.

(on white paper), and ‘Alex.[ander] Geb.[irge] | [18]86 Hbh.

[Haberhauer]’ (on brown paper);

e ?-? with circles of brown locality label paper:

e ?-G with handwritten label ‘Prov. Samark.[and] | Hbhr.

[Haberhauer] [18]81’ (on brown paper);

e ?-? with circles of brown locality label paper;

Of these specimens, only two males are designated as

‘Origin.[ale]’ though they all originated from Central Asia and

211

hence may belong to the type series and should be considered syn-

types: ‘...wie andere mir vorliegende centralasiatische aus dem

Fergana-Gebiet... Wir versandten die centralasiatischen Stücke als

var. Antiqua. (op. cit.: 334).

aquilina Staudinger, 1901

‘Lycaena F. Orbitulus Prun. v. Aquilina Stgr. Staudinger, 1901: 81. Type locality:

‘Lap.[ponia]’.

Syntypes 66,49:

e 3 with handwritten label ‘Lapponia | Keit.[el]’ (on pale bluish

paper):

e 24,29 with circles of pale bluish locality label paper;

e 6 with handwritten (Staudinger) label ‘Finmarken | Schm.[?]’

(on white paper);

e 24,29 with circles of white locality label paper.

Likewise most of the Staudinger species-group taxa becoming

available after his death in 1900 (i. e. in Staudinger, 1901), the type

specimens were not designated with characteristic ‘Origin.’ labels.

Since there are no other specimens determined as aquilina Stgr. in

the collection, and since the specimens above are placed under the

handwritten drawer label ‘v. Aquilina Stgr. | Aquila Auriv.[illius]’

(on white paper, unknown hand, in pencil) and because of the same

type locality stated on the label and in the original description, they

are accepted here with confidence as syntypes and provided with a

printed label ‘aguilina Staudinger, 1901 | SYNTYPUS |

Y. Nekrutenko det. 3.11.1998’ (on yellow paper).

araxiana Kocak, 1980

‘Agrodiaetus (Sublysandra) myrrhus araxiana n. subsp.’ Koçak, 1980b: 266, Figs 6-9.

Type locality: “Türkei, Kars, Akçay 1500-1700 m’.

Paratypes 3 d:

e 34 with labels: printed ‘Tiirkei/Kars | Akçay | 1500-1700 m |

22.7.[19]77 | leg. Eckweiler’ (on white paper), handwritten

(Koçak) on printed form ‘PARATYPUS <4 | Agrodiaetus |

(Sublysandra) | myrrha araxiana | KOCAK 1979 (on red

paper) — cf. Häuser & Eckweiler (1997: 109).

212

ardschira Brandt, 1938

‘Lycaena ardschira sp. n.’ Brandt, 1938: 672, Taf. 4, Figs 6-8. Type locality: Iran,

‘Comèe, 2600 m’.

Paratype d:

e 6 with labels: printed ‘Iran Fars | Straße Ardekan - Talochsroe

| Comèe ca 2600 m 1937 | coll. Brandt’ (on yellow paper), print-

ed ‘61.’ (on white faded paper), printed ‘Paratypus’ (on red

paper) and handwritten ‘Lycaena | ardschira B.[randt]’ (on

white paper) — cf. Häuser & Eckweiler (1997: 63).

argiva Staudinger, 1886

‘Lyc.[aena] Argiva Stgr. (Argus var.?)’ Staudinger, 1886: 204. Type locality: ‘Von

dieser Art (oder Localform) besitze ich 3 d and 1 © aus dem Alai, 2 d und 1 © aus

‘Margelan’, die auch wohl aus dem südlich davon gelegenen Alai-Gebirge stammen, und

ein d, welches am 10. Mai bei Namangan gefunden wurde’.

Syntypes 68,3%:

e d with labels: handwritten (Staudinger) ‘Argiva Ster. (on white

paper), handwritten ‘Alai | [18]85 Hbh.[Haberhauer]’ (on brown

paper), printed ‘Origin. (on pink paper);

e 6 with labels: handwritten ‘Alai | [18]85 Hbh.[Haberhauer]’

(on brown paper) and printed “Origin. (on pink paper);

e 6, 2 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper);

e 26,29 with labels: handwritten ‘Margelan | Maur.[er] [18]84’

(on brown paper) and printed ‘Origin.’ (on pink paper); one d

and one 2 with handwritten (Staudinger) label ‘wie Roxane d

(2) Gr. Gr.’ (on white paper);

e G with labels: handwritten (Staudinger) ‘Argus var.? | Argiva

Ster. (on white paper), handwritten ‘Namangan | Hbh.[Haber-

hauer] [18]84’ (on brown paper), printed ‘Origin. (on pink

paper).

arionides Staudinger, 1887

‘Lycaena Arionides Stgr. Staudinger, 1887b: 141, Taf. 7, Fig. 1, a-c. Type locality:

‘Christoph fing diese schöne Art Ende Juli, Anfang August bei Wladiwostok; Dörries

sandte sie mir später noch in einiger Anzahl vom Ussuri ein.’

215

Syntypes 94, 59:

e 6 with labels: handwritten (Staudinger) ‘Arionides | Stgr.’ (on

white paper), handwritten ‘Ussuri | Dörr.[ies]’ (on violet-blue

paper) and printed ‘Origin. (on pink paper); |

e 3,2% with circles of violet-blue locality label paper and print-

ed ‘Origin.’ (on pink paper);

e d with labels: handwritten “Wladiwostock | Chr.[istoph]’ (on

violet-blue paper), small date label ‘9/8’ (recto white, verso red)

and printed ‘Origin. (on pink paper);

e 64,39 with circles of violet-blue locality label paper, some

with small date labels as above, and printed “Origin. (on pink

paper). |

armena Rebel, 1901

‘Thecla Myrtale Klug var. Armena n. var. d 9’ Rebel, 1901: 165. Type locality:

‘...anfangs Juli 1901 in einem Flussthale bei Kasikoparan (Russ.[isch]-Armenien, südlich

von Kars gelegen)...” ‘Mir liegen zur Beschreibung zwei Pärchen vor: eines aus der

Sammlung des k. k. naturhistorischen Hofmuseum, das andere aus jener des Herrn

Bohatsch’.

Syntypes d, 9:

e G, % with labels: printed ‘Russ.[ische] Armen.[ien] |

Kasikoparan | 1901 Korb’ (on white paper) and handwritten

(Püngeler) on printed form ‘Cotype <underlined> | v. armena

Rbl. 3 (8) |von M. Korb | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Myrtale Klug | v. Armena Rbl’ (on white paper, in printed

box).

armena Staudinger, 1871

‘Lycaena Hylas Esp. v. Armena’ Staudinger, 1871a: 12. Type locality: ‘Arm.[enia]’.

Lectotype à:

e d with labels: handwritten ‘Achalzich’ (on white faded paper),

printed ‘Coll. Led.’ (on white paper) and ‘Origin. (on pink

paper), ‘Zool. Mus. | Berlin’ (on pale yellow paper), printed,

214

with handwritten (Wagener, in pencil) insertions ‘Abgebildet in

Hesselbarth, | van Oorschot & Wagener: | Tagfalter der Türkei |

Tafel 706 Figur 41’ (on white paper), handwritten (Nekrutenko)

on printed form ‘LECTOTYPUS & | armena Stgr. |

Y. Nekrutenko design.’ (on yellow paper) — Nekrutenko in

Hesselbarth et al. (1995: 658, Taf. 106, Fig. 33).

Paralectotypes, 2d, 9:

e ¢ with labels: handwritten ‘Achalzich’ (on white faded paper),

printed ‘Coll. Led.’ (on white paper) and ‘Origin.’ (on pink

paper), “Zool. Mus. | Berlin’ (on pale yellow paper), handwrit-

ten (Nekrutenko) on printed form ‘PARALECTOTYPUS 4 |

armena Stgr. | Y. Nekrutenko design. (on yellow paper);

e à with labels: handwritten “Caucasus | Hbhr. [Haberhauer]’ (on

green paper) and printed “Origin. (on pink paper), handwritten

(Nekrutenko) on printed form ‘PARALECTOTYPUS à | arme-

na Ster. | Y. Nekrutenko design.’ (on yellow paper);

e © with labels: handwritten ‘Achalzich’ (on white faded paper),

printed ‘Coll. Led.’ (on white paper) and ‘Origin.’ (on pink

paper), “Zool. Mus. | Berlin’ (on pale yellow paper), handwrit-

ten (Nekrutenko) on printed form ‘PARALECTOTYPUS &

(2) | armena Stgr. | Y. Nekrutenko design.’ (on yellow paper),

printed, with handwritten (Wagener, in pencil) insertions

‘Abgebildet in Hesselbarth, | van Oorschot & Wagener: |

Tagfalter der Türkei | Tafel 706 Figur 40° (on white paper).

armeniaca A. Bang-Haas, 1906

“Chrysophanus virgaureae v. armeniaca n. var. Bang-Haas, A. 1906: 128. Type locali-

ty: ‘Eine Anzahl dd aus Armenien (Tschorum).... — Junior subjective synonym of

Papilio virgaureae virgaureae Linnaeus, 1758 (Hesselbarth er al., 1995: 470).

Lectotype &:

e 6 with labels: handwritten (A. Bang-Haas?) ‘Chrys.[ophanus]

Virg.[aureae] | v. Armeniaca | B-H | Tschorum Armen.[ien]’ (on

white paper) and printed ‘Origin. (on pink paper). Designated

with a handwritten label on printed form ‘'LECTOTYPUS |

armeniaca A. B.-H. | Y. Nekrutenko design.’ (on yellow paper)

— Nekrutenko in Hesselbarth er al. (1995: 470).

215

Paralectotype CE

e J with printed label ‘Origin.’ (on pink paper). Designated with

a handwritten label on printed form ‘PARALECTOTYPUS 4 |

armeniaca A. B.-H. | Y. Nekrutenko design.’ (on yellow paper).

arsacia Lederer, 1869

‘Lycaena Panagaea H. Sch. Var. Arsacia Led. Lederer, 1869: 78, Taf. 4, Fig. 9). Type

locality: ‘...bei Astrabad in Persien... Um Astrabad fand er [Haberhauer] nur wenige

Lepidopteren, dagegen zeigten sich Hadschyabad und Schaku, 120 Werste davon entfernt,

reich daran’ (op. cit.: 73). — Junior subjective synonym of Lycaena anisophthalma

Kollar, 1849 (Staudinger, 1901: 80).

Syntypes 36d, @:

e 3d with labels: handwritten ‘Lycaena | Arsacia’, ‘Astrabad’

(both on white paper) and printed ‘Origin.’ (on pink paper);

e 3, % with labels: handwritten ‘Hadschyabad’ (on white paper)

and printed ‘Coll. Led.’ (on white paper);

e & with printed label ‘Coll. Led.’ (on white paper).

The specimens are provided now with a printed label: ‘arsacia |

Lederer, 1869 | SYNTYPUS | Y. Nekrutenko det. 3.11.1998’ (on

yellow paper).

asabinus Herrich-Schäffer, [1851]

‘[Polyommatus] Asabinus’ Herrich-Schäffer, [1851] 1(49): Tab. 109, Figs 527, 528;

‘Helius Kad. [1852] 6(55): 32. Type locality: ‘Aus Kleinasien.

Types: not found.

astraea Freyer, [1851]

‘Lycaena Pap.[ilio] Astraea’ Freyer, [1851] 6(98): 161, Tab. 583, Figs 1,2. Type locali-

ty: ‘Von Herrn Kindermann unter obigen Namen mir zugesendet. Bei Amasia’.

Syntypes d, 9:

e 6 with labels: handwritten ‘Astraea | Frey.[er] h.<n.[euere]?>

Beitr.[äge] t. 583 | Led. | Amasia Kind.[ermann]’ (recto yellow,

in printed box, verso white), printed ‘4679’ (on white paper);

e © without any label.

216

Catalogus Generalis: 4679 — see Appendix.

The specimens are provided now with a printed label ‘astraea

Reyer | 1851] I SYNTYPUS | Y. Nekrutenko det. 7.12.1998’ (on

yellow paper).

aurantiaca Staudinger, 1901

‘Thestor Hb. Nogelii HS. v. (et ab.) Aurantiaca Stgr. Staudinger, 1901: 72. Type locali-

ty: ‘As.[ia] min.[or] (Kaisarie); Ant.[iochia] (Aintab)’.— Junior subjective synonym of

Thestor nesimachus Oberthür, 1893.

Syntypes 68,2%:

e ¢ with labels: handwritten (Staudinger) ‘v. Aurantiaca | Stgr.’

(on white paper, glued to a larger piece of paper and then pinned

to the specimen), handwritten ‘Kaiserie <sic> | [18]86

Man.[issadijan]’ (on yellow paper) and small handwritten date

label ‘1/6’ (on white paper);

e © with a circle of yellow locality label paper, small date label

‘15/5’ (on yellowish paper), printed ‘Zool. Mus. | Berlin’ (on

pale yellow paper), printed ‘Paratypus’ (on red paper), hand-

written ‘8.6 Mann | Kaiserie’ (on white paper), handwritten

‘Thestor nogelii | aurantiaca Stgr.’ (on white paper) and printed,

with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türkei. |

Tafel 92 Figur 50’ (on white paper);

e 3 (abdomen removed), with labels: printed “Zool. Mus. |

Berlin’ (on pale yellow paper), printed ‘Paratypus’ (on red

paper), handwritten ‘Kaiserie’ (on white paper), handwritten

‘Thestor nogelii | aurantiaca Stgr.’ (on white paper) and printed,

with handwritten insertion‘Genit.-Unters. | Nr. 202 | Zool. Mus.

Berlin’ (on white paper);

e 43, 2 with a circle of yellow locality label paper, three male

specimens also have small date labels ‘12/5’ and ‘29/5’ (on yel-

lowish paper) and ‘1/6’ (on white paper); last one has also

labels: printed “Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Wagener) ‘Thestor nogelii | aurantiaca Ster. |

[18]’86 Mann. Kaiserie’ (on white paper) and printed, with

handwritten (Wagener) insertions ‘Abgebildet in Hesselbarth |

DAT

van Oorschot & Wagener: | Tagfalter der Türkei. | Tafel 92 Figur

36’ (on white paper).

aureomicans Heyne, 1897

‘Polyommatus virgaureae L. nov. var. aureomicans’ Heyne, 1897: 9. Type locality:

‘Fluggebiet: Mersina (südliches Kleinasien, Taurus)’.

Syntypes d , 2:

d with labels: handwritten “Hadjin’ (on white paper), hand-

written ‘7/95’ (on white paper), handwritten (Püngeler) ‘als

aureomicans | d 5/1912 v.[om] Bartel | Piing.[eler]’ (on white

paper) and handwritten (Püngeler) on printed form ‘Cotype

<underlined> | v. aureomicans Heyne & | Püngeler’ (on pink

paper);

? with labels: handwritten (Püngeler) ‘Taurus | (Mersina) |

Heyne (recto) | v. aureomicans © | Cotype <underlined> | ex

coll. Bernard | 3.[19]09 v.[om] M. Bartel (verso)’ (on white

paper) and handwritten (Püngeler) on printed form “Cotype

<underlined> | v. aureomicans Heyne % | Püngeler’ (on pink

paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Virgaureae L. | v. Aureomicans Heyne’ (on white paper, in

printed box).

bella Herrich-Schäffer, [1844]

‘Lycaena Bella’ Herrich-Schäffer, [1844] 1(7): Tab. 49, Figs 227, 228; [1845] 1(10): 127.

Typ

e locality: ‘Ein Mannchen erhielt Hr. Bischoff aus der Gegend von Brussa’.—

Junior subjective synonym of Papilio argus aegidion Meisner, 1818. Infrasubspecific.

Types: not found.

bellus Kardakoff, 1928

‘Zep

Typ

hyrus taxila Brem. f. 3 bellus (nom. coll.)’ Kardakoff, 1928: 271, Taf. 7, Fig. 15.

e locality: ‘...aus dem Ussurigebiet...’ — Infrasubspecific.

Syntype d:

218

& with labels: printed, with handwritten (Kardakoff?) insertion

‘Narva, S. Ussurigebiet | 5.8. [19]27. N. Kardakoff’ (on white

paper), handwritten (Kardakoff?) ‘Zephyrus taxila Brem L. | f.

6 bellus Kardakoff” (on white paper) and printed ‘Typus’ (on

red paper, in box).

The specimens are placed under a typewritten drawer label ‘f. 4

bellus’ <underlined with red pencil> (on a stripe of white paper, in

blue type ribbon).

berber Le Cerf, 1932

‘Lycaena coridon Poda ssp. berber, nova’ Le Cerf, 1932: 163. Type locality: “Types :

3d et 22 (1 H.[olo-] T.[ype]), Maroc, Moyen Atlas: Daya Chikker, Ksar el Kebir,

Timesmount, Tameghilt, Moussah ou Salah, de 1.500 à 3.000 m., 10 au 26-VIII-1930. Le

male «Holotype» est de Ksar el Kebir.’

Paratypes d, 9:

e G with labels: handwritten (Le Cerf?) on printed form

‘Lyc.[aena] coridon | ssp. berber | 3 Cotype Le Cerf | Coll. F.

Le Cerf’, handwritten (Le Cerf?) ‘Maroc | Moyen Atlas | Ksar

el Kebir | 20/25-VIII-1930 | F. Le Cerf’ and printed ‘'COTYPE’

(on white paper, in red ink);

e © with labels: handwritten (Le Cerf?) on printed form

‘Lyc.[aena] coridon | ssp. berber | $ Cotype Le Cerf | Coll. F.

Le Cerf’, handwritten (Le Cerf?) ‘Maroc | Moyen Atlas | Ksar

el Kebir | 20/25-VIII-1930 | F. Le Cerf’ and printed ‘COTY PE’

(on white paper, in red ink).

betulina Staudinger, 1887

‘Thecla Betulina’ Staudinger, 1887b: 127, Taf.16, Fig. 6. Type locality: ‘...nur zwei

Männchen, welche von Dörries am Suifun, an Apfelbäumen fliegend, gefangen wurden’.

Syntypes 26:

e 6 with labels: handwritten (Staudinger) ‘Betulina | Stgr.’ (on

white paper), handwritten ‘Suifun | [18]85 Dörr.[ies]’ (on vio-

let-blue paper) and printed ‘Origin. (on pink paper);

e d with circle of violet-blue locality label paper, handwritten

label ‘474. and printed ‘Origin.’ (on pink paper).

219

birunii Eckweiler & ten Hagen, 1998

‘Polyommatus (Agrodiaetus) posthumus birunii subspec. nov. Eckweiler & ten Hagen,

1998: 116, Abb. 13-16. Type locality (holotype): ‘Iran, Tehran, Elburs-Gebirge 15 km

E Firuzkuh (Paß), 2400 m’.

Paratype d :

e 3 with printed labels: ‘Iran, Tehran | Elburs-Geb. 15 km E |

Firuzkuh (Paß) | 2300-2400 m | 18.VII.1997 | leg. W. ten

Hagen’ (on white paper). Designated with a printed label

‘Paratypus | Polyommatus (Agrodiaetus) | posthumus birunii

subspec. nov. | des. ECKWEILER & TEN HAGEN 1998’ (on bluish

paper, in box).

bitonides Staudinger, 1871

‘Lycaena Biton Brem. (an ‘Bitonides’ nominandus?)’ Staudinger, 1871a: 13. Type

locality: ‘Daür. [Dahuria]; Amur’ (see also Staudinger, 1892b: 163). — nomen nudum.

Types: not found.

boisduvalii Herrich-Schäffer, [1843]

‘Lycaena Boisduvalii’ Herrich-Schäffer, [1843] 1(1): Tab. 2, Figs 7-9; [1844] 1(9): 121.

Type locality: ‘Aus dem südlichen Russland’ — Junior subjective synonym of Lycaena

eroides Frivaldszky, 1835.

Types: not found.

borussia Dadd, 1909

‘Lycaena corydon borussia’ Dadd, 1909a: (1xii1); 1909b: (36). Type locality: ‘...aform

from Berlin... The same form is reported by Herr Zobel from Osterode, East Prussia...’

Neotype d:

e 3 with labels: handwritten ‘Nieder- | finow [bei Eberswalde,

DDR], 22.7.1982, | [leg.] Dr. Kames’ (on white paper, in pen-

cil), handwritten (Schurian) ‘Polyommatus (Ly- | sandra) cori-

don | borussia Dapp (1909) | det. SCHURIAN 1989’ (on white

paper), on printed form with handwritten (Schurian) insertion

‘Neo- Typus | d’ (recto red, verso white), handwritten

(Schurian) ‘Neotypus & | Lycaena corydon | borussia Dadd

1909 | design. Schurian, 1989’ (recto red, verso white) —

Schurian (1989: 305, Abb. 1).

220

brevicaudis Vorbrodt, 1912

‘Thecla F. linceus Esp. (=spini Schiff.) brevicaudis Püngeler 1. 1. Vorbrodt, 1912: 106.

Type locality: ‘Zermatt, in etwa 40 Stücken erzogen’.

2-Syntypes 46 , 49:

e ¢ with labels: printed, with handwritten (Püngeler) insertion

‘Zermatt | Rhamnus alpina | Püng.[eler] e[x] /.[arva] fin.[e]

7.1907 (on white paper) and handwritten (Püngeler) on printed

form ‘Type <double underlined> | v. brevicaudis Püng.[eler] |

3’ (on pink paper);

e 33,4% with same printed labels with dates of ‘fin. 7.1901’ and

‘fin. 7.1908’ (all ex larvae) and handwritten (Püngeler) on print-

ed form ‘Cotype <underlined> | v. brevicaudis Püng.[eler] | 4

(2)’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Spini Schiff. | v. Brevicaudis Püng.’ (on white paper, in

printed box).

brillantina Staudinger, 1887

‘Thecla Brillantina Stgr.’ Staudinger, 1887b: 130, Taf. 6, Fig. 3, a, b,c. Type locality:

*...erhielt ich nur von der Insel Askold und vom Suifun durch Dörries in einer grösseren

Anzahl meist gezogener Stiicke’.

Syntypes 5d, 59:

e ¢ with labels: handwritten (Staudinger) ‘Brillantina | Stgr.’ (on

white paper), handwritten ‘Suifun | [18]85 Dörr.[ies]’ (on bluish

paper), printed ‘Origin.’ (on pink paper);

e © with circle of bluish locality label paper and printed ‘Origin.’

(on pink paper);

e 6 with labels: handwritten ‘Askold | [18]83 Dorr.[ies]’ (on

bluish paper) and printed ‘Origin. (on pink paper);

e 36,49 with circle of bluish locality label paper and printed

‘Origin.’ (on pink paper); one © also has a handwritten label

‘12’ (on white paper).

221

caerulea Staudinger, 1871

‘Lycaena Damon Schiff. ab. 2 (et v.?) Caerulea; Damone © var. Ld. Hor. 1. c. T. IV, 8;

Var.

intermedia Iphigeniae et Damonis Ld. I. c. (® caerul., 4 marg. lato nigro).’

Staudinger, 1871a: 13. Type locality: ‘Hyrc.[aniae] alp.[es]’.

Syntypes 56,99:

222

? with labels: handwritten ‘Hadschyabad” (on white faded

paper), printed, with handwritten insertion ‘Damone | Coll.

Led. (on white paper), printed ‘Origin.’ (on pink paper), print-

ed ‘Zool. Mus. | Berlin’ (on pale yellow paper), handwritten

(Wagener, in ball pencil), handwritten ‘caerulea’ (on white

paper);

2 with labels: handwritten “‘Hadschyabad’ (on white faded

paper), printed, with handwritten insertion ‘Damone | Coll.

Led.’ (on white paper), printed ‘Origin. (on pink paper), print-

ed ‘Zool. Mus. | Berlin’ (on pale yellow paper), handwritten

(Wagener) ‘Lycaena | caerulea | Stgr’ (on white paper) hand-

written ‘27 (on white notebook cross-lined paper), printed,

with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Tiirkei. |

Tafel 727 Figur 57’ (on white paper);

d with labels: handwritten ‘Hadsch.[yabad] | Hbhr.

[Haberhauer]’ (on yellow paper), printed ‘Origin. (on pink

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Wagener) “Lycaena | caerulea | Stgr.’ (on white

paper), handwritten ‘26’ (on white notebook cross-lined paper),

printed, with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türkeı. |

Tafel 721 Figur 56’ (on white paper);

2 with labels: handwritten ‘Hadsch.[yabad] | Hbhr.

[Haberhauer]’ (on yellow paper), printed ‘Origin.’ (on

pink paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow

paper), handwritten (Wagener) ‘Lycaena | caerulea | Stgr.

(on white paper), printed, with handwritten (Wagener)

insertions ‘Abgebildet in Hesselbarth | van Oorschot &

Wagener: | Tagfalter der Türkei. | Tafel 72/7 Figur 58’ (on white

paper);

e G with labels: handwritten ‘Schahkuh | Chr.[istoph]’ (on yellow

paper), printed ‘Origin.’ (on pink paper), handwritten date label

‘18/5’ (‘18° on white, ‘V’ on pink background, recto white/pink,

verso pink);

36 with circle of yellow locality label paper, printed ‘Origin.

(on pink paper), handwritten date label ‘18/V’ (‘18° on white,

“V’ on pink background, recto white/pink, verso pink);

2 with circle of yellow locality label paper, printed ‘Zool. Mus.

| Berlin’ (on pale yellow paper), handwritten (Wagener)

‘Schahkuh’ (on white paper), handwritten (Wagener in ball pen-

cil) ‘caerulaea | Berlin’ (on white paper), printed, with hand-

written (Wagener) insertions ‘Abgebildet in Hesselbarth | van

Oorschot & Wagener: | Tagfalter der Türkei. | Tafel 727 Figur

59’ (on white paper);

e 26,49 with circle of yellow locality label paper and small

Christoph’s date labels. — ? above and these specimens

undoubtedly belong to the type series though not provided with

Staudinger’s ‘Origin.’ labels (cf. Hesselbarth et al., 1995: 728,

Taf. 121, Fig. 56-60; Häuser & Eckweiler, 1997: 68);

e © with labels: handwritten (Püngeler) ‘Hyrcania (recto) | als

caerulea 2 | vom Stgr (verso)’ (on white paper, recto in printed

box), handwritten (Püngeler) on printed form ‘Cotype <under-

lined> | v. caerulea Stgr. 2 | vom Autor. | Püngeler’ (on pink

paper) — cf. Häuser & Eckweiler (1997: 68).

caerulescens A. Bang-Haas, 1912

‘Callophrys rubi v. caerulescens n. var. Bang-Haas, A., 1912: 106. Type locality: ‘Aus

dem Kaukasus (Elisabethpol).” — Junior subjective synonym of Callophrys rubi chaly-

beitincta Sovinsky, 1905 (Hesselbarth er al., 1995: 516).

Lectotype d:

e d with labels: handwritten (O. Bang-Haas - sic) ‘Callophrys |

Rubi | v. Caerulescens | B[ang] H[aas]’ (on white paper), print-

ed ‘Elisabethpol | Kaukasus’ (on white paper) and printed

‘Origin. (on pink paper), handwritten (Nekrutenko) on printed

form ‘LECTOTYPUS à | caerulescens O.<sic! recte A.> B.-H.

| Y. Nekrutenko design.’ (on yellow paper) — Nekrutenko in

223

Hesselbarth ef al. (1995: 516), handwritten (Wagener) ‘= chaly-

beitincta Sovinsky’ (on white paper, in ball pencil).

Paralectotypes d, 9:

e d with labels: printed ‘Elisabethpol | Kaukasus’ (on white

paper) and printed ‘Origin.’ (on pink paper), handwritten

(Nekrutenko) on printed form ‘PARALECTOTYPUS G |

caerulescens O. <sic! recte A.> B.-H. | Y. Nekrutenko design.’

(on yellow paper);

e © with labels: printed ‘Elisabethpol | Kaukasus’ (on white

paper) and handwritten (Püngeler) ‘v. caerulescens © | 1/1914

v.[om] B-Haas | Püng.[eler]’.

The specimens are placed under handwritten (Püngeler) drawer

label ‘Rubi L. | ab. Caerulescens Courv. (on white paper, in print-

ed box).

cala Strand, 1919

‘Lycaena semiargus Rott. ab. cala m.’ Strand, 1919: 18. Type locality: ‘Marburg.’ —

Infrasubspecific.

Holotype 4 (by monotypy):

e d with labels: printed ‘Deutschland | Marburg i.[n] H.[essen] |

E. Strand, [190]8. (on white paper), handwritten (Strand) on

printed form ‘Lycaena | semiar- | gus Rott. | ab. cala m. à |

Strand det.’ (on white paper) and printed “Type’ (on dirty-pink

paper).

candalus Herrich-Schäffer, [1851]

‘[Lycaena] Candalus’ Herrich-Schäffer, [1851] 1(49): Tab. 105, Figs 502-505; [1852]

6(55): 26-27. Type locality: ‘Aus Kleinasien’.

Types: not found.

candens Herrich-Schäffer, [1844]

‘Polyommatus candens’ Herrich-Schäffer, [1844] 1(7): Tab. 49, Figs 229-231; [1847]

1(26): Tab. 73, Fig. 355; ‘Polyommatus Candens’ Bischoff [1845] 1(10): 133. Type

locality: ‘Ein Pärchen erhielt Hr. Bischoff aus Kleinasien’.

Types: not found.

224

carmon Herrich-Schäffer, [1851]

‘[Lycaena] Carmon Kad. Herrich-Schäffer, [1851] 6(48): 24; [1851] 1(49): Tab. 105,

Figs 506, 507. Type locality: ‘Ein Mann von Herr Kaden aus Kleinasien’.

Type (holotype by monotypy): not found (cf. Häuser & Eckweiler,

1997: 68).

caspius Lederer, 1869

“Polyomm.[atus] Caspius Led.’ Lederer, 1869: 76, Taf. 4, te Bs IMDS localıay: bei

Astrabad in Persien...’ (op. cit.: 73).

Holotype d (by monotypy):

e ¢ with labels: handwritten ‘Hadschyabad’ and (Lederer?)

‘Polyomat.[us] <sic> | Caspius’ (both on white paper), printed

‘Origin.’ (on pink paper) and ‘Coll. Led.’ (on white paper).

There are one more male and one female specimen with similar

locality labels but without ‘Coll. Led.’ labels provided by

Staudinger with printed ‘Origin.’ Since Lederer clearly declared

‘Ich erhielt nur dies eine Exemplar’ (op. cit.: 77), only one of these,

belonging to Lederer, is de facto a type; other specimens subse-

quently considered by Staudinger as ‘Origin.[ale]’, 1. e. ‘types’,

have no nomenclatural status.

The specimen is provided now with a printed label ‘caspius

Lederer, 1869 | HOLOTYPUS | (par monotypiam) | Y. Nekrutenko

det. 26.10.1998’ (on red paper).

caucasica Lederer, 1870

‘Lycaena Corydon Scop. Var. caucasica Led. (Polona Led. Wien. Ent. Mtsch. T. VIII,

p. 166, non Z.)’ Lederer, 1870a: 23. Type locality: not stated.

Neotype d:

e JS with labels: handwritten ‘Caucas[us]’ (on white faded paper),

printed ‘Coll. Led.’ (on white paper), printed ‘Origin. (on pink

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Schurian) on printed lined form ‘Polyommatus |

(Lysandra) | corydonius cauca- | sicus LED., 1870 | det.

Schurian 1988’ (on white paper), handwritten (Schurian)

‘Neotypus 3 <written over printed ‘Paratypus’> | Lycaena |

225

corydonius caucasica | LEDERER 1870 | det. Schurian, 1988’

(on red paper, in box) — Schurian (1988: 139).

caudatus Staudinger, 1901

‘Chrysophanus Hb. Thetis Klug. v. Caudatus Stgr.’ Staudinger, 1901: 73. Type locali-

ty: ‘Arm.[enia] m.[eridionalis]; Pont.[us] or.[ientalis]; Kurd.[istan] s.[eptentrionalis]

oc.[cidentalis]. — Junior subjective synonym of Lycaena thetis thetis Klug, 1834.

Lectotype à:

e d with labels: handwritten (Staudinger) ‘Thetis II Gen. | v.

Caudata’ (on white paper), handwritten ‘Kasikoparan | Cauc. m.

Chr.[istoph]’ (on green paper), handwritten ‘thetis caudatus

Staudinger, 1901 | LECTOTYPUS | Nekrutenko design. 1983’

(on yellow paper) — Nekrutenko in Hesselbarth er al. (1995:

491).

Paralectotypes 38,42:

e 3d, 42 with circles of green locality label paper —

Nekrutenko in Hesselbarth et al. (1995: 491).

caudatus Staudinger, 1901

‘Chrysophanus Hb. Sarthus Stgr. v. Caudatus Stgr. Staudinger, 1901: 75. Type locali-

ty: ‘Buch.[{ara] s.[eptentrionale] or.[ientale] (Karategin).’ — Junior primary homonym of

Chrysophanus thetis caudatus Staudinger, 1901: 73 (above; precedence in pages);

replacement name: Lycaena ophion Hemming, 1933: 278.

Syntypes 5 d :

e d with labels: handwritten (Staudinger) ‘v. Caudatus | Stgr.’ (on

white paper), handwritten ‘Karategin | [18]89 Maur.[er]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 45 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper); one specimen has also handwritten

(Staudinger) ‘Sarthus var. | Caudatus Stgr. (recto) | Caspius var.

| Catapyrrus | Stgr. in IL. (verso)’ (on white paper); one specimen

has also a handwritten label ‘v. Catapyrrus’ (on white paper, in

pencil); printed Staudinger label ‘Origin.’ missing.

226

charybdis Staudinger, 1886

‘Lyc.[aena] Charybdis Stgr. Staudinger, 1886: 226. Type locality: ‘...erhielt ich aus

der Umgegend von Margelan und Namangan...’

Syntypes 88,5%:

e ¢ with labels: handwritten (Staudinger) ‘Charybdis | Stgr.’ (on

white paper), handwritten ‘Margelan | Maur.[er] [18]84’ (on

brown paper), printed ‘Origin.’ (on pink paper);

e 58,3% with circles of brown locality. label paper and printed

“Origin. (on pink paper); one male specimen has a handwritten

(Staudinger) label ‘Charybdis | Stgr.’ (on white paper);

e G with labels: handwritten ‘Namangan | Hbh. [Haberhauer]

[18]84’ (on brown paper), printed ‘Origin. (on pink paper);

e 3,2% with circles of brown locality label paper and printed

“Origin. (on pink paper).

christophi Staudinger, 1874

‘Lycaena Christophi n. sp.’ Staudinger, 1874: 87. Type locality: ‘Herr Christoph... ent-

deckte diese Art zuerst bei Schahrud im nördlichen Persien.’ ... ‘Im voriger Jahr fand er

die Art einzeln bei Krasnowodsk in Turkestan’; restricted by the lectotype designation:

‘Schahrud [Iran, Semnän Province, Shährüd]’ (Olivier, van der Poorten & De Prins, 1998:

Diy

Lectotype d:

e d (antennae missing), with labels: handwritten (Christoph)

‘Schahrud | Chr[istoph]’ (on yellow paper), printed ‘Origin. (on

pink paper), printed ‘christophi Staudinger, | 1874 | LECTO-

TYPUS | Olivier, v.d. Poorten & De Prins design. | Phegea 1998

26(3): 91, pl. 1, f. 1, pl. 2, f. 1’ — Olivier, van der Poorten &

De srans (1998: 91 pk. 1, 2, fie. 1); Tshikolovets (1997a;, 136,

238, pl. 40, fig. 12(14).

Paralectotypes 66, 122:

e 5d, 11% with circles of yellow locality label paper and printed

labels ‘Origin, some have small handwritten date labels, d

and 4 have no ‘Origin.’ labels;

205

e 3 (abdomen missing), 2 with labels: handwritten (Püngeler)

‘Regn.[1i] Pers.[iae] | sept.[entrionalis]’ (on white paper, in print-

ed box) and handwritten (Püngeler) on printed form ‘Cotype

<underlined> | Christophi Stgr. 3 (®)l vom Autor | Püngeler’

(on pink paper).

chrysopis Grum-Grshimailo, 1888

‘Lycaena Chrysopis m.’ Grum-Grshimailo, 1888: 306. Type locality: “Hindukush

orient.’

Paralectotypes à , 2:

e ¢ with labels: handwritten (Staudinger) ‘Chrysopis | Gr. Gr.

(on white paper), handwritten (Grum-Grshimailo, in Russian)

‘Beik | 12. VII.[18]87’ (on white paper), handwritten

‘Hindukusch or.[ientalis] | [18187 Gr. Gr.’ (on brown paper);

e © with circle of brown locality label paper and handwritten

(Grum-Grshimailo, in Russian) label ‘Beik | 11. VII.[18]87’ (on

white paper).

In evidence that these specimens, forwarded by Grum-

Grshimailo to Staudinger, belong to the type series, a quotation

from a subsequent publication by the author (redescription and type

locality restriction) follows: ‘J’ai découvert Chrysopis dans les

monts Kounjout, où elle volait sur les bords d’un ruisseau, qui se

déverse dans le Beik méridional’ (Groum-Grshimailo, 1890:

416-417, pl. 10, fig. 5a, 5b). Lectotype designated by D’Abrera

(1993: 498, figs. [not numbered] ‘det. Balint’ — deposited in The

Natural History Museum, London; see also Tshikolovets, 1997a:

158).

The specimens are provided now with a printed label ‘chrysopis

| Grum-Grshimailo, 1888 | SYNTYPUS | Y. Nekrutenko det

5.12.1998’ (on yellow paper).

cilissa Lederer, 1861

‘Cigaritis Cilissa’ Lederer, 1861: 147, Taf. 1, Fig. 1. Type locality: ‘Bei Mersin und

Antiochia’.

228

Syntypes 6, 29:

e JS with labels: handwritten ‘Antiochia’ (on white paper), print-

ed ‘Coll. Led.’ (on white paper) and ‘Origin.’ (on pink paper);

e 2° with labels: handwritten ‘Antiochia’, circles of white local-

ity label paper and printed labels ‘Coll. Led.’ (on white paper)

and ‘Origin.’ (on pink paper).

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Transcaspica | Stgr.’ (on white paper).

cinnides Staudinger, 1901

‘Lycaena F. Bellargus Rott. ab. Cinnides Stgr. Staudinger, 1901: 86. Type locality: not

stated. — Infrasubspecific.

Types: not found (probably not designated).

clara Staudinger, 1901

‘Lycaena F. Baton Berg. v. et ab. Clara (Stgr.) Chr. Mém. Rom. III p. 51’ Staudinger,

1901: 81. Type locality: ‘Syr.[ia]; Kurd.[istan]; Tura; Ferg.[ana] m.[eridionalis] etc.’ —

Junior primary homonym of Lycaena clara Edwards, 1880; replacement name:

Scolitantides astabene Hemming, 1932: 181.

Types: not found.

cognata Staudinger, 1892

‘Thecla Orientalis Murr. (Diamantina Obrth) und var. ? Cognata Stgr.’ Staudinger, 1892b:

152 (footnote). Type locality: ‘Dorries brachte 11 Pärchen aus dem Sutschan-Gebiet...

Denn es liegen mir 3 fast ganz mit diesen übereinstimmende & 4 vom Uss.[uri], Suif.[un]

und Ask.[old] vor und von Ask.[old] auch 4 völlig übereinstimmende 9 9°.

Lectotype d :

e d (abdomen removed), with labels: handwritten (Staudinger)

‘v. Cognata 89 | Stgr.’<sic> (on white paper), handwritten

‘Sutschan | [18]90 Dörr.[ies]’ (on violet-bluish paper), printed

“Origin. (on pink paper), three slovenly scribbled handwritten

(Dantchenko): ‘8.03.[19]96 | Dantchenko | abdomen’ and ‘4’

(both on white paper, in red ink), and ‘S. Cognata <blot> | Stg.

| lectotypus | Dantchenko | A. V. gesign. (sic!) | 1996’ (on red

paper, in black ink), printed “Lectotype: - | Thecla orientalis |

229

var. cognata Staudinger | Designated by | S. Matsuda, 1996’ (on

white paper and glued to a larger piece of red coloured paper)

— Matsuda (1996: 175, Fig. la - Ic).

Paralectotypes 49, 109:

230

& with labels: handwritten ‘Askold | Dörr.[ies] [18]83’ (on

bluish-violet paper), printed ‘Origin. (on pink paper), and two

handwritten (Dantchenko) ‘Dantchenko | abdomen |

8.03.[19]96’ and ‘5’ (both on white paper, in red ink), printed

‘Paralectotype: - | Thecla orientalis | var. cognata Staudinger |

Designated by | S. Matsuda, 1996’ (on white paper and glued to

a larger piece of red coloured paper) — Matsuda (1996: 175,

Fig. 3a — 3c).

& with labels ‘Ussuri | Dörr.[ies] [18]82’ (on bluish-violet

paper), printed ‘Origin.’ (on pink paper), printed ‘Paralectotype:

- | Thecla orientalis | var. cognata Staudinger | Designated by |

S. Matsuda, 1996’ (on white paper and glued to a larger piece

of yellow coloured paper) — Matsuda (1996: 175, Fig. 2a — 2c).

32 with circles of bluish-violet locality label paper and printed

‘Origin. (on pink paper), one of them with handwritten

(Dantchenko) labels ‘8.03.96 | Dantchenko | abdomen’ and ‘3’

(both on white paper, in red ink), another with handwritten

(Dantchenko) ‘abdomen | Dantchenko | 8.03.96’ and ‘6’ (both

on white paper, in red ink), all with printed labels

‘Paralectotype: - | Thecla orientalis | var. cognata Staudinger |

Designated by | S. Matsuda, 1996’ (on white paper glued to a

larger piece of yellow coloured paper) — Matsuda (1996: 175,

Figs 4a—c, 5a—c, 7a-c);

2 with labels: handwritten (Staudinger) ‘Orientalis var. |

Cognata Stgr. | 2 dimorph’ (on white paper), handwritten

‘Sutschan | [18]90 Dorr.[ies]’ (on bluish-violet paper), printed

‘Origin.’ (on pink paper), ‘Paralectotype: - | Thecla orientalis |

var. cognata Staudinger | Designated by | S. Matsuda, 1996’

(printed on white paper glued to a larger piece of yellow

coloured paper) — Matsuda (1996: 175, Fig. 6a — 6c);

22 not designated as paralectotypes though placed aside and

having Staudinger’s printed labels ‘Origin. and circles of

bluish-violet locality label paper, each with handwritten

(Staudinger) labels ‘v.? Cognata 22 | von Askold <under-

lined>’ and ‘Brillantina | Orientalis hybr.?’ (both on white

paper).

An additional 23 and 4% housed in another cabinet were appar-

ently not found by Matsuda and also constitute an integral part of

the type series:

e ¢ with labels: handwritten (Staudinger) ‘Thecla v. ? | Cognata

Stgr. | type from Sutschan <sic! in English>’ (on white paper),

a circle of bluish-violet paper) and printed ‘Origin.’ (on pink

paper); |

e 3 with labels: handwritten (Staudinger) ‘Orientalis var. |

Cognata Stgr.’ (on white paper), handwritten ‘Sutschan | [18]85

Dörr.[ies]’ (on bluish-violet paper), printed ‘Origin. (on pink

paper);

e ? with labels: handwritten (Staudinger) ‘v.Cognata 99 |

Taxila <illegible> <crossed out> | von Askold | früher

beschrieben’ (on white paper), a circle of bluish-violet paper

and printed ‘Origin.’ (on pink paper);

e © with a circle of bluish-violet paper and printed ‘Origin. (on

pink paper);

e © with labels: handwritten (Staudinger) ‘Cognata ;, | dimorph?’

(on white paper), a circle of bluish-violet paper and printed

‘Origin.’ (on pink paper);

e © with labels: handwritten ‘Askold | [18]83 Dörr.[ies]’ (on

bluish-violet paper) and printed ‘Origin.’ (on pink paper).

corydonius Herrich-Schäffer, [1852]

‘[Lycaena] Corydonius’ Herrich-Schäffer, [1852] 1(54): Tab. 123, Figs 595, 596; Lycaena

corydon Zeller [Lokalvarietät] Corydonius Kef. [1852] 6(55): 27. Type locality: ‘Aus

der Türkei... Aus Siidrussland.’

Neotype d:

e 3 with labels: handwritten ‘Helenendorf’ (on white faded

yellowish paper), printed ‘Zool. Mus. | Berlin’ (on pale yel-

low paper), handwritten (Schurian) on printed lined form

Zou

‘Polyommatus | (Lysandra) | corydon corydonius | H.-S. 1852 |

det. Schurian 1988’ (on white paper), handwritten (Schurian)

on printed form ‘Gen. Präparat | Nr. 221/1986 | Schurian’ (on

white paper), handwritten (Schurian) on printed form

‘Neotypus d | Lysandra corydonius | H.-S. (1852) | teste

SCHURIAN 1985’ (on red paper), and ‘Neotypus 9 <written

over printed ‘Paratypus’> | [Lycaena] | corydon corydonius |

H.-S. 1852 | des. Schurian, 1988’ (on red paper, in box) —

Schurian (1988: 137, Taf. 1, 2, Fig. 6).

crımaea A. Bang-Haas

Unpublished.

Syntype d:

e 3 with labels: handwritten (Püngeler) ‘Krim (recto) | als

damone Ev. | v. crimaea B-H &@ | 3/1913 v.[vom] Autor |

Püng.[eler] (verso)’, handwritten date label ‘27/VI | 1911’ (on

white paper), handwritten (Forster) on printed form

‘Agrodiaetus | poseidon 9 | krymaea Shelj. | det. W. Forster

1943’, printed, with handwritten (Püngeler) insertion ‘Cotype

<underlined> | v. crimaea B-Haas 4° (on pink paper).

The specimen is placed under a handwritten (Püngeler) drawer

label ‘Poseidon HS. | v. Crimaea B-Haas’ (on white paper, in print-

ed box).

cyanea Staudinger, 1899

‘Lycaena Damone var. Cyanea’ Staudinger, 1899: 142. Type locality: ‘zwei Pärchen

durch Haberhauer von Achalzich erhielt und 3 4 à in Lederers Sammlung aus Hankynda

vorfand’; (restricted by the lectotype designation): Hankynda (Stepanakert = Xankändi,

Nagorno-Karabakh — Forster (1956: 85).

Lectotype à:

e ¢ with labels: handwritten ‘Hankynda’ (on white paper), print-

ed, with handwritten insertion ‘Coll. Led. | Kindermanni’ (on

white paper), printed ‘Origin. (on pink paper), printed “Zool.

Mus. | Berlin’ (on pale yellow paper), handwritten (Forster) on

printed form ‘Lecto-Typus 3 | Lycaena damone | cyanea Stgr. |

W. Forster München’ (on pink paper), printed, with handwritten

252

(Wagener) insertions ‘Abgebildet in Hesselbarth | van Oorschot

& Wagener: | Tagfalter der Türkei. | Tafel 723 Figur 36° (on

white paper) — Forster (1956: 85); Hesselbarth er al. (1995:

Taf. 123, Fig. 36); Häuser & Eckweiler (1997: 70), Eckweiler

& Häuser (1997: 122, Pl. 6).

Paralectotypes 48, 9:

e d with labels: handwritten ‘Hankynda’ (on white paper), print-

ed, with handwritten insertion ‘Coll. Led. | Damocles’ (on white

paper), printed ‘Origin. (on pink paper) and a circle of green

paper (?-see locality label of pseudocyanea Forster below);

e d with labels: printed, with handwritten insertion ‘Coll. Led. |

Damocles’ (on white paper), and printed ‘Origin.’ (on pink

paper);

e JS with labels: handwritten ‘Achalzich | Hbhr. [Haberhauer]

[18]82’ (on green paper) and printed ‘Origin.’ (on pink paper);

e © with a circle of green paper, small date label ‘2/8’ and print-

ed label ‘Origin.’ (on pink paper);

e 3 with labels: handwritten (Staudinger) ‘Derbent | Beckr.

[Becker]’ (on green paper), handwritten (Staudinger)

‘Kindermanni | ab.?’ (on white paper), printed ‘300. (on white

paper faded brown), handwritten (Forster) on printed form

‘Polyommatus | (Agrodiaetus) | carmon à | cyanea Stgr. | det.

W. Forster 1947’ (on white paper).

cyprius Stichel, 1911

‘Thestor romanovi cyprius Stich., subsp. nov.’ Stichel, 1911: 76. Type locality: ‘Sehr

local, auf Höhen von 3000-3500 m nächst Sultanabad.’ — Junior subjective synonym of

Thestor romanovi Christoph, 1881 (Hesselbarth et al., 1995: 506).

Syntypes d, 9:

e ¢ with labels: cuttings from the proof-sheet of the original

description ‘Thestor romanovi cyprius Stich. | ...larum omnium

latiore a subspe...’ and ‘Sultanabad’ (on white paper, the text is

on one side only, so that they are for sure cut out from a proof

sheet), printed ‘Nachträgl. der | Coll. Piingeler | hinzugefügt.’

(on greyish paper) and printed “Typus’ (on red paper);

255

e ? with labels: printed ‘Nachträgl. der | Coll. Püngeler |

hinzugefügt. (on greyish paper) and printed “Typus’ (on red

paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Romanovi Chr.’ (on white paper, in printed box).

cytis Christoph, 1877

‘Lycaena Cytis Chr.’ Christoph, 1877: 234, Tab. 5, Figs 5, 6. Type locality: ‘Ich fing

den, wie es scheint, sehr seltenen Schmetterling im Juli bei Schahkuh... von 8 bis

10,000’ Höhe’.

Syntypes 6, 9:

e 6, 2 with labels: handwritten (Püngeler) ‘Hyrcania’ (on white

paper, in printed box), and handwritten (Püngeler) on printed

form ‘Cotype <underlined> | Cytis Chr & (®)l von Stgr. |

Püngeler’ (on pink paper); male specimen with small handwrit-

ten (Christoph) date label ‘11/7’ (recto white, verso green),

female specimen with small square piece of white paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label “Cytis Chr.’ (on white paper, in printed box).

dagmara Grum-Grshimailo, 1888

‘Lycaena Dagmara m. Grum-Grshimailo, 1888: 306. Type locality: “Darwaz’.

Syntypes 6, 2:

e 3 with labels: handwritten (Staudinger) ‘Dagmara | Gr. Gr. |

Chingoe-darja | Darwas.’ (on white paper), handwritten

‘Darwas | [18]85 Gr.[um] Gr.[shimailo]’ (on brown paper),

handwritten (Grum-Grshimailo, in Russian) “Tobi | dara.’ (on

white paper) and small handwritten (Grum-Grshimailo) date

label ‘10.VI | [18]85. (on white paper);

e © with circle of brown locality label paper, handwritten (Grum-

Grshimailo, in Russian) “Tobi | dara.’ (on white paper) and

small handwritten (Grum-Grshimailo) date label ‘10.VI |

[18]85. (on white paper).

234

In evidence that these specimens, forwarded by Grum-

Grshimaïlo to Staudinger, belong to the type series, a quotation

from a subsequent publication by the author (redescription and type

locality restriction) follows: ‘Je n’ai trouvé Dagmara qu’une seule

fois, près du col Agwaï-Polousak et en suite sur l’ancien “saï’ du

Khingoob-Darya, près de Tobi-Dara. De 7,000 à 5,500 p.

(Commencement de Juin)’ (Groum-Grshimailo, 1890: 412-414,

pl. 9, fig. 2a, 2b; see also Häuser & Eckweiler, 1997: 70).

The specimens are provided now with a printed label ‘dagmara |

Grum-Grshimaïlo, 1888 | SYNTYPUS | Y. Nekrutenko det.

16.11.1998’ (on yellow paper).

dama Staudinger, 1892

‘Lycaena Dama Stgr. n. sp.’ Staudinger, 1892a: 234. Type locality: ‘...wurde von

Manissadjian Ende Juli bei Malatia in einer grössere Anzahl von d d und weniger © 2

entdecket’.

Syntypes 68,42:

e d with labels: handwritten (Staudinger) ‘Dama | Stgr. (on

white paper), handwritten ‘Malatia | [18]84 Man.[issadjian]’

(on yellow paper), printed ‘Origin.’ (on pink paper), printed

‘Zool. Mus. | Berlin’ (on pale yellow paper), printed, with hand-

written (Wagener) insertions ‘Abgebildet in Hesselbarth | van

Oorschot & Wagener: | Tagfalter der Türkei. | Tafel 778

Figur 29° (on white paper):

e 5d, 42 with circles of yellow locality label paper, printed

‘Origin.’ (on pink paper), some with small date labels, and

printed ‘Zool. Mus. | Berlin’ (on pale yellow paper); one of the

male specimens also has new handwritten labels ‘20. (on cross-

lined notebook paper) and ‘dama’ (on white paper), one male

specimen — handwritten label (Staudinger) ‘Dama | Stgr.’ (on

white paper) and handwritten date label ‘25/7’ (on white faded

paper), one female specimen — a handwritten label ‘21. (on

cross-lined notebook paper), handwritten (Wagener) ‘Lycaena

dama | Stgr. 1892 © | Malatia 84 Man.’ (on white paper) and

printed, with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türkei. |

Tafel 718 Figur 52 <recte: 49>’ (on white paper) —

235

Hesselbarth ef al. (1995: 727, Taf. 118, Fig. 29, 49); cf. Häuser

& Eckweiler (1997: 70-71), Eckweiler & Häuser (1997: 133,

Pl. 3) and Olivier, De Prins, van der Poorten & Puplesiene

(1999: 202-204).

damonides Staudinger, 1899

‘[Lycaena] Damone Ev. var. Damonides’ Staudinger, 1899: 138. Type locality: ‘In den

Gebirgen Nordpersiens, besonders bei Hadschyabad und Schahkuh, sowie im

südostlichen Transcaucasien (bei Ordubad [Azerbaijan, Naxcivan], auch bei

Kasikoporan...’

Lectotype d:

3S (abdomen missing) with labels: handwritten (Staudinger) ‘v.

Damonides | Stg. (on white paper), handwritten ‘Ordubat |

Chr.[istoph]’ (on green paper), handwritten (unknown hand) ‘L.

Iphigenia ? | Ordubad 10.6.[18]81’ (on faded paper, the text

strongly faded, almost invisible), printed ‘Origin.’ (on pink

paper), printed “Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Forster) on printed form ‘Agrodiaetus | poseidon

3 | damonides Stgr. det W. Forster 1948’ (on white paper),

handwritten (Forster) “Cotypus | Lycaena | damone | damonides

Stgr. (on pink paper), handwritten (Forster) “Lectotypus |

Lycaena | damone | damonides Stgr. | W. Forster 1948’ (on pink

paper), printed, with handwritten (Wagener) insertions

‘Abgebildet in Hesselbarth | van Oorschot & Wagener: |

Tagfalter der Türkei. | Tafel 727 Figur 67’ (on white paper) —

Forster (1961: 13); Hesselbarth er al. (1995: Taf. 121, Fig. 67);

Hauser & Eckweiler (1997: 72); Eckweiler & Häuser (1997:

138, Pl. 5); Olivier, Puplesiene, van der Poorten, De Prins &

Wiemers (1999: 15, Fig. 2).

Paralectotypes:

236

33, © all with circles of green locality label paper, one male

has also handwritten (unknown hand) label ‘Ordub.[ad]

22.6.[18]81’ (on faded paper, the text strongly faded, almost

invisible) and printed ‘Origin.’ (on pink paper), one male —

handwritten (unknown hand) label ‘Ordub.[ad] 25.6.[18]81’ (on

faded paper, the text strongly faded, almost invisible), printed

‘Zool. Mus. | Berlin’ (on pale yellow paper), and two new

labels: handwritten (Wagener) ‘Lycaena damone damo- | nides

Stgr. 1899 | Ordubad’ (on white paper) and printed, with hand-

written (Wagener) insertions ‘Abgebildet in Hesselbarth | van

Oorschot & Wagener: | Tagfalter der Türkei. | Tafel /2/

Figur 6/’ (on white paper) (figured in Hesselbarth er al., 1995:

Taf. 121, Fig. 61, not ‘2’ as stated in the legend); female speci-

men has handwritten (unknown hand) label ‘Ordub.[ad]

22.6.[18]81’ (on faded paper, the text strongly faded, almost

illegible); |

e ? with labels: handwritten (unknown hand) ‘Ordub.[ad]

10.6.[18]81’ (on faded paper, the text strongly faded, almost

invisible), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Forster) on printed form ‘Agrodiaetus | poseidon

2 | damonides Stgr. det W. Forster 1948’ (on white paper),

handwritten (Forster) ‘Cotypus | Lycaena 2 | damone |

damonides Ster. (on pink paper), printed, with handwritten

(Wagener) insertions ‘Abgebildet in Hesselbarth | van Oorschot

& Wagener: | Tagfalter der Türkei. | Tafel /2/ Figur 66° (on

white paper) — Olivier, Puplesiene, van der Poorten, De Prins

& Wiemers (1999: 16).

All above specimens are placed under a_ handwritten

(Staudinger) drawer label ‘v. Damonides | Stgr. (on white paper).

darius Eckweiler & ten Hagen, 1998

‘Polyommatus (Agrodiaetus) darius spec. nov. Eckweiler & ten Hagen, 1998: 110,

Abb. 2-6. Type locality (holotype): ‘Iran, Tehran, Elburs, Demavend, Polur, 2400 m’.

Paratype d:

e d with printed labels: ‘Iran, Tehran | Elburs, Demavend | Polur

2400 m | 18.V11.1997 | leg. W. ten Hagen’ (on white paper),

‘Paratypus | Polyommatus (Agrodiaetus) | darius spec. nov. |

des. ECKWEILER & TEN HAGEN 1998’ (on bluish paper, in box).

decolor Staudinger, 1886

‘Lycaena Argiades Pall. var. Decolor Stgr. Staudinger, 1886: 203. Type locality: ‘Aus

dem Umgegend von Margelan...’

237

Syntypes 36, 9:

e 6 with labels: handwritten (Staudinger) ‘var. Decolor | Ster.

(on white paper), handwritten ‘Margelan | Maur.[er] [18]83’ (on

brown paper) and printed ‘Origin. (on pink paper);

e 23, 2 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

decolorata Staudinger, 1886

‘Lycaena Argiades Pall. ... aberr. (eventuell auch var.) Decolorata. Staudinger, 1886:

204. Type locality: ‘...aus Wien, Ungarn und von Bulgarien.’

Syntypes 38,32:

e à with labels: handwritten (Staudinger) ‘ab. Decolorata | Stgr’

(on white paper), handwritten ‘Wien’ (on yellow paper) and

printed ‘Origin. (on pink paper);

3d with labels: handwritten ‘Eperies | Dahlstr. (on white paper)

and printed ‘Origin. (on pink paper);

e 6, ? with circles of white locality label paper and printed

“Origin. (on pink paper);

? with labels: handwritten ‘Balkan | Kr.[üper]’ (on yellow

paper) and printed ‘Origin. (on pink paper);

e © with circle of white locality label paper and printed ‘Origin.

(on pink paper).

dentata Staudinger, 1892

‘Thestor Callimachus var. Dentata’ Staudinger, 1892a: 233. Type locality: ‘bei

Mardin.’ — Junior subjective synonym of Lycaena callimachus Eversmann, 1848.

Syntypes 56 , 59:

e ¢ with labels: handwritten (Staudinger) ‘var. Dentata | Stgr.’

(on white paper), handwritten ‘Mardin | [18]91 Manis.[sadjian]’

(on yellow paper) and printed ‘Origin. (on pink paper);

e 44,59 with circles of yellow locality label paper and printed

‘Origin.’ (on pink paper); one male specimen also has a hand-

written (Staudinger) label ‘Callimachus | v. Dentata | Mardin.’

(on white paper).

238

depuncta Hirschke, 1903

‘Lycaena Argiades Pall. ab. (et var.) depuncta’ Hirschke, 1903: 270. Type locality:

“...ın oberen Czernatal...’ — Infrasubspecific.

Syntype d

e JS with labels: printed, with handwritten (Hirschke?) insertion

‘Hirschke. Wien | Czernathal | V 1901 | Type (recto) | Depuncta

Hirschke | 3 | von | H. Hirschke 3/04 (verso, Püngeler)’ (on

white paper, recto in printed box), handwritten ‘alcetas’ (on

white paper, ın pencil), handwritten (Püngeler) on printed form

‘Cotype <underlined> | ab. depuncta Hirschke 3 | 3.1904 vom

Autor | Püngeler’ (on pink paper).

The specimen is placed under a handwritten (Püngeler) drawer

label ‘Argiades Pall. | ab. Depuncta Hirschke’ (on white paper, in

printed box).

diana Miller, 1913

‘Lycaena diana nov. sp. Miller, 1913: 220. Type locality: ‘L. diana fing ich im

Kagysmanschen Bezirk zum grössen Teil in der Umgegend des Bergsee’s Tschanglé,

nördlich vom Arax. ... Die Höhe, in der sie leben, beträgt wie gesagt, etwa 1800-2000 m;

Fangdatum bei Tschanglé 15.-17.VI.[1911]’ (op. cit.: 221, 222).

Synfypes 6, 2:

e 3 with labels: printed ‘Kagyzman | 2500m <sic> Juni |

Russ.[isch] Armen.[ien]’ with handwritten (Püngeler) insertion

on verso ‘als diana Miller | & 4/1912 von | B-Haas |

Püng.|eler]’ (on white paper), handwritten (Püngeler) on print-

ed form ‘Cotype <underlined> | diana Miller 3 | von B-Haas |

Püngeler’ (on pink paper);

e © with labels: printed ‘Kagyzman | 2500m <sic> Juni |

Russ.[isch] Armen.[ien]’ with handwritten (Püngeler) insertion

on verso ‘als diana Miller 2 | 4/1912 von] B-Haas |

Püng.|eler]’ (on white paper), handwritten (Püngeler) on print-

ed form ‘Cotype <underlined> | diana Miller 3 (2) | von

B-Haas | Püngeler’ (on pink paper).

The lectotype was selected and designated from the type series

unequivocally designated by Miller (“Vorliegendes Material 7 d

239

3 2’) deposited now in Zoological Museum, Moscow University

(Nekrutenko in Hesselbarth ef al., 1995: 655; cf. Antonova, 1981:

211). Up to the present, only 74 and 2? have been found. The

forementioned specimens which Püngeler obtained from Bang-

Haas may hardly belong to Miller’s type series, and this doubt is

clearly expressed on the labels.

difficillima Forster, 1956

‘Agrodiaetus transcaspica difficillima ssp. nov.’ Forster, 1956: 76, Taf. 10, Figs 5, 10,

Taf.

11, Figs 5, 10. Type locality: ‘Nordpersien, Schahkuh’ (holotypus).

Paratypes 3d , 39:

240

d with labels: handwritten (Staudinger) ‘Damone var ?’ (on

white paper), handwritten ‘Hadsch. ? | Hbhr. [Haberhauer]’ (on

yellow paper), printed ‘290. (on white paper faded brown),

handwritten (Forster) on printed form ‘Para-Typus à | Agr.[odi-

aetus] transcaspica | ssp. difficillima Forster | W. Forster

München’ (on pink paper);

dS with labels: printed ‘314.’ (on white paper faded brown),

printed ‘Origin.’ (on pink paper), handwritten (Forster) on print-

ed form ‘Co-Typus d | Lycaena damone | iphidamon

Staudinger’ (on pink paper), handwritten (Forster) on printed

form ‘Para-Typus d | Agr.[odiaetus] transcaspica | ssp. difficil-

lima Forster | W. Forster München’ (on pink paper);

® with labels: handwritten ‘Schakuh | Chr.[istoph]’ (on yellow

paper), printed ‘289.’ (on white paper faded brown), small date

label ‘22/7’ (recto white, verso green), a small piece of white

paper, handwritten (Forster) on printed form ‘Co-Typus © |

Lycaena damone | ssp. damonides Stgr.’ (on pink paper), hand-

written (Forster) on printed form ‘Para-Typus 2 | Agrodiaetus

trans- | caspica difficillima | Forster | W. Forster München’ (on

pink paper);

8,2% with labels: printed, with handwritten insertions ‘HYR-

CANIA, (Schahkuh) | <male with handwritten date 25.7>’ (on

white paper, in box) and handwritten (Forster) on printed form

‘Para-Typus 3 (2) | Agr[odiaetus] transcaspica | ssp. difficil-

lima Forster | W. Forster Miinchen’ (on pink paper); female

specimens also have handwritten labels ‘transcaspica’ (on white

cross-lined notebook paper).

dilutior Staudinger, 1881

‘Polyomm.[atus] Dimorphus Stgr. n. sp. ... var dilutior’ Staudinger, 1881: 283. Type

locality: ‘...bei Margelan (und, wie ich glaube, auch im Tianschen) kommt sie mit

Phoenicurus zusammen (ob an anderen Lokalitäten fliegend?) vor’.

Lectotype d:

e 3 (abdomen missing), with labels: handwritten ‘Margelan |

Hbhr. [Haberhauer]’ (on brown paper), printed ‘Origin. (on

pink paper), printed ‘ex collect. | Staudinger’ (on photographic

paper) and printed *Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Nekrutenko) on printed form in box ‘Figurat | par

Y. Nekrutenko in: | Vestn. zool. 1984, 6: | tab. ex adv. p. 48° (on

white paper), handwritten (Nekrutenko) ‘dilutior Stgr. |

Lectotypus | Y. Nekrutenko design. | 10.1X.1983’ (on yellow

paper) — Nekrutenko (1984: 45, pl. [1]).

Paralectotypes 56:

e 54 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

dimorphus Staudinger, 1881

‘Polyomm.{atus] dimorphus’ Staudinger, 1881: 282. Type locality: ‘Von dieser merk-

würdigen Art sandte Haberhauer 6 Stück (1 8,5 2) meist geflogen von Lepsa ein’.

Lectotype d:

e d with labels: handwritten ‘Lepsa | Hbhr. [Haberhauer]’ (on

brown paper), printed “Origin. (on pink paper, in box), printed

‘ex collect. | Staudinger’ (on photographic paper), printed

‘Zool. Mus. | Berlin’ (on yellow paper), handwritten

(Nekrutenko) on printed form in box ‘Figurat | par

Y. Nekrutenko in: | Vestn. zool. 1984, 6: | tab. ex adv. p. 49’ (on

white paper), handwritten (Nekrutenko) ‘dimorphus Stgr. |

Lectotypus | Y. Nekrutenko design. | 10.1X.1983’ (on yellow

paper) — Nekrutenko (1984: 46, pl. [2]).

241

Paralectotypes 5°:

e 52 with handwritten (Nekrutenko) labels ‘dimorphus Ster. |

Paralectotypus | Y. Nekrutenko design. | 10.IX.1983’ (on blue

paper) — Nekrutenko (1984: 44).

dobrogensis Caradja, 1895

‘Thestor Nogelii HS. var. Dobrogensis Caradja’ Caradja, 1895: 34. Type locality:

Romania, Dobrogea, ‘... Ende Juni [1865] 2 Stunden von Tultscha auf Bergen an

Astragalus ponticus gefligen (Mn. [Mann]).’— Junior subjective synonym of Thecla

Nogelii Herrich-Schäffer, [1851] (Hesselbarth er al., 1995: 510).

Syntypes 36,29:

e 6 with handwritten label “Tultscha | Mn. [Mann]’ (on yellow

paper);

e © with a circle of yellow locality label paper;

e 24 with labels: handwritten “Tultscha’ and printed ‘Coll. Led.’

(both on white paper);

e © with printed label ‘Coll. Led.’ (on white paper).

As supportive evidence that these specimens belong to the type

series, a quotation from the original description follows (Caradja,

1895: 1: ‘Der rühmlichst bekannte Entomologe [Mn. = Joseph

Mann - op. cit.: 6] giebt uns neben eingehender Schilderung der

vor ihm besuchtet Gegend eich reichhaltiges Verzeichniss der

währen der Monate Mai, Juni und Juli [1865] bei Tultscha... Ein

Theil dieser Thiere befindet sich im K. K. Hofmuseum zu Wien, ein

anderer Theil derselben kam mit Lederer’schen Sammlung in der

Besitz des Herrn Dr. Otto Staudinger’).

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Dobrogensis | Caradja | alle typen <sic>’ (on white

lined notebook paper); each provided now with a printed label:

‘dobrogensis | Caradja, 1895 | SYNTYPUS | Y. Nekrutenko det.

13.11.1998’ (on yellow paper).

dombaiensis Alberti, 1969

‘Lycaena anteros dombaiensis nov. ssp.’ Alberti, 1969: 197, Taf. 2, Fig. lc. Type locali-

ty: ‘...am Tschutschchur-Paß (vereinfacht geschrieben Tschutschur-Paß) bei Dombai in

Höhen von 2200-2900 m im Juli und August...’

242

Paratypes 46:

e 43 with labels: printed, with handwritten (Alberti) insertions

“N. W. Kaukasus | Dombai 2300 (2800) m | Tschutschur | 78.

(20., 19/21) 7.1965 | leg. Alberti’ (on white paper, in blue ball

pencil, in box), handwritten (Alberti) on printed form ‘ssp. à |

dombaiensis | det.<crossed out> B. Alberti (recto) | Paratypus

(verso) (on white paper, recto in box, handwritten text in blue

ball pencil), printed ‘Paratypus’ (on red paper, in box).

dschagataicus O. Bang-Haas, 1915

‘Lycaena pheretiades dschagataicus n. var. B.-H. Bang-Haas, O., 1915: 101. Type

locality: ‘... am Nordabhange des Karakorum, in den gebirgen südlich der Stadt Chotan

(Chinesische Tatarei)...’ (p. 93).

Types: not found.

duplicata A. Bang-Haas, 1910

‘Lycaena damone v. duplicata n. v. Bang-Haas, A., 1910: 30. Type locality: ‘Aus dem

Juldus-Gebiete 3 dd’.

Syntypes 36:

e 3 with labels: handwritten (O. Bang-Haas) ‘L.[ycaena]

Damone | v. Duplicata | BH.’ (on white paper), handwritten (O.

Bang-Haas) ‘Lyc.[aena] Überg.[ang] zu | Damone | v. Melania

Stg | nach Pgl. [Püngeler] 1/1 [19]09’ (on white paper) printed

‘Juldus’ (on white paper) and handwritten ‘Orig.’ (on pink

paper);

e 26 with labels: printed ‘Juldus’ (on white paper) and printed

‘Origin.’ (on pink paper) — cf. Häuser & Eckweiler (1997: 74).

egea Herrich-Schaffer, [1852]

‘[Lycaena] Egea Friv.’ Herrich-Schäffer, [1852] 6(55): 25-26; ‘Lycaena Bavius’ [1847]

1(26): Tab. 74, Figs 357-360. Type locality: ‘Aus der Türkei’.

Types: not found.

243

epidolus Herrich-Schäffer, [1843]

‘Lycaena Epidolus’ Herrich-Schäffer, [1843] 1(1): Tab. 4, Figs 18, 19; ‘[Lycaena]

Epidolus Boisd. [1844] 1(7): 114. Type locality: ‘Bei Constantinopel. Von Herrn

Frivaldszky und Herrn Keferstein erhalten.’ — Junior primary homonym of Lycaena epi-

dolus Boisduval, 1840; junior subjective synonym of Lycaena menalcas Freyer, 1837.

Types: not found (cf. Häuser & Eckweiler, 1997: 75).

erotides Staudinger, 1392

‘Lyc.[aena] Eros O. var. Erotides Stgr.’ Staudinger, 1892d: 319. Type locality: ‘Die 13

vorliegenden d d vom Kentei...’

Syntypes 5d:

e 3 with labels: handwritten (Staudinger) ‘v. Erotides | Stgr.’ (on

white paper), and ‘Kentei | [18]89 Dorr.[ies]’ (on green paper),

printed “Origin. (on pink paper);

e 34 with circles of green locality label paper and printed label

“Origin. (on pink paper);

e 3 with labels: handwritten ‘Mongol. | (Changai) (recto) |

erotides à | v.[om] Stgr. 4.1900 (verso)’ (on white paper, recto

in printed box) and handwritten (Püngeler) on printed form

‘Cotype <underlined> | v. erotides Stgr. à | vom Autor |

Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Eros O. | v. Erotides Stgr.’ (on white paper, in printed box).

erschoffii Lederer, 1869

‘Lycaena Erschoffii Led.’ Lederer, 1869: 80, Taf. 4, Figs 4, 5. Type locality: ‘...bei

Astrabad in Persien...’ (op. cit.: 73).

Syntypes 2d, 9:

e 24, 2 with labels: handwritten “Hadschyabad’ (on white

paper), printed ‘Coll. Led.’ (on white paper), printed ‘Origin.

(on pink paper) and handwritten (Hauser) on printed form

‘SYNTYPUS | Lycaena | erschoffii | LEDERER | det. Chr.

L. Häuser | V/.[19]94’ (on pink paper) — Häuser & Eckweiler

(OO We).

244

erythrea Staudinger, 1892

“Cigaritis Siphax Luc. ab. (var.) Erythrea Stgr.’ Staudinger, 1892c: 280. Type locality:

‘von Collo (Provinz Constantine)’.

Syntypes 48, 2:

e 3 with labels: handwritten (Staudinger) ‘ab. (var.?) Erythrea |

Stgr.’ (on white paper), handwritten ‘Collo | Seriz’ (on white

paper) and printed ‘Origin.’ (on pink paper);

e 26, 2 with circles of yellow locality label paper and printed

label ‘Origin. (on pink paper),

e ¢ with labels: handwritten ‘Bône | [18]91 Vall.[?]’ and printed

‘Origin.’ (on pink paper).

erzurumensis Eckweiler & Hesselbarth, 1978

‘Agriades pyrenaicus erzurumensis n. subsp. Eckweiler & Hesselbarth, 1978: 65,

Abb. 14. Type locality (holotype): “Türkei, Prov. Erzurum: Palandöken Dag,

2900-3100 m.’ — Junior subjective synonym of Lycaena dardanus Freyer, [1843]

(Hesselbarth et al., 1995: 622).

Paratypes 36:

e 36 with printed labels: “Türkei/Erzurum | Palandöken Dag |

2900-3100 <sic> m | 27.7-31.7.[19]77 | leg. Eckweiler’ (on

white paper) and ‘Paratypus à | Agriades pyrenaicus | ssp. erzu-

rumensis | des. ECKWEILER et | HESSELBARTH 1978 (on

red paper).

estonica Hoyningen-Huene, 1883

‘Polyommatus [virgaureae] var. Estonica’ Hoyningen-Huene, 1883: 49. Type locality:

‘Die var. Estonica fliegt in Esthland, unter 59'/,° nördl. Breite...’

Syntypes 4d , 49:

e à with labels: handwritten (Staudinger) ‘Var. Estonica | Huene’

(on white paper), printed ‘Estonia. | HUENE’ (on dirty-violet

paper, in box) and circle of green paper;

e 33,42 with same printed labels and circles of green paper.

Part of the type material was sent by the author through to

Staudinger (and to S. Alphéraky) to get an advice before publish-

ing a description (Hoyningen-Huene, 1883: 50).

245

The specimens are provided now with a printed label ‘virgaureae

estonica Huene, 1883 | SYNTYPUS | Y. Nekrutenko det.

26.10.1998’ (on yellow paper).

euphemia Staudinger, 1887

‘Lycaena Euphemia Stgr. (Euphemus Hb. var.?)’ Staudinger, 1887b: 142, 288, pl. 13,

fig. 6. Type locality: ‘... von Raddefka (nur 2 dd), Ussuri, Askold (nur 1 ©) und von

Sidemi in Korea und nordlich von Peking.’

Syntypes 56,39:

e 6 with labels: handwritten (Staudinger) ‘Euphemus var. |

Euphemia Stgr.’ (on white paper), handwritten ‘Raddefskaja |

Chr.[istoph].’ (on violet-blue paper), small date label ‘19/7’ and

printed ‘Origin’ (on pink paper);

e 6, $ with circles of violet-blue locality label paper and print-

ed ‘Origin’ (on pink paper);

e © with labels: handwritten ‘Askold | Dörr.[ies] [18]83’ (on vio-

let-blue paper) and printed ‘Origin’ (on pink paper);

e 3 with labels: handwritten ‘Sidemi | Jank.[owsky?]’ (on violet-

blue paper) and printed ‘Origin’ (on pink paper);

e 3, 2 with circles of violet-blue locality label paper and print-

ed ‘Origin’ (on pink paper);

e JS with labels: handwritten ‘Ussuri | Dörr.[ies] [18]82’ (on vio-

let-blue paper) and printed ‘Origin’ (on pink paper);

eversmanni Staudinger, 1886

‘Lycaena Eversmanni Stgr. n. sp. Staudinger, 1886: 205. Type locality: ‘...in den

Gebirgen in einer Höhe von 1500—2000 Meter bei Margelan, Osch, Namangan und

Samarkand im Juni und Juli vorzukommen’ (see also Forster, 1941: 70-73, Taf. 2, Figs 1,

2, Taf. 3, Figs 1, 2). — Junior primary homonym and junior objective synonym of

Lycaena eversmanni Lang, 1884: 370; senior objective synonym of Plebejus ardis Balint

& Johnson, 1997: 14, syn. n.

Syntypes 129, 119:

e 3 with labels: handwritten (Staudinger) ‘Eversmanni | Stgr.’

(on white paper), ‘Margelan | Hbhr. [Haberhauer]’ (on brown

paper), and printed ‘Origin. (on pink paper);

246

d with labels: handwritten ‘Namangan | Hbh. [Haberhauer]

[18]84’ (on brown paper) and printed ‘Origin. (on pink paper);

S with labels: handwritten ‘Prov. Samarkand | Hbhr. [Haberhauer]

[18]81’ (on brown paper) and printed ‘Origin.’ (on pink paper);

S with labels: handwritten ‘Alai | [18]85 H.[aberhauer]’ (on brown

paper) and printed ‘Origin.’ (on pink paper);

6 with labels: handwritten ‘Osch | Hbh. [Haberhauer] [18]82’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

78,11% with printed labels ‘Origin.’ (on pink paper) and circles of

brown locality label paper, 25, 2 also have handwritten labels

‘2000 [metres] | 4/6 [18]80’ (on blue paper).

fergana Staudinger, 1881

‘Lyc.[aena] Loewii Z. var.? Fergana Stgr. Staudinger, 1881: 262. Type locality: ‘Die

Stücke bei Margelan wurden von Ende Mai bis Juni gefangen.’

Syntypes 4d , 59:

e d with labels: handwritten ‘Margelan | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

e 34,49 with circles of the brown locality label paper and print-

ed ‘Origin. (on pink paper);

e © with labels: handwritten ‘Saisan | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

Two of the male syntypes also have handwritten labels ‘08 9/61’

and ‘30/5’ (on blue paper).

fervida Staudinger, 1901

‘Callophrys Rubi L. v. (gen. aest ?) Fervida Stgr. Staudinger, 1901: 70. Type locality:

‘Ib.[eria] m.[eridionalis], Maur.[itania], As.[ia] min.[or].— Junior subjective synonym of

Papilio rubi Linnaeus, 1758.

Syntypes:

e The types were not designated by Staudinger; under a hand-

written (Staudinger) drawer label ‘v. Fervida | Stgr.’, 22 male

and female specimens are placed from different localities: from

S. Spain to Central Asia.

247

firdussiü Forster, 1956

‘Agrodiaetus damone firdussii ssp. nov. Forster, 1956: 58, Taf. 8, Figs 16, 17, Taf. 9,

Figs 16, 17. Type locality (holotypus): ‘Nordiran, Schahkuh, Westabhang, Geröllzone,

1800-2000 m’.

Paratypes d , 29:

e 3,2% with labels: printed, with handwritten insertions ‘HYR-

CANIA, (Schahkuh) [18198 <in male>, 21.7.[1898],

14.7.[18]98 <in females>’ (on white paper, in box) [leg. Funke

— teste Forster, /. cit.: 58] and handwritten (Forster) on printed

form ‘Para-Typus 4 (2) | Agrodiaetus damone | firdussii

Forster | W. Forster München’ (on pink paper) — cf. Häuser &

Eckweiler (1997: 77).

fortunata Staudinger, 1870

‘Lycaena Fortunata nov. sp. Staudinger, 1870a: 99. Type locality: ‘Diese Art wurde

von... Dr. Bauer... auf Teneriffa entdeckt. Dr. Bauer sammelte im Winter und Frühling

1864 auf Madeira und den Canarischen Inseln... — Junior subjective synonym of

Polyommatus webbianus Brullé, [1838] (Staudinger, 1901: 77).

Syntypes 26, 9:

e ¢ with labels: handwritten (Staudinger) “Teneriffa | Bauer’ (on

pale yellow paper) and printed ‘Origin. (on pink paper);

e d with same handwritten (Staudinger) label;

e © with circle of the pale yellow locality label paper and print-

ed ‘Origin.’ (on pink paper).

The specimens are placed under a handwritten drawer label

“Webbianus | Brullé | Fortunata | Stgr.’; each provided now with a

printed label ‘fortunata Staudinger, | 1870 | webbianus Brullé,

[1838] | SYNTYPUS | Y. Nekrutenko det. 1.11.1998’ (on yellow

paper).

frivaldszkyi Lederer, 1855

‘Thecla Frivaldszkyi Kindermann’ Lederer, 1855a: 100, Taf. 1, Fig. 1. Type locality:

‘Den Schmetterling fand Herr Kindermann auf Bergen in der Nahe von Ust-Buchtarminsk

am 2. Juni kurz nach dem Schmelzen des Schnees; er erbeutete 15, meist geflogene

Stiicke und theilte mir ein Parchen mit.’

248

Syntypes d, 9:

e 3 with labels: handwritten (Lederer?) ‘Frivaldsz- | kyi

Kind.[ermann] Led.[erer] | Sibiria occ. (recto) | Frivaldsz- | kyi

(verso)’ (on white paper, recto in printed box), printed ‘Origin.

(on pink paper), ‘Coll. Led.’ (on white paper);

e © with labels: handwritten (Kindermann) ‘Altai’ (on white

paper), printed ‘Origin.’ (on pink paper), ‘Coll. Led.’ (on white

paper).

galba Lederer, 1855

‘Lycaena Galba Kollar’ Lederer, 1855b: 190, Taf. 1 (not 2 as stated in the text), Fig. 4.

Type locality: ‘Ich erhielt nur 8 Stücke [Beirut]’ — /. c.: 187.

Lectotype d:

e 3 with labels: printed, in box ‘Origin.’ (on dirty-pink paper),

printed ‘Coll. Led.’ (on white paper), handwritten (Lederer?)

‘Beirut’ (on white paper), printed “Zool. Mus. | Berlin’ (on pale-

yellow paper), handwritten (Nekrutenko) label ‘galba Lederer,

1855 | Lectotypus d | Y. Nekrutenko design. | 7.X1.1983’ (on

yellow paper) — Nekrutenko (1984: 31, fig. 2, /).

Paralectotypes 3 © :

e © with same labels as lectotype and handwritten (Nekrutenko)

‘galba Lederer, 1855 | Paralectotypus © | Y. Nekrutenko design.

| 1984’ (on blue paper);

e 22 with same printed labels as lectotype, one with a handwrit-

ten label ‘Beirut’ (on white paper).

gamra Lederer, 1855

‘Lycaena Gamra Kollar in lit” Lederer, 1855b: 189, Taf. 1, Fig. 3. Type locality:

“[Beirut]” — /. c.: 187. — Junior subjective synonym of Lycaena jesous Guérin, 1847

(Staudinger, 1901: 77).

Syntypes

e ? — © with printed labels ‘Coll. Led.’ (on white paper) and

‘Origin.’ (on pink paper);

e ? — 26 with printed labels ‘Coll. Led.’ (on white paper);

249

e 24,39 labelled ‘Beirut | Zach’

Lederer (1855b: 189-190) does not mention the number of spec-

imens he saw, but apparently he had material of both sexes that

were collected ‘im Juni und Juli auf Brombeerblüthen’. They were

certainly taken by Franz Zach in 1854, who stayed in Beirut and

collected in its neighbourhood from 22 July 1853 until early

August 1854. Therefore the specimens collected by Zach are cer-

tainly syntypes.

gerhardi Staudinger, 1895

‘Thecla Acaciae Fab. var. Gerhardi Stgr.’ Staudinger, [1895]: 241. Type locality: ‘...bei

Mardin und Aintab (nördliches Mesopotamien, ersteres auch zum südlichen Kurdistan

gerechnet).... — Junior subjective synonym of Thecla abdominalis Gerhard, [1850]

(Hesselbarth et al., 1995: 531).

Syntypes 5d, 4°:

e d with labels: handwritten (Staudinger) ‘var. Gerhardi | Stgr.’

(on white yellowish lined notebook paper), handwritten

‘Mardin | [18]92 Man[issadjian]’ (on yellow paper) and printed

‘Origin.’ (on pink paper);

e 34,29 with circles of yellow locality label paper and printed

‘Origin.’ (on pink paper), one male specimen also has a hand-

written date label ‘24/5’, one also a handwritten (Staudinger)

label ‘var. Gerhardi | Stgr. (on white-yellowish lined notebook

paper, with an insertion in pencil ‘Mardin’);

e © with labels: handwritten (Staudinger) ‘var. Gerhardi | Stgr.’

(on white yellowish lined notebook paper), handwritten

‘Mardin | [18]88 Sont.[?]’ (on yellow paper) and printed

‘Origin.’ (on pink paper);

e ? with circle of yellow locality label paper and printed

‘Origin.’ (on pink paper);

e JS with labels: handwritten ‘Aintab | [18]87 Man[issadjian]’ (on

yellow paper), handwritten date label ‘21/5’ (on white paper)

and printed ‘Origin.’ (on pink paper);

e 3 with labels: handwritten (Püngeler) ‘Mesopot.[amia] (recto)

| acaciae var. | Gerhardi ¢ | von | A. Bang-Haas 1/[19]02

(verso)’ (on white paper, recto in box), handwritten (Piingeler)

250

on printed form ‘Cotype <underlined> | v. Gerhardi Stgr. ¢ |

von Bang-Haas | Püngeler’ (on pink paper);

e © with labels: handwritten (Püngeler) ‘Mesopot.[amia] (recto)

| v. Gerhardi 2 | ex coll Deckert | 5/1902 v.[om] M. Bartel

(verso) (on white paper), handwritten (Püngeler) on printed

form ‘Cotype <underlined> | v. Gerhardi Stgr 2 | von

M. Bartel | Piingeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Acaciae F. | v. Gerhardi Stgr.’ (on white paper, in printed

box).

gigas Staudinger, 1871

‘Lycaena Loewii Z. v. Gigas’ Staudinger, 1871a: 10. Type locality: ‘Lyd.[iae] alp.[es]’

(restricted: ‘Im Taurus fing Lederer eine riesige Form dieser Art... und die ich in meinem

Catalog als v. Gigas bezeichnete.’ — Staudinger, 1878: 234-235). — Junior subjective

synonym of Lycaena loewii loewii Zeller, 1847 (Hesselbarth et al., 1995: 613).

Syntypes 3d, 2°:

e G with labels: handwritten (Staudinger) ‘Taurus | Ld.

[Lederer]’ (on yellow paper) and printed ‘Origin.’ (on pink

paper);

e ? with circle of yellow locality label paper and printed

‘Origin.’ (on pink paper);

e d with labels: handwritten ‘Külek’ (on white paper) and print-

ed: ‘Coll. Led.’ (on white paper) and ‘Origin. (on pink paper);

e 6, © with labels: handwritten ‘Syrien’ (on white paper) and

printed: ‘Coll. Led.’ (on white paper) and ‘Origin.’ (on pink

paper).

gisela Püngeler, 1901

‘Lycaena gisela n. sp.’ Piingeler, 1901: 180, Taf. 2, Fig. 12, 12a, 12b. Type locality: not

stated: ‘Ausgangpunkt war die unter 41° 7’ nördl. Breite und 80° 81’ östl. L. v. Gr. gele-

gene Stadt Aksu...’

Syntypes 48,42:

e 6 with labels: printed, with handwritten (Püngeler) insertion

‘Ost-Turkestan | (Aksu) | Riickbeil 1900’ (on white paper) and

251

handwritten (Püngeler) on printed form ‘Type <double under-

lined> | gisela Püng. 3 | abgebildet Iris 1901 | Püngeler’ (on

pink paper); Ä

e 34,49 with labels: printed, with handwritten (Püngeler) inser-

tion ‘Ost-Turkestan | (Aksu) | Rückbeil 1900’ (on white paper)

and handwritten (Püngeler) on printed form ‘Cotype <under-

lined> | gisela Püng. 8 (?)| Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Gisela Püng.’ (on white paper, in box); the holotype desig-

nation (as “Type’) has not been published.

glaucias Lederer, 1870

‘Lycaena Glaucias Led. Lederer, 1870b: 10, Taf. 1, Figs 6, 7). Type locality: ‘...bei

Astrabad in Persien... ...in 8000 Fuss Höhe das kleine, nur im Sommer von Hirten

bewohnte Dorf Hadschyabad’ (op. cit.: 4); “Nur 7 Stücke erhalten.’

Syntypes 36, ©:

e d with labels: handwritten (Lederer) ‘Glaucias | m.[ihi]’ (on

white paper, in printed box), and ‘Hadschyabad’ (on white

paper), printed ‘Coll. Led.’ (on white paper) and ‘Origin. (on

pink paper);

e 25, ¢ with labels: handwritten ‘Hadschyabad’ (on white

paper), one male with printed ‘Coll. Led.’ (on white paper), all

with printed labels ‘Origin. (on pink paper) — cf. Hauser &

Eckweiler (1997: 79).

grumi Staudinger, 1901

‘Lycaena F. Eversmanni Stgr. v. ? Grumi Stgr. Staudinger, 1901: 79. Type locality:

‘Transalai (sec.[undum] Maurer)’ (see also Forster, 1941: 72-73, Taf. 2, Figs 6-8, Taf. 3,

Figs 6-8).

Syntypes 48,3%

e d with labels: handwritten (Staudinger) ‘v.? Grumi | Stgr.’ (on

white paper), “Transalai | Pamir? | [18]88 Maur.[er]’ (on brown

paper), and printed ‘Origin’ (on pink paper);

e 34,39 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper).

252,

haberhaueri Staudinger, 1886

‘Lyc.[aena] Sieversii Chr. var. Haberhaueri Stgr. Staudinger, 1886: 206. Type locali-

ty: ‘...Ende Mai bei Namangan wie Samarkand in Anzahl gefangen.

Syntypes 94,89:

e 3 with labels: handwritten (Staudinger) ‘Sieversi var? |

Haberhaueri | Stgr. (on white paper), ‘Namangan | Hbhr.

[Haberhauer] [18]84’ (on brown paper) and printed ‘Origin.’

(on pink paper); |

e 6d, 62 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper); one of the male specimens also

has a small handwritten date label ‘24/5’ (on white paper); one

of the male specimens also has a handwritten (Staudinger) label

‘Sieversi var.? | Haberhaueri | Stgr.’ with insertion in pencil

‘Namangan’ (on white paper);

e 6 with labels: ‘Prov. Samark.[and] | Hbhr. [Haberhauer]

[18181 (on brown paper) and printed ‘Origin. (on pink

paper);

e 5,2% with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper).

helena Staudinger, 1862

‘Lycaena Helena’ Staudinger, 1862: 265. Type locality: ‘...liegt mir in fünf, theilweise

gut erhaltenen Paaren vom Taygetos-Gebirge, aus dem siidlichen Theil des Peloponnes

vor, wo sie Anfangs Juni in einer Höhe von etwa 6-7000° durch Dr. Krüper erbeutet

wurde’.

Syntypes d, 2%:

e 3 with labels: handwritten ‘Taygetos | Kr.[üper]’ (on yellow

paper) and printed ‘Origin.’ (on pink paper);

e 22 with circles of yellow locality label paper and printed

label ‘Origin. (on pink paper); one has a small date label

SiG.

The specimens are placed under a handwritten drawer label

‘Helena | Stgr.’ (on white paper, in box).

253

helius Herrich-Schäffer, [1852]

‘[Polyommatus] Helius Kad. Herrich-Schäffer, [1852] 6(55): 32. Type locality: ‘Aus

Kleinasien.” — Junior objective synonym of Polyommatus asabinus Herrich-Schäffer,

[1851] (see above).

Types: not found.

heringi Kardakoff, 1928

‘Cyaniris argiolus L. heringi (gen. aest. nov.) Kardakoff, 1928: 272, Taf. 7, Figs 20, 21.

Type locality: ‘...ich sammelte ihn hauptsächlich in der Umgegend von Wladiwostok

im 2/2 V. [second half of May]. — Infrasubspecific.

Syntypes 20,22:

e G, % with labels: printed, with handwritten insertion

“Wladiwostok | S. Ussurigebiet | 7.V. (15.V. in female)192/ (N.

Kardakoff’ (on white paper), handwritten (Kardakoff?)

‘Cyaniris argiolus L. | v. heringi Kardakoff’ (on white paper)

and printed “Typus’ (on red paper, in box) — these male and

female specimens are figured in Kardakoff (1928: Taf. 7,

Fig. 20 and 21 respectively);

e G, $ with labels: printed, with handwritten insertion

“Wladiwostok | S. Ussurigebiet | 70.V. (21.V. in female)192/

(N. Kardakoff’ (on white paper), handwritten (Kardakoff?)

‘Cyaniris argiolus L. | g. v. heringi Kard. er >

N. Kardakoff” (on white paper), printed ‘Nachträgl. der | Coll.

Piingeler | hinzugefiigt’ (on greyish paper) and printed

‘Cotypus’ (on red paper, in box).

hispana Herrich-Schaffer, [1851]

‘[Lycaena| Corydon var. Hispana’ Herrich-Schäffer, [1851] 1(49): Tab. 104,

Figs 500-501; [1852] 6(55): 27. Type locality: ‘Exemplare aus Spanien sind auffall-

end klein und lebhaft gefarbt.’

‘Lectotype’ d:

e 6 with labels: handwritten ‘hispana’ (on white paper, in ball

pencil), printed ‘Coll. Led.’ (on white paper), printed ‘Zool.

Mus. | Berlin’ (on pale yellow paper), handwritten (Schurian)

on printed lined form ‘Polyommatus | (Lysandra) | hispanus H.-

S. 1852 | det. Schurian 1988’ (on white paper), handwritten

254

(Schurian) on printed form ‘Gen. Präparat | Nr. 206/1984 |

Schurian’ (on white paper), handwritten (Schurian) on printed

form ‘Lectotypus d | Lycaena hispana | H.-S. | (1852) | teste.

SCHURIAN, 1984’ and ‘Lectotypus 3 <written over printed

‘Paratypus’> | /Lycaena] | corydon hispana | H.-S. 1852 |

design. Schurian, 1988? (on red paper, in box) — Schurian

(1988: 132).

‘Paralectotypes’ 3 d :

e 6 with labels: handwritten ‘hispana’ (on white paper, in ball

pencil), handwritten (Staudinger) ‘Barcelona’ (on bluish-green

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Schurian) on printed lined form ‘Paralectotypus

3 | [Lycaena] | corydon hispana | H.-S. 1852 | det. Schurian

1988’ (on white paper), handwritten (Schurian) on printed form

‘Gen. Präparat | Nr. 207/1984 | Schurian’ (on white paper),

handwritten (Schurian) on printed form ‘Para- | Lectotypus ¢ |

Lysandra hispana H.-S. | (1852) | teste. SCHURIAN, 1984’ (on

red paper);

e 26 with circles of bluish-green locality label paper and hand-

written (Schurian) labels on printed lined form ‘Paralectotypus

3 | [Lycaena] | corydon hispana | H.-S. 1852 | det. Schurian

1988’ (on white paper) — Schurian (1988: 133).

The paper of Schurian (op. cit.) contains no evidence that a lecto-

type (and paralectotypes) have been selected from Herrich-

Schaffer’s syntypes. — The lectotype designation is unavailable

and the specimen designated as such by Schurian (1988: 132) loses

its status of lectotype (ICZN Art. 74.2).

hopfferi Herrich-Schäffer, [1851]

‘[Lycaena] Hopfferi’ Herrich-Schaffer, [1851] 1(49): Tab. 107, Figs 512-514; [1851]

6(48): 24. Type locality: ‘Aus Kleinasien; von Herr Kindermann’.

Syntypes d, 9:

e 6 with labels: handwritten ‘Hopfferi | Kind.[ermann] ms.

HSch. f | 512-14. Gerh. t. 21. f. 2 | Amasia Kind[ermann]’

(recto yellow, in printed box, verso white), printed ‘4721’

(on white paper);

255

Catalogus Generalis: 4721 — see Appendix; cf. Häuser &

Eckweiler (1997: 80).

The specimens are provided now with a printed label “hopfferi

Herrich-Schäffer, [1851] | SYNTYPUS | Y. Nekrutenko det.

30.11.1998’ (on yellow paper).

hunza Groum-Grshimailo, 1890

‘Lycaena Hunza Gr. Gr.’ Groum-Grshimailo, 1890: 397, pl. 15, fig. 2. Type locality:

‘La Lyc. Hunza vole tout le long de la rivière Mazar, qui prend sa source dans les monts

Moustag (Hımalaya occidental), sur le col Beik et un peu au-dessous de celui-ci dans les

monts Kounjout (la elle est moins typique). Altitude de 12,700 p. (Kourgan-Goudjabai et

l’embouchure du Mazar), à 15,500 p. (Beik). ... Juin et commencement de Juillet.’

Syntypes d, 9:

e 6 with labels: handwritten (Groum-Grshimailo, in Russian)

‘Ak Su’ | 20.7.[18]87’ (on white paper), printed “Transalai’ (on

white paper faded yellow, in box), printed ‘Grumm-Grsh.’ (on

white paper, in box), handwritten (Püngeler) ‘hunza & | Cotype

<underlined> | 11./[19]09 ex coll. | Dieckmann’ (on white

paper) and handwritten (Püngeler) on printed form ‘Cotype

<underlined> | hunza Gr.Gr. à | Püngeler’ (on pink paper);

‘Transalai’ (on white paper faded yellow, in box), printed

‘Grumm-Grsh.’ (on white paper, in box), handwritten

(Püngeler) ‘hunza 2 | Cotype <underlined>, 11./[19]09 | ex

coll. | Dieckmann’ (on white paper) and handwritten (Püngeler)

Piingeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Stoliczkana Feld. | v. Hunza Gr. Gr.’ (on white paper, in

printed box).

hyacinthus Herrich-Schaffer, [1847]

‘Lycaena Hyacinthus’ Herrich-Schäffer, [1847] 1(26): Tab. 72, Figs 345-348; [1852]

6(55): 29. Type locality: ‘Aus der Türkei’.

Types: not found.

256

hyrcana Lederer, 1869

‘Lycaena Hyrcana Led. Lederer, 1869: 78, Taf. 4, Figs 6, 7. Type locality: ‘...bei

Astrabad in Persien...’ (op. cit.: 73). — Junior primary homonym of Lycaena hyrcana

Felder, 1860; replacement name: Cupido morgiana Kirby, 1871: 369.

Lectotype d:

e d with labels: handwritten ‘Lycaena | Hyrcana’ (on white

paper), ‘Hadschyabad’ (on white paper), printed ‘Origin. (on

pink paper), printed ‘Coll. Led.’ (on white paper), handwritten

(Nekrutenko) on printed form ‘LECTOTYPUS & | hyrcana

Led. | Y. Nekrutenko design. (on yellow paper) — Nekrutenko

in Hesselbarth er. al. (1995: 612).

Paralectotypes 4,3%:

e 3,32 with same handwritten locality label ‘Hadschyabad’ and

printed ‘Origin’ (on pink paper), one paralectotype female with

printed ‘Coll. Led.’ (on white paper) but without locality label,

handwritten (Nekrutenko) on printed form ‘PARALECTOT Y-

PUS 3 (®) | hyrcana Led. | Y. Nekrutenko design.’ (on yellow

paper).

hyrcana Neuburger, 1903

“Chrysophanus ochimus H.-S. var. hyrcana 3 Neubgr. Neuburger, 1903: 59. Type

locality: “Habitat: Hyrcania (Persia septentr.)’.

Syntype &:

e JS with labels: handwritten ‘Persia | 3’ (on white paper faded

yellow), handwritten (Piingeler) ‘Chrysophanus | ochimus | v.

hyrcana Neubgr.’ (on white paper), a small square piece of

white paper, printed ‘Type’ (on red paper).

iberica Staudinger, 1901

‘Zephyrus Dalm. Quercus L. v. Iberica Stgr. Staudinger, 1901: 71. Type locality:

‘Ib.[eria] c.[entralis] et m.[eridionalis], Maur.[itania]’.

Syntypes 24, 29:

e à with labels: handwritten (Staudinger) ‘Quercus var. | Iberica

| Stgr.’ (verso), (on white paper), handwritten ‘Sn. Ildef.[onso] |

297

18[84] m.[ihi]’ (on pale green paper) and small date label ‘27/7’

(on white strongly faded brownish paper);

e 3,2% with circles of yellow locality label paper and small date

labels ‘10/7’ (3), ‘25/7’ and ‘28/7’.

Likewise most of the Staudinger species-group taxa becoming

available after his death in 1900 (i. e. in Staudinger, 1901), the type

specimens were not designated with characteristic ‘Origin.’ labels.

As may be seen from the original description, the type locality

statement was pretty broad, so that all forementioned specimens are

labelled such as to fit Staudinger’s meaning of the ‘terra typica’ to

constitute a syntype series.

The specimens are provided now with a printed label ‘iberica

Staudinger, 1901 | SYNTYPUS | Y. Nekrutenko det. 30.11.1998’

(on yellow paper).

ignitus Herrich-Schaffer, [1846]

‘Polyomm.{atus] ignitus Bisch. Herrich-Schäffer, [1846] 1(13): 163. [1846] 1(14):

Tab. 69, Fig. 332; Type locality: “Von Hrn. Wagner, von der Südseite des Ararat.’ —

Junior subjective synonym of Polyommatus thetis Klug, 1834 (Staudinger, 1871a: 8;

19072):

Types: not found.

ignorata Staudinger, 1901

‘Lycaena F. Meleager Esp. v. Ignorata Stgr. (Versicolor Stgr. in 1.)’ Staudinger, 1901: 86.

Type locality: “Taur.[us] m.[eridionalis] or.[ientalis] (Akbes).’ — Junior subjective

synonym of Lycaena meleager versicolor Heyne, [1895] (Hesselbarth er al., 1995: 687).

Syntypes 68,4:

e 5 with labels: handwritten (Staudinger) ‘v. Ignorata | Stgr. |

Versicolor in |. (recto) | v. Versicolor | Stgr. in l. (verso)’ (on

white paper), handwritten ‘Eibes | 18[94] Man.[issadjian]’ (on

yellow paper) and printed ‘Origin.’ (on pink paper);

e 6 with circle of yellow locality label paper, small date label

‘11/6’ and printed label ‘Origin.’ (on pink paper);

e 3 with labels: handwritten ‘Eibes | 18[94] Haradj.[?]’ (on yel-

low paper) and printed ‘Origin. (on pink paper);

258

e 264 with circles of yellow locality label paper and printed label

“Origin. (on pink paper);

e 4° with circles of yellow locality label paper and printed label

‘Origin.’ (on pink paper), one has a small date label ‘17/8’;

e 6 with labels: handwritten ‘Asia min. | (Hadjin <sic> | Funke

[18]91’ (on white paper, in printed box) and handwritten

(Piingeler) on printed form ‘Cotype <underlined> | v. ignorata

Stgr. 3 | vom Autor | Püngeler’ (on pink paper).

iliensis Staudinger, 1886

‘Pol.[yommatus| Phoenicurus Led. var. Iliensis Stgr” Staudinger, 1886: 203. Type

locality: ‘...vom Kuldja-District...’ (see also Rühl in Rühl & Heyne, [1893]: 207).

Beetotype 3:

e 6 with labels: brown paper circle, printed ‘Origin. (on pink

paper), printed ‘ex collect. | Staudinger’ (on photographic

paper) and printed “Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Nekrutenko) on printed form in box ‘Figurat | par

Y. Nekrutenko in: | Vestn. zool. 1984, 6: | tab. ex adv. p. 49’ (on

white paper), handwritten (Nekrutenko) label ‘iliensis Stgr. |

Lectotypus | Y. Nekrutenko design. | 10.IX.1983’ (on yellow

paper) — Nekrutenko (1984: 47, pl. [2]).

Paralectotypes 64 , 29:

e 3 with labels: handwritten (Staudinger) ‘var. Iliensis | Stgr.’ (on

white paper), handwritten ‘Kuldja | 85 Rckbl. [Rückbeil]’ (on

brown paper), printed ‘Origin’ (on pınk paper);

e 3,2% with circles of brown locality label paper and printed

“Origin. (on pınk paper);

e 6 with labels: handwritten (Staudinger) ‘Phoenicurus var. |

Ihensis IStgr. (on white paper), handwritten ‘Kuldja |

Alph.[éraky]’ (on brown paper), handwritten (Alphéraky) date

label ‘13/5’ (on white paper, in pencil) and printed ‘Origin’ (on

pink paper);

e 34 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper); one has also handwritten (Alphéraky)

date label ‘13/5° (on white paper, in pencil).

259

iphicarmon Eckweiler & Rose, 1993

‘Polyommatus (Agrodiaetus) iphigenia iphicarmon ssp. nov. Eckweiler & Rose, 1993:

360, Abb. 13-19. Type locality (holotype): ‘Türkei, Isparta, Egridir, Aksu, Dedegöl

Dag, 1500-1800 m’.

Paratypes 2 d :

e 24 with printed labels: “Türkei, Isparta | Egridir/Aksu |

Dedegöl Dag | 1500-1800 m | 23.-25.VII.1991 | leg. Eckweiler’

(on white paper) and ‘Paratypus 9 | Polyommatus (Agrodiae-

tus) | iphigenia iphicarmon | ECKWEILER & Rose 1993’ (on red

paper) — see also Häuser & Eckweiler (1997: 81); Eckweiler

& Häuser (1997: Pl. 8).

iphicles Staudinger, 1886

‘Lyc.{aena] Iphicles Stgr. Staudinger, 1886: 227. Type locality: ‘...(Haberhauer sandte

sie mir als Jolas), erhielt ich nur 3 im Alai (am 13. Juni) gefangene à , ein am 10. Juni bei

Osch gefangenes d und 2 am 6. Juli bei Samarkand (in den Gebirgen) gefangene 2. —

Junior subjective synonym of Lycaena gigantea Groum-Grshimailo, 1885 (Staudinger,

1901: 88).

Syntypes 46 , 29:

e d with labels: handwritten (Staudinger) ‘Iphicles | Stgr.’ (on

white paper), handwritten ‘Alai | [18]85 Hbhr. [Haberhauer]’

(on brown paper), printed ‘Origin. (on pink paper);

e 25 with circles of brown locality label paper and printed label

‘Origin. (on pink paper); one of them has a handwritten

(Haberhauer?) label ‘Iolas 16/6’ (on white faded paper, in vio-

let ink) — see Type locality section above;

e 3 with labels: handwritten ‘Osch | Hbhr. [Haberhauer]’ (on

brown paper), handwritten ‘32, handwritten date label *10/6’

and printed ‘Origin. (on pink paper);

[Haberhauer] [18]81’ (on brown paper), and printed ‘Origin.’

(on pink paper);

‘6/7’ (on white paper, in pencil) and printed label ‘Origin. (on

pink paper).

260

iphidamon Staudinger, 1899

‘[Lycaena] Damone Ev. var. Iphidamon’ Staudinger, 1899: 144. Type locality: ‘bei

Stücken aus Nord-Persien’; restricted by the lectotype designation: “Schahkuh’.

Lectotype d:

e 3 (abdomen missing), with labels: handwritten ‘Schakuh |

Chr.[istoph]’ (on yellow paper), a circle of green paper, printed

‘315. (on white paper faded brown), handwritten (Forster) on

printed form ‘Lecto-Typus à | Polyommatus iphigenia | iphida-

mon Stgr. | W. Forster München’ (on pink paper) — Forster

(1960: 118); Häuser & Eckweiler (1997: 81); Eckweiler &

Hauser, 1997: Pl. 8).

Paralectotypes 84 , 29:

e 3 with labels: handwritten ‘Hadsch. | Chr.[istoph]’ (on yellow

paper), date label ‘26/6’ (on dark-green paper) printed ‘316.’

(on white paper faded brown), handwritten (Forster)

‘Cotypus 3 | Lycaena damone Ev. | iphidamon Stgr.’ (on pink

paper);

e d with a circle of yellow locality label paper and printed

‘Origin.’ (on pink paper);

e JS with labels: handwritten ‘Schakuh | Chr.[istoph]’ (on yellow

paper), printed ‘311. (on white paper faded brown), handwrit-

ten (Forster) on printed form ‘Polyommatus | (Agrodiaetus) à |

iphigenia | iphidamon Stgr. | det. W. Forster 1947’ (on white

paper);

‘312. (on white paper faded brown), printed, with handwritten

insertion ‘Coll. Led. | Iphigenia’ (on white paper), handwritten

(Forster) on printed form ‘Polyommatus | (Agrodiaetus) 9 |

iphigenia | iphidamon Stgr. | det. W. Forster 1947’ (on white

paper);

e ¢ (abdomen missing), with labels: handwritten ‘Hadsch. |

Hbhr. [Haberhauer]’ (on yellow paper), printed ‘317. (on white

paper faded brown), printed ‘Origin.’ (on pink paper), hand-

written (Forster) ‘Cotypus d | Lycaena damone Ev. | iphidamon

Stgr.’ (on pink paper);

261

e d with labels: handwritten ‘Astrabad’ (on white paper), printed

‘310. (on white paper faded brown), printed ‘Coll. Led. with

insertion ‘Damon’ (on white paper), printed ‘Origin.’ (on pink

paper), handwritten (Forster) ‘Cotypus d | Lycaena damone Ev.

| iphidamon Stgr.’ (on pink paper);

e & with labels: handwritten ‘Hyrcania | Chr.[istoph] (on yellow

paper), printed ‘Origin. (on pink paper), printed ‘309. (on

white paper faded brown), handwritten (Forster) “Cotypus |

Lycaena | damone | iphidamon Stgr.’ (on pink paper);

label *1/6’ (on white paper), small date label ‘17/6’ (on white

paper faded brown), printed ‘313. (on white paper faded

brown), printed ‘Origin. (on pink paper), handwritten (Forster)

‘Cotypus 3 <sic> | Lycaena damone | Ev. | iphidamon Stgr.’ (on

pink paper), handwritten (Forster) on printed form ‘Ne- Allo-

Typus <corrected Type>, ? | Polyommatus (A grodiaetus) | iphi-

genia iphidamon | Stgr. | W. Forster München’ (on pink paper);

e 24, one with labels: handwritten ‘Hadschyabad’ (on white

paper), printed, with handwritten insertion ‘Coll. Led. |

Iphigenia’ and printed ‘Origin.’ (on pink paper), another with a

circle of yellow locality label paper and printed ‘Origin.’ (on

pink paper) is placed at the series end under a handwritten

(Staudinger) drawer label ‘ab. Transcaspica | Stgr.’ (on white

paper).

iphigenia Herrich-Schaffer, [1847]

‘Lycaena Iphigenia Friv.[aldszky]’ Herrich-Schäffer, [1847] 1(26): Tab. 73, Fig. 354;

[1851] 6(48): 24. Type locality: ‘Aus der Türkeï’.

Syntypes2eom2

e 3 with labels: handwritten ‘Iphigenia | Friv.[aldszky] HSch. f.

354 | Gerh.[ard] | t. 20. f. 2. Frey.[er] | Constantinop.[el]

Friv.[aldszky]’ (on white paper, in printed box) and printed

‘4724’ (on white paper);

e 4,29 without any label.

Catalogus Generalis: 4724 — see Appendix; cf. Häuser &

Eckweiler (1997: 81).

262

The specimens are provided now with a printed label ‘iphigenia

Herrich-Schäffer, [1847] | SYNTYPUS | Y. Nekrutenko det.

11.12.1998’ (on yellow paper).

iphigenides Staudinger, 1886

‘Lyc.{aena| Kindermanni Ld. ... var. Iphigenides Stgr. Staudinger, 1886: 213. Type

locality: ‘nur 4 reine d erhielt, von denen 3 Ende Mai bei Namangan, das vierte in den

Gebirgen bei Margelan gefunden ist’; restricted by the lectotype designation: Namangan.

Lectotype d:

e G with labels: handwritten (Staudinger) ‘var. Iphigenides |

Stgr.” (on white paper), handwritten ‘Namangan | Hbhr.

[Haberhauer] [18]84’ (on brown paper), printed ‘Origin. (on

pink paper), handwritten (Forster) on printed form ‘Lecto-

Typus & | Polyommatus (Agrodiaetus) | iphigenia iphigenides

Stgr. | W. Forster München’ (on pink paper) — Forster (1960:

122); Häuser & Eckweiler (1997: 82).

Paralectotypes 36:

e ¢ with labels: handwritten ‘Margelan | Maur.[er] [18]84’ (on

brown paper) and printed ‘Origin. (on pink paper);

e d with labels: a circle of locality label brown paper, printed

‘Origin.’ (on pink paper) handwritten (Forster) ‘Cotypus & |

Lycaena kinder- | manni | var. iphigenides | Stgr.’ (on pink

paper), handwritten (Forster) on printed form ‘Polyommatus |

(Agrodiaetus) | iphigenia | iphigenides Stgr. | det. W. Forster

1947’ (on white paper).

iris Staudinger, 1886

‘Lyc.[aena] Iris Stgr.’ Staudinger, 1886: 207. Type locality: ‘aus den Gebirgen bei

Margelan und Samarkand, wo sie im Juli gefanden wurde.’ — Junior primary homonym

and junior objective synonym of Lycaena iris H. C. Lang, 1884; replacement name

Polyommatus neoiris Tshikolovets (1997a: 139) is a junior objective synonym of Lycaena

iris H. C. Lang, 1884.

Syntypes 94, 39:

e d with labels: handwritten (Staudinger) ‘Iris | Stgr. (on white

paper), ‘Margelan | Hbhr. [Haberhauer]’ (on brown paper) and

‘8/7’ (on white paper) and printed ‘Origin.’ (on pink paper);

263

34 with handwritten date labels, ‘10/7 [18]80’, ‘16/7’, ‘12/7’

(on white paper) and printed ‘Origin.’ (on pink paper);

? with circle brown locality label paper and printed label

‘Origin.’ (on pink paper);

dé with labels: handwritten ‘Prov. Samark.[and] | Hbhr.

[Haberhauer] [18]81’ (on brown paper), small handwritten date

label ‘20/7’ (on white paper) and printed ‘Origin. (on pink

paper);

48,29 with circles of brown locality label paper, printed label

“Origin. (on pink paper), and some with small date labels

‘12/V”, ‘6/5’ (on white paper), (figured in Tshikolovets, 1997a:

139, 238, pl. 40, figs 15(17; 1997b: 37, figs 7-10); one male

specimen also has a handwritten (Staudinger) label ‘Iris’ (on

white paper).

isaurica Staudinger, 1871

‘Lycaena Isaurica Lederer in coll.’ Staudinger, 1871b: 327. Type locality: ‘[Lederer]

fing

diese Stücke [44 ] bei Karli Boghas im Taurus, im südlichen Kleinasien’.

Syntypes 3d, 2:

264

6 with labels: handwritten (Lederer) ‘Isaurica | m.[ihi]’ (on

white paper, in printed box), handwritten “Karli Boghas’ (on

white paper), printed ‘Origin.’ (on pink paper); printed ‘Zool.

Mus. | Berlin’ (on pale yellow paper), printed, with handwritten

(Wagener) insertions ‘Abgebildet in Hesselbarth | van Oorschot

& Wagener: | Tagfalter der Türkei. | Tafel 705 Figur 7’ (on white

paper);

d with labels: handwritten ‘Karli Boghas’ (on white paper),

printed ‘Coll. Led.’ (on white paper), printed ‘Origin.’ (on pink

paper);

6 with labels: handwritten “Taurus | Ld. [Lederer]’ (on yellow

paper), printed ‘Origin.’ (on pink paper);

® with labels: handwritten ‘Karli Boghas’ (on white paper),

printed ‘Coll. Led.’ (on white paper), printed ‘Zool. Mus. |

Berlin’ (on pale yellow paper), handwritten (Wagener)

‘Lycaena | isaurica | Stgr. 1871’ (on white paper), printed, with

handwritten (Wagener) insertions ‘Abgebildet in Hesselbarth |

van Oorschot & Wagener: | Tagfalter der Türkei. | Tafel 705

Figur 7’ (on white paper).

ismenias Klug, 1834

‘Lycaena Ismenias’ Klug, 1834, Decas Quarta: Signature k[in], Taf. 40, Abb. 1, 2. Type

locality: ‘Prope Ambukohl mense Iulio semel capta’.

Types: not found; Catalogus Generalis: 4091 — see Appendix.

Jordanus Staudinger, 1897

‘Jolaus Jordanus Stgr. n. sp. Staudinger, 1897: 153, Taf. 5, Fig. 5. Type locality:

“...wurde vor einigen Jahren von Herrn Paulus im Jordanthal (Paläastina) entdeckt...

Syınypes 28,28:

e 6 with labels: handwritten (Staudinger) ‘Jolaus ? | Jordanus |

Stgr. (on white paper), handwritten ‘Jordanthal | [18]95 Paulus’

and printed “Origin. (on pink paper);

e 6,22 with printed ‘Palaes- | tina’ (on white paper) and printed

‘Origin.’ (on pink paper); one female specimen also has a hand-

written label (Staudinger) ‘Jolaus | Jordanus 2’ (on white

paper).

Juennana Forster, 1940

‘Glaucopsyche (Phengaris) atroguttata juennana ssp. nov. Forster, 1940: 877. Type

locality (holotype): ‘China, Jünnan, Umgebung von Tali’.

Holotype d:

e 5 with labels: printed, with handwritten insertion ‘China

Yunnan | Tali | 6.9.[19]/4, | R. Mell’ (on white paper), hand-

written (Forster) on printed form ‘Holo-Type | Phengaris

atroguttata | ssp. yuennana <sic> Forst.’ | W. Forster München’

(on pink paper), handwritten (Mell) field label “Tali <illegible>

| 6.1X.[19]14 | <illegible> (recto) | <text in Chinese> (verso)’

(on poor quality paper, in pencil, folded).

265

Allotype © :

Yunnan | Tali | 23.9.[19]74, | R. Mell’ (on white paper), hand-

written (Forster) on printed form ‘Allo-Type | Phengaris

atroguttata | ssp. yuennana <sic> Forst.’ | W. Forster München’

(on pink paper), handwritten (Mell) field label “Tali (J. 3 <illeg-

ible>) | 23.1X.[1914] <illegible> (recto) | <text in Chinese>

(verso)’ (on poor quality paper, in pencil, folded).

Paratypes 36:

e 3d with labels: printed, with handwritten insertion ‘China

Yunnan | Tali | 3., 7., 11.9.[19]74, | R. Mell’ (on white paper),

handwritten (Forster) on printed form ‘Para-Typus | Phengaris

atroguttata | ssp. yuennana <sic> Forst. | W. Forster München’

(on pink paper); the paratypes also have handwritten (Mell)

original field labels (on poor quality paper, folded).

For paratypes in Zoologisches Forschungsinstitut und Museum

Alexander Koenig, see also Nekrutenko (1998: 133).

Juldusa Staudinger, 1886

‘Lyc.[aena] Kindermanni Ld. var. Juldusa Stgr. Staudinger, 1886: 213. Type locality:

“...von Alpheraki auf dem Juldus- (Tian Schan) Gebirge gefangene...’

Lectotype d:

e d with a circle of brown locality label paper, printed ‘Origin.’

(on pink paper), handwritten (Forster) on printed form

‘Polyommatus | (Agrodiaetus) | iphigenia à | juldusa Stgr. | det.

W. Forster 1947’ (on white paper), handwritten (Forster) on

printed form ‘Lecto-Typus 6 | Lycaena kindermanni | juldusa

Stgr. | W. Forster München’ (on pink paper, in box), handwrit-

ten (unknown hand) ‘juldussa’ (on white paper) Forster

(1960: 124); Hauser & Eckweiler (1997: 83); Eckweiler &

Hauser (1997: Pl. 9). |

Paralectotypes 7¢, 29:

e d with labels: handwritten (Staudinger) ‘var. Juldusa | Stgr.’ (on

white paper), handwritten “Tian Schan | Alph.[eraky]’ (on

266

brown paper), handwritten date label ‘15/VII’ and printed

‘Origin.’ (on pink paper);

e 48,2% with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper), one male and one female speci-

men also have labels: handwritten (Forster) on printed form

‘Polyommatus | (Agrodiaetus) | iphigenia & (®) | juldusa Stgr.

| det. W. Forster 1947’ (on white paper) and handwritten on

printed form ‘Cotypus d (2) | Lycaena kindermanni | juldusa

Stgr.’ (on pink paper); one female specimen also has a date label

14/7’).

Juliae Kardakoff, 1928

‘Deudorix arata Brem. juliae (subsp. nov.)’ Kardakoff, 1928: 271, Taf. 7, Fig. 12. Type

locality: ‘Die von mir gesammelte Sommertiere stammen aus Wladiwostok,

Russ.[ische] Insel, Narva [Narwa] und Barabasch’.

Syntypes 2°:

S. Ussurigebiet | 23.V1.[19]21 N. Kardakoff’ (on white paper),

printed ‘Nachträgl. der | Coll. Püngeler | hinzugefügt’ (on

greyish paper), handwritten (Kardakoff?) ‘Deudorix arata

Brem. | v. juliae Kardakoff’ (on white paper) and printed

‘Typus” (on red paper, in box);

See ssumeb- wiles. Vi <erossed, corected to “6'> [19121

N. Kardakoff’ (on white paper), printed ‘Nachträgl. der | Coll.

Püngeler | hinzugefügt’ (on greyish paper), handwritten

(Kardakoff?) “Deudorix arata Brem. | v. juliae Kardakoff’ (on

white paper) and printed “Typus’ (on red paper, in box).

kenteana Staudinger, 1892

‘Lyc.{aena| Cleobis Brem. var. Kenteana’ Staudinger, 1892d: 316. Type locality:

‘Kentei’.— Junior subjective synonym of Lycaena cleobis ida Grum-Grshimailo, 1891

(Staudinger, 1901: 79).

Syntypes 4d, 29:

e d with labels: handwritten (Staudinger) ‘var. | Kenteana | Stgr.

(recto) | v. Ida GrGr. | v. Kenteana Stgr. (verso)’ (on white

267

paper), handwritten ‘Kentei | [18]89 Dörr.[ies]’ (on green paper)

and printed ‘Origin. (on white paper);

e 34,29 with circles of green locality label paper and printed

labels ‘Origin.’ (on pink paper).

kindermannii Lederer, 1852

‘Lycaena * Kindermannii Gerh.[ard]’ Lederer, 1852: 21 (nomen nudum; * designates

‘was nicht echt europäisch’ — op. cit.: 15); ‘Lycaena Kindermannii’ Lederer, 1852: 37.

Type locality: ‘...meine Kindermannii aus der Gegend von Amasia.’ — Junior

subjective synonym of Lycaena carmon Herrich-Schäffer, [1851] (Staudinger, 1899:

139):

Syntypes 26 , 9:

e 6 with labels: handwritten ‘Kindermani <sic> | Gerh.[ard] 2 3’

(on white paper), handwritten ‘Amasia’ (on white paper) and

printed, with handwritten insertion ‘Kindermani <sic> | Coll.

Led.’ (on white paper);

e d with printed label with handwritten insertion ‘Kindermanni |

Coll. Led.’ (on white paper);

with handwritten insertion‘Kindermanni | Coll. Led.’ (on white

paper).

This species-group taxon was unintentionally named as pre-

sumed to be published by Gerhard but actually it was not (cf.

Häuser & Eckweiler, 1997: 85). The first of forementioned speci-

mens has a label exactly showing its relation to Gerhard; the others

were labelled with ‘Coll. Led. after Lederer’s collection was

acquired by Staudinger, and insertions on printed labels made in

the same writing characters as accepted in the Staudinger collection

may show that they were designated in some way in the collection

of Lederer.

The specimens are placed under a handwritten (Staudinger)

drawer label ‘Carmon HS. | Kindermanni | Ld. (on white faded

paper); each provided now with a printed label ‘kindermannii |

Lederer, 1852 | SYNTYPUS | Y. Nekrutenko det. 10.11.1998’ (on

yellow paper).

268

kogistana Grum-Grshimailo, 1888

‘Lycaena kogistana m.’ Grum-Grshimailo, 1888: 306. Type locality: ‘Darwaz’.

Syntype d : |

e ¢ with labels: handwritten (Staudinger) ‘Kogistana Grum. |

Agwai-Polusali Pass | Darwas Geb.[irge].. (on white paper),

handwritten “Darwas | [18]85 Gr.[um] Gr.[shimailo]’ (on brown

paper), handwritten (Grum-Grshimailo?, in Russian) ‘Karysen

<hardly legible>’ (on white paper) and small handwritten

(Grum-Grshimailo) date label ‘4. VIT [18]85. (on white paper).

In evidence that this specimen, forwarded by Grum-Grshimailo

to Staudinger, belongs to the type series, a quotation from a subse-

quent publication by the author (redescription and type locality

restriction) follows: ‘Elle n’a été trouvée qu’une seule fois, près du

col Agwai-Polousak, où elle volait sur des clairières d’“artcha”

[Juniperus sp.] (près de 9,500 p.). Commencement de Juin’

(Groum-Grshimailo, 1890: 407, pl. 8, fig. 6a, 6b).

The specimen is provided now with a printed label ‘kogistana |

Grum-Grshimaïlo, 1888 | SYNTYPUS | Y. Nekrutenko det.

16.11.1998’ (on yellow paper).

korlana Staudinger, 1901

‘Lycaena F. Prosecusa Ersch. v. Korlana Stgr.’ Staudinger, 1901: 90. Type locality:

‘Tian.[schan] c.[entralis] m.[eridionalis] (Korla)’ ‘sec.[undum] 3 2 2 descripta’ -1. c.

Syntype ©:

e ? with handwritten (Staudinger) labels ‘Prosecusa var.? |

Korlana Stgr.’, “von Korla’ (both on white paper) and a square

piece of pink paper.

The specimen is provided now with a printed label ‘prosecusa

korlana | Staudinger, 1901 | SYNTYPUS | Y. Nekrutenko det.

10.11.1998’ (on yellow paper).

kwangtungensis Forster, 1942

‘Zephyrus kwangtungensis spec. nov. Forster, 1942: 570, Taf. 16. Type locality (holo-

type): ‘China, Kwangtung, Lung tao shan’.

269

Holotype 6:

e 3 with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao Shan | 3.5.[19]18, | R. Mell’ (on white

paper), handwritten (Forster) on printed form ‘Holo-Type <cor-

rected to “Typus’> ¢ | Zephyrus | kwangtungensis Forst. |

W. Forster Miinchen’ (on pink paper);

Paratypes 2 d :

e 24 with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao Shan | 3.8 [19178 and 7.5.[19]77 |

R. Mell’ (on white paper), handwritten (Forster) on printed

form ‘Para-Typus 6 | Zephyrus | kwangtungensis Forst. |

W. Forster München’ (on pink paper); both have also hand-

written (Mell?) original field labels (one in Chinese) (on poor

quality paper, folded).

laetifica Püngeler, 1898

‘Lycaena cyllarus Rott. var. laetifica. m. Püngeler, 1898: 57. Type locality: ‘Hab.[itat]

Ili flimen. later on redescribed as a species ‘Lycaena laetifica m. Püngeler, 1899: 95.

e 3 with labels: printed, with handwritten (Püngeler) insertion

‘Asia centr.[alis] | (Zli) | Rückbeil 1897’ (on white paper) and

handwritten (Püngeler) on printed form “Type <double under-

lined> | laetifica Püng. 3 | Püngeler’ (on pink paper);

e 6, ? with labels: printed, with handwritten (Püngeler) inser-

tion ‘Asia centr.[alis] | (Zli) | Rückbeil 1897’ (on white paper)

and handwritten (Püngeler) on printed form ‘Cotype <under-

lined> | laetifica Püng. 3 (2)! Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Cyllarus Rott. | v. Laetifica Püng.’ (on white paper, in box);

the holotype designation (as “Type’) has not been published.

lampon Lederer, 1870

‘Polyommatus Lampon Led. Lederer, 1870b: 8, Taf. 1, Figs 2, 3. Type locality: ‘...bei

Astrabad in Persien... ...von Hirten bewohnte Dorf Hadschyabad... (op. cit.: 3, 4).

270

Syntypes 23,2%:

e d with labels: handwritten “Hadschyabad’ (on white paper) and

printed ‘Origin.’ (on pink paper);

(on white paper);

e JS, 2 with labels: handwritten “Hadschyabad’ (on white paper),

printed ‘Origin. and printed ‘Eigentum | Collection |

Staudinger’ (both on pink paper).

lamponides Staudinger, 1901

‘Chrysophanus Hb. Lampon Ld. v. Lamponides Stgr. Staudinger, 1901: 73. Type

locality: ‘Pers.[ia] s.[eptentrionale] (Schahrud).

Syntypes 4d, 29:

e d with labels: handwritten ‘Schahrud | Chr.[istoph]’ (on yellow

paper), small handwritten label ‘17/5’ and printed ‘Origin. (on

pink paper);

e 34,29 with circles of yellow locality label paper.

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Lamponides | Stgr.’

latedisjuncta Alberti, 1973

‘L.[ycaena] pyrenaica latedisjuncta n. subsp.’ Alberti, 1973: 221. Type locality: ‘...mit

Holotypus d und typischer Population vom Kasbek’.— Junior subjective synonym of

Lycaena dardanus Freyer, [1843] (Hesselbarth et al., 1995: 622).

Types (‘...einige Paratypen auch im Zoologischen Museum

Berlin...’ — op. cit.: 221): not found.

lederi A. Bang-Haas, 1907

‘Lycaena lycormas Btl. v. lederi n. var. Bang-Haas, A., 1907: 69. Type locality: ‘Aus

dem Sajangebiete (Munku) erhielt ich eine Anzahl männlichen Stiicke...’

Syntypes 36:

e 3 with labels: handwritten (O. Bang-Haas) ‘Lyc.[aena]

Lycormas | v. Lederi | BH. | (Sajan M.[ontes])’ (on white paper),

271

handwritten (O. Bang-Haas) ‘Sajan | M.[ontes]’ (on white

paper) and handwritten ‘Orig.’ (on pink paper);

e 26 with labels: printed ‘Sajan’ (on white paper) and handwrit-

ten ‘Orig.’ (on pink paper).

livia Klug, 1834

‘Lycaena Livia’ Klug, 1834: Decas Quarta: Signature k(ili), Tab. 40, Figs 3, 4 <in text;

figured figs 3-6>. Type locality: [Egypt] ‘Patria: inter Kineh [Kineh, Qena] et Assuan

[Aswan] Novembre’.

Syntypes 2d:

e 3 with labels: handwritten ‘Livia | Kl.[ug] Symb.[olae]

t.[abula] 40 f.[igurae] 3-6 | Arab.[ia] fel.[ix] Ehrbg.

[Ehrenberg]’ (recto yellow, in printed box, verso white), printed

‘4117’ (on white paper);

e 3 with labels: handwritten ‘Aegypt | Ehrbg. [Ehrenberg]’ (on

dirty green paper), printed ‘4118’ (on white paper).

Catalogus Generalis: 4117, 4118 — see Appendix.

The specimens are provided now with a printed label ‘/ivia Klug,

1834 | SYNTYPUS | A. Olivier & Y. Nekrutenko det. 30.11.1998’

(on yellow paper).

lorestanus Eckweiler, 1997

‘Polyommatus (Agrodiaetus) demavendi lorestanus subsp. nov. Eckweiler, 1997: 8,

Farbtafel 1. Type locality (holotype): ‘Iran, Lorestan, Dorud, Saravand,

2000-2300 m’.

Paratypes 2d, 9:

e 23, 2 with printed labels: ‘Iran/Lorestan | Dorud/Saravand | 2.-

5.VIII.1979 | leg. GORGNER’ (on white paper) and

‘Paratypus 4 (2) | Polyommatus (Agrodiaetus) | demavendi

lorestanus | subsp. nov. | Eckweiler, 1997’ (on bluish paper, in

box) — see also Häuser & Eckweiler (1997: 87).

lorquinii Herrich-Schäffer, [1850]

‘Lyc.[aena] Lorquinii’ Herrich-Schäffer, [1850] 1(43): Tab. 92, Figs 442-444; [1852]

6(55): 25. Type locality: not stated: ‘Ein Pärchen von Herr Keferstein’.

202

Types: not found.

lucifera Staudinger, 1867

‘Lycaena lucifera Kinderm. in litt.’ Staudinger, 1867: 100. Type locality: ‘...findet sich

in nur einem männlichen Exemplar in Gruner’s Sammlung... und stammt aus Ust-

Kamenogorsk im Altai (westliches Sibirien)’.

Holotype d (by monotypy): not found (see also Forster, 1941:

Bee Var 2, Fie. 12, Taf. 3, Fig. 12).

lusitanica Staudinger, 1892

‘Laeosopis Roboris Esp. var Lusitanica Stgr. Staudinger, 1892a: 232. Type locality:

‘ein solches d am 6. Juni [18](84) bei Lissabon und besitzte noch ein anderes Stück

dort; Herr Max Korb fand diese var. Lusitanica in Anzahl und in sehr typischen Stücken

in Algarbien im südlichen Portugal’.

Syntypes 3d, 9:

e JS with labels: handwritten (Staudinger) ‘var. Lusitanica | Stgr.’

(on white paper), handwritten ‘Lusitania | Eaton’ (on yellow

paper) and printed “Origin. (on pink paper);

e 6 with labels, handwritten (Staudinger) ‘Lissabon | [18]84

m.[ihi]’ (on yellow paper), handwritten on printed form ‘Visto y

dibujado | en 1959 |R. AGENJO det.’ (on white paper), small

handwritten <illegible>, and printed ‘Origin. (on pink paper):

e 3 with labels: handwritten (Staudinger) ‘Algarbia | Korb’ (on

yellow paper) and printed ‘Origin. (on pink paper);

e ? with a small square piece of yellow paper and printed

‘Origin.’ (on pink paper).

maculifera Staudinger, 1892

‘Thestor Romanovi Christ. var. Maculifera Stgr. Staudinger, 1892a: 233. Type locali-

ty: ‘Mardin in Mesopotamien. — Junior subjective synonym of Thestor romanovi

Christoph, 1882 (Hesselbarth er al., 1995: 506).

Syntypes 6d , 79:

e 6 with labels: handwritten (Staudinger) ‘var. Maculifera | Stgr.’

(on white paper), handwritten “Mardin | [18]91 Manis.[sadjian]’

(on yellow paper), printed ‘Origin.’ (on pink paper);

213

e 54,7% with circles of yellow locality label paper and printed

‘Origin.’ (on pink paper), one of the male specimens also has a

handwritten (Staudinger) label ‘var. Maculifera | Stgr.’ with an

insertion in pencil ‘Mardin’ (on white paper), some also with

small date labels, not always legible.

mandschurica Staudinger, 1892

‘Lycaena Astrarche Brgstr. var. Mandschurica’ Staudinger, 1892b: 161. Type locality:

‘Diese Mandschurica erhielten wir in Anzahl von Herz, der sie im nördlichen China bei

Ta-schiao-sy (nördlich von Peking) von Anfang Juni bis Ende Juli fand...” — Junior sub-

jective synonym of Lycaena chinensis Murray, 1874 (Staudinger, 1901: 83).

Syntypes 34,39:

e 5 with labels: handwritten (Staudinger) ‘Myrmecias | var.

Mandschurica | Stgr.’ (on white paper), and ‘Peking | [18]85

Herz’ (on white paper), small handwritten date label ‘15/6’,

printed ‘Origin. (on pink paper);

e d with labels: handwritten (Staudinger) ‘Myrmecias var. |

Mandschurica | Stgr. (on white paper), circle of white paper,

small handwritten date label ‘9/6’, printed ‘Origin. (on pink

paper);

e 4,3% with circles of white locality label paper, small hand-

written date labels ‘8/6’, ‘10/7’, ‘28/7’ and printed labels

‘Origin.’ (on pink paper).

maraschi Forster, 1956

‘Agrodiaetus damone maraschi ssp. nov. Forster, 1956: 57. Type locality (holotype):

‘Syria sept., Taurus, Marasch’.— Junior subjective synonym of Agrodiaetus damone wag-

neri Forster, 1956 (Hesselbarth et al., 1995: 729).

Paratype à:

e ¢ with labels: handwritten “Taurus | Karli Boghas | 1895 (recto)

| Lycaena | actis HS. à | E. Funke 1.[18]96 (verso)’ (on white

paper, recto in printed box) and handwritten (Forster) on print-

ed form ‘Para-Typus & | Agrodiaetus damone | maraschi

Forster | W. Forster Miinchen’ (on pink paper, in box) — see

also Häuser & Eckweiler (1997: 88).

274

marchica Hannemann, 1928

‘Chrysophanus amphidamas marchica’ Hannemann, 1928: 110. Type locality:

‘Strausberg [bei Berlin]’.

Syntypes 34,29:

e 364,22 with labels: handwritten (Hannemann?) ‘15.7.[19]28 |

Strausberg | E. Hañemañ <sic> (on white paper) and printed

‘Paratypus’ (on pink paper).

The specimens are placed under a handwritten (Herring) drawer

label ‘Lycaena | amphidamas | marchica Hann. and (on the same

pin) printed ‘Paratype’ (on pink paper). Since the original descrip-

tion contains no holotype designation (the type material is referred

to as Typen’), these specimens are to be considered as syntypes

(subsequent designation with labels ‘Paratypus’ by Herring is

apparent).

marcida Lederer, 1870

‘Lycaena Marcida Led. Lederer, 1870b: 10, Taf. 1, Fig 8. Type locality: ‘...bei

Astrabad in Persien... Nur 2 Männchen erhalten...’ (op. cit.: 3).

Syntypes 26d:

e à with labels: handwritten (Lederer) ‘Marcida | m.[ihi]’ (on

white paper, in printed box), and ‘Hadschyabad’ (on white

paper), printed ‘Eigentum | Collection | Staudinger’ (on pink

paper), printed ‘Origin.’ (on pink paper), printed ‘Zool. Mus. |

Berlin’ (on pale yellow paper);

e d with labels: handwritten ‘Hadschyabad’ (on white paper),

printed ‘Coll. Led.’ (on white paper), printed ‘Origin.’ (on pink

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Schurian) note on a piece of paper “Hybrid: daph-

nis brandti x marcida Led. det. Schurian 1992’.

margelanica Staudinger, 1881

“Polyomm.[atus] Dimorphus Stgr. n. sp. ...var. Margelanica’ Staudinger, 1881: 282. Type

locality: ‘...die größere Margelan-Varietat, die ganz gut als var. Margelanica bezeich-

net werden kann...’

275

Lectotype à:

e d with labels: circle of brown paper, handwritten ‘30/5 80’ (on

white-yellowish paper, in black ink), printed ‘Origin. (on pink

paper), printed ‘ex collect. | Staudinger’ (on photographic

paper), printed ‘Zool. Mus. | Berlin’ (on yellow paper), hand-

written (Nekrutenko) on printed form ‘Figurat | par

Y. Nekrutenko in: | Vestn. zool. 1984, 6: | tab. ex adv. p. 48’ (on

white paper, in box), handwritten (Nekrutenko) ‘margelanica

Stgr. | Lectotypus | Y. Nekrutenko design. | 10.1X.1983’ (on

yellow paper) — Nekrutenko (1984: 44, pl. [1]).

Paralectotypes 6d , 79:

e d with labels: handwritten ‘Margelan | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin’ (on pink paper);

e 54,7% with circles of brown locality label paper and printed

‘Origin’ (on pink paper).

marpurgica Strand, 1919

‘Lycaena argus L. cum ab. marpurgica Strand n. ab.’ Strand, 1919: 18. Type locality:

‘Marburg.’ — Infrasubspecific.

Syntypes 39 :

e 39% with labels: printed ‘Deutschland | Marburg i.[n] H.[essen]

| E. Strand, 7.04[1904].’ (on white paper), handwritten (Strand)

‘Lycaena | argus L. | Strand det. | ab. marpur- | gica m 2’ (on

stiff white paper, printed line is glued on the middle part of the

label); one specimen has a printed label ‘Type’ (on dirty-pink

paper), two other have the same printed labels, with handwrit-

ten (Strand) insertion ‘Co-Type’.

mauretanica Staudinger, 1892

‘Thecla Ilicis Esp. v. Mauretanica Stgr.’ Staudinger, 1892c: 279. Type locality: ‘Ein d

von Tunis stimmt mit Stiicken aus Collo (Prov. Constantine)...’

Syney Pes! omnes

e ¢ with labels: handwritten (Staudinger) ‘var. | Mauretanica |

Stgr.’ (on white paper), handwritten “Tunis | [18]91 Vaulog.’ (on

white paper) and printed ‘Origin.’ (on pink paper);

276

e d with labels: handwritten ‘Collo | Seriz’ (on yellowish paper)

and printed ‘Origin.’ (on pink paper);

e 34, 2 with circles of yellowish locality label paper and print-

ed labels ‘Origin. (on pink paper).

maxima O. Bang-Haas, 1927

‘Lycaena cyane maxima O. B.-Haas, subsp. nov.’ O. B.-Haas, 1927: 52, Taf. 7, Figs 24,

25. Type locality: ‘China sept. or.: Kansu, Lanchowfu’.

Syntypes à, 9:

e ¢ with labels: handwritten (O. Bang-Haas) ‘Lyc.[aena] cyane |

maxima O B Haas’ (on white paper), printed ‘Lanchowfu |

Kansu’ (on white paper), printed, with handwritten insertions

‘Horae Macrolep. | Vol. I abgebildet | t. 7, f. 24 | beschrieb.

p. 52’ (on white paper), printed “Type | O. Bang-Haas’ (on pink

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper);

printed, with handwritten insertions ‘Horae Macrolep. | Vol. I

abgebildet | t. 7, f. 25 | beschrieb. p. 52’ (on white paper), print-

ed “Zool. Mus. | Berlin’ (on pale yellow paper).

maxima Staudinger, 1901

‘Cigaritis B. Cilissa Ld. v. Maxima Stgr. Staudinger, 1901: 76. Type locality:

‘Kurd.[istan] oc.[cidentalis] (Malatia); Mesop.[otamia] s.[eptentrionale] (Mardin)’.

Syntypes 54,39:

e d with labels: handwritten (Staudinger) ‘v. Maxima | Stgr.’ (on

white paper), handwritten ‘Malatia | [18]84 Man.[issadjian]’

(on yellowish paper), and small handwritten ‘18/6’;

e 23,22 with circles of yellowish locality label paper and small

date labels ‘24/6’, ‘16/6’, ‘16/6’ and ‘19/6’;

e d with handwritten label ‘Mardin | [18]91 Manis.[sadjian]’ (on

yellow paper);

e 3, 2 with circles of yellow locality label paper.

The specimens are provided now with printed labels ‘cilissa

maxima | Staudinger, 1901 | SYNTYPUS | Y. Nekrutenko det.

1998’ (on yellow paper).

BTIT!

mayencis Eitschberger & Steiniger, 1975

‘Eumedonia eumedon mayencis n. subsp. Eitschberger & Steiniger, 1975: 106,

Figs 13-24. Type locality: ‘Deutschland, Rheinland-Pfalz, Mainz-Mombach, Mainzer

Sand’.

Paratypes 36:

e 34 with labels: printed, with handwritten insertions “Germania

Mainz | Großer Sand | 26.5.[19]74 | leg. Schurian’ (on white

paper), printed 'PARATYPUS o <sex not stated> | Eumedonia

eu- | medon mayencis | EITSCHBERGER & | STEINIGER,

1975’ (on deep red paper).

melania Staudinger, 1886

‘Lyc.[aena] Kindermanni Ld. ... var. Melania Stgr. Staudinger, 1886: 213. Type locali-

ty: ‘...nur 2 frische d aus dem südlichen Alai erhielt...’

Syntypes 2 d :

e d with labels: handwritten (Staudinger) ‘var. Melania | Stgr.’

(on white paper), handwritten ‘Alai | [18185 Hbhr.

[Haberhauer]’ (on brown paper), printed ‘Origin.’ (on pink

paper), handwritten (Forster) on printed form ‘Polyommatus

(Agrodiaetus) 3 | iphigenia | iphigenides Stgr. | f. melania Stgr.

det. W. Forster 1947’ (on white paper), handwritten (Häuser) on

printed form ‘SYNTYPUS | Lycaena | kindermanni | var. |

melania | STAUDINGER | det. Chr. L. Häuser | VZ.[19]94 (on

pink paper);

e d with labels: circle of locality label brown paper, printed

‘Origin.’ (on pink paper) and handwritten (Hauser) on printed

STAUDINGER | det. Chr. L. Häuser | V/.[19]94’ (on pink paper)

— Hauser & Eckweiler (1997: 88-89); Eckweiler & Hauser

(HONFEP EL):

melantho Klug, 1834

‘Lycaena Melantho’ Klug, 1834: Decas Quarta: Signature k(iv), Tab. 40, Figs 10, 11.

Type locality: ‘Habitat in Syria” — Junior subjective synonym of Papilio spini

Fabricius, 1787 (Hesselbarth et al., 1995: 521).

278

Syntypes 2d

e ¢ with labels: handwritten label “Var. Melantho | Kl.[ug]

Symb.[olae] t.[abula] 40 | f.[igurae] 10, 11 | Syria Ehrbg.

[Ehrenberg]’ (recto yellow, in printed dotted line box, verso

white) and printed ‘4353’ (on white paper);

e 6 without any label.

Catalogus Generalis: 4353 — see Appendix.

The specimens are provided now with a printed label ‘melantho

Klug, 1829 | SYNTYPUS | A. Olivier & Y. Nekrutenko det.

30.11.1998’ (on yellow paper).

melibaeus Staudinger, 1878

‘Pol.[yommatus] Alciphron Rott. (Hipponoë Esp.), v. Melibaeus Stgr. Staudinger, 1878:

231. Type locality: ‘Amasia und Bosz-Dagh’.

Lectotype d:

e ¢ with labels: handwritten ‘Amasia’ (on white paper), printed

‘Coll. Led.’ (on white paper), printed ‘Origin.’ (on pink paper),

printed “Zool. Mus. Berlin’ (on white paper), handwritten

(Lederer) ‘V. Melibaeus | Kdm. ilit. <sic>* (on pale green paper,

in printed box), handwritten (Wagener) on printed form

‘Lectotypus | Lycaena | alciphron | melibaeus | P. S. Wagener’

(on red paper, in box) — Wagener in Hesselbarth ef al. (1995:

oar 127; Fig. 38).

Paralectotypes 3d, 2°:

e d with labels: handwritten ‘Amasia’ (on yellow paper),

printed ‘Origin. (on pink paper), printed ‘Zool. Mus. Berlin’

(on white paper), handwritten (Wagener) on printed form

‘P[ara]Lectotypus | Lycaena | alciphron | melibaeus | P. S.

Wagener’ (on red paper, in box);

e d with labels: handwritten ‘Bos Dagh’ (on white paper), print-

ed ‘Coll. Led.’ (on white paper), printed ‘Origin.’ (on pink

paper), printed ‘Zool. Mus. Berlin’ (on white paper), handwrit-

ten (Wagener) on printed form in box ‘Pl[ara]-Lectotypus à |

Lycaena | alciphron | melibaeus | P. S. Wagener’ (on red paper);

279

ed ‘Coll. Led.’ (on white paper), printed ‘Origin.’ (on pink

paper), printed ‘Zool. Mus. Berlin’ (on white paper), handwrit-

ten (Wagener) on printed form “P[ara]-Lectotypus $ | Lycaena

| alciphron | melibaeus | P. S. Wagener’ (on red paper, in box);

e JS, 2 with labels: handwritten ‘Hadschyabad’ (on white paper),

printed ‘Coll Led.’ (on white paper), printed ‘Origin.’ (on pink

paper) — these specimens, though labelled by Staudinger as

‘Origin.[ale]’, do not represent the type series;

d with labels: handwritten ‘Szavalan | Persia Chr.[istoph]’ (on

yellow paper) and printed ‘Origin.’ (on pink paper);

3, 2 with circles of yellow paper and printed labels “Coll Led.’

(on white paper) and‘Origin. (on pink paper) — these speci-

mens, though labelled by Staudinger as ‘Origin.[ale]’, do not

represent the type series.

melli Forster, 1940

‘Zephyrus melli spec. nov.’ Forster, 1940: 871, Taf. 22, Figs 4, 5, Taf. 23, Figs 4, 5. Type

locality (holotype): ‘China, Kwangtung, Tsha-yuen-shan’.

Holotype d :

e 6 with labels: printed, with handwritten insertion ‘China |

Tsha-jiu-san | | 76. V77.[19]11, | Mell S. V (on yellow paper),

printed ‘China Kwantung | R. Mell’ (on white paper) handwrit-

ten (Forster) on printed form ‘Holo-Type à | Zephyrus| melli

Forst. | W. Forster München’ (on pink paper).

Allotype © :

Kwantung | Tsha yuen shan | 27. 6. [19]71 | R. Mell’ (on white

Zephyrus| melli Forst. | W. Forster München’ (on pink paper),

handwritten (Mell) original field label ‘Te 27 V [19]11 (recto) |

<text in Chinese> (verso)’ (on white faded paper, in pencil,

folded).

280

Paratypes 6d , 29:

e 26, 2 with labels: printed, with handwritten insertion ‘China |

Tsha-jiu-san | | <different dates>: VII. [19]11,| Mell S. V’ (on

yellow paper), printed ‘China Kwantung | R. Mell’ (on white

paper), original field labels (folded), handwritten (Forster)

on printed form ‘Para-Type d (2) | Zephyrusl melli Forst. |

Dr. W. Forster München’ (on pink paper);

e 2d, ® with labels: printed ‘China, Canton | Mell S. V. (on

white paper), printed ‘China Kwantung | R. Mell’ (on white

paper), original field labels (folded), handwritten (Forster)

on printed form ‘Para-Type d (2) | Zephyrusl melli Forst. |

Dr. W. Forster München’ (on pink paper);

e d with labels: printed, with handwritten insertion ‘China

Kwantung | Tsha yuen shan | 23.6. [19]/7 IR. Mell’ (on white

paper), handwritten (Mell) original field label (folded), hand-

written (Forster) on printed form ‘Para-Type do | Zephyrusl

melli Forst. | Dr. W. Forster München’ (on pink paper);

e 3 with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao shan | 30.6. [19]77 | R. Mell’ (on white

paper), handwritten (Mell) original field label (folded), label,

handwritten (Forster) on printed form ‘Para-Type ¢d | Zephyrusl

melli Forst. | Dr. W. Forster Miinchen’ (on pink paper).

merisina Lorkovic, 1943

‘Everes argiades merisina n. Lorkovié, 1943: 450, Taf. 25, Fig. 11, Taf. 26, Figs 9, 10.

Type locality: ‘China, Kwang-tung, Gao-fung.’

Holotype &:

e d with labels: printed, with handwritten insertion ‘China |

Tsha-jıu-san | | 22. VII.[19]10, | Mell S. V. (on yellow paper),

handwritten (Forster <sic>) on printed form ‘Holo-Type <cor-

rected to Typus> 6 | Everes argiades Pall.| merisina Lorkovic’

(on pink paper), handwritten (Mell) field label ‘Te 22 VII

(recto) | <typewritten> in (verso)’ (on white paper, in pencil,

folded) and a square piece of paper with handwritten ‘la’ in

pencil (on white paper).

281

Allotype ©

Tsha-jıu-san | | 27. VIL.[19]10,| Mell S. V. (on yellow paper),

handwritten (Forster <sic>) on printed form ‘Allo-Type <cor-

rected to Typus> à | Everes argiades Pall.| merisina Lorkovic’

(on pink paper), handwritten (Mell) field label ‘Te 21 VI 1a’

in pencil (on white paper, folded).

Paratypes 154, @:

e 54 with labels: printed, with handwritten insertion ‘China |

Tsha-jiu-san | <different dates>: VII., VIII, IX [19]10,| Mell S.

V (on yellow paper), printed ‘China Kwantung | R. Mell’ (on

white paper), handwritten (Forster <sic>) on printed form

‘Para-Typus d | Everes argiades Pall.\ merisina Lorkovic’ (on

pink paper);

e 24 with labels: printed ‘China Kwantung | R. Mell’ (on white

paper), handwritten (Forster <sic>) on printed form ‘Para-

Typus d | Everes argiades Pall.| merisina Lorkovic’ (on pink

paper);

e 54, 2 with labels: printed, with handwritten insertion ‘China

Kwantung | Tsha yuen shan | <different dates> &., 9., [19]10. |

R. Mell’ (on white paper), handwritten (Forster <sic>) on print-

ed form ‘Para-Typus 9 (9) | Everes argiades Pall.| merisina

Lorkovic’ (on pink paper);

e 24 with labels: printed, with handwritten insertion ‘China

Kwantung | Man tai shan | <different dates> &., 9., [19]75. |

R. Mell’ (on white paper), handwritten (Forster <sic>) on print-

ed form ‘Para-Typus d (2) | Everes argiades Pall.| merisina

Lorkovic’ (on pink paper);

e ¢ with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao shan | 27. [19]17. | R. Mell’ (on white

paper), handwritten (Forster <sic>) on printed form ‘Para-

Typus d (2) | Everes argiades Pall.| merisina Lorkovic’ (on

pink paper).

Some paratypes have also original field labels (on poor quality

paper, folded).

282

mesopotamica Staudinger, 1892 |

‘Lycaena Poseidon Led. var. Mesopotamica Stgr. Staudinger, 1892a: 233. Type gear

ty: ‘Die von Manissadjian bei Malatia Anfangs Juli in Anzahl gefundenen Stücke...” —

Junior subjective synonym of Lycaena poseidon poseidon Herrich-Schäffer, [1851]

(Schurian et al., 1992: 217).

Syntypes 68,22:

e 3 with labels: handwritten (Staudinger) ‘Poseidon var. |

Mesopotamica | Stgr. (on white paper), circle of yellow paper,

small date label ‘2/7’ (on dirty green paper), printed ‘Origin.’

(on pink paper), printed “Zool. Mus. | Berlin’ (on pale yellow

paper), printed, with handwritten (Wagener) insertions

‘Abgebildet in Hesselbarth | van Oorschot & Wagener: |

Tagfalter der Türkei. | Tafel 778 Figur /2’ (on white paper) —

Hesselbarth et al. (1995: 726, Taf. 118, Fig. 12); Häuser &

Eckweiler (1997: 90);

e 43 with circles of yellow paper, three of these with small date

labels ‘2/7’ ‘3/7’ and ‘5/7’ (on white faded yellow paper), one,

dated °3/7’, also has a handwritten (Staudinger) label ‘Malatia’

(on white paper faded brownish); all with printed labels

‘Origin.’ (on pink paper), two with printed “Zool. Mus. | Berlin’

(on pale yellow paper), two with new, handwritten in ball pencil

‘mesopotamica’ (on white paper);

paper), small date label ‘11/7’ (on dirty green paper), printed

‘Origin.’ (on pink paper), printed “Zool. Mus. | Berlin’ (on pale

yellow paper), handwritten (Wagener) ‘Lycaena | mesopotami-

ca | Stgr. 1892 | Malatia’ (on white paper), printed, with hand-

written (Wagener) insertions ‘Abgebildet in Hesselbarth | van

Oorschot & Wagener: | Tagfalter der Türkei. | Tafel 778 Figur

49° (on white paper) — Hesselbarth et al. (1995: 726, Taf. 118,

Fig. 49 as ‘Original’ ‘Dama Stgr.’); cf. Häuser & Eckweiler

(1997: 90);

ed label ‘Origin.’ (on pink paper);

e JS with labels: handwritten (Püngeler) ‘Mesopot.[amia] (recto)

| ‘mesopotamica | 4 | vom Stgr 4.99 (verso)’ (on white paper,

283

recto in printed box), handwritten (Püngeler) on printed form

‘Cotype <underlined> | v. mesopotamica Stgr. | $ | vom Autor.

| Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler)

drawer label ‘Poseidon HS. | v. Mesopotamica Stgr.’ (on white

paper, in printed box).

minima Staudinger, 1901

‘Cigaritis B. Cilissa Ld. v. Minima Stgr. Staudinger, 1901: 76. Type locality:

‘Taur.[us] s.[eptentrionalis] (Hadjin).. — Junior subjective synonym of Cigaritis cilissa

Lederer, 1861 (Hesselbarth et al., 1995: 455).

Syntypes 3d, 9:

e d with handwritten label ‘Hadjin | [18]88 K. O. (on yellowish

paper) and small handwritten date label ‘6/7’;

e 26, 2 with same small (date) labels ‘5/7’, ‘6/7’ and ‘12/7’.

The specimens are provided now with a printed label ‘cilissa

minima | Staudinger, 1901 | SYNTYPUS 1 Y. Nekrutenko det. 1998’

(on yellow paper).

miris Staudinger, 1881

‘Lyc.[aena] Miris Stdgr. n. sp.’ Staudinger, 1881: 263. Type locality: ‘Diese neue Art

wurde zuerst von Christoph bei Schahrud (Nord-Persien am Fuße des Gebirges) Mitte

Mai gefunden; dann fand sie Haberhauer in einigen Stücken bei Saisan, später ein Stück

bei Lepsa und 1880 in einigen 20 Exemplaren bei Margelan (Central Turkestan)’.

Syntypes 128,42:

e ¢ with labels: handwritten ‘Schahrud | Chr.[istoph]’ (on yellow

paper), printed ‘Origin. (on pink paper);

e 3d, 2 with circles of yellow locality label paper and printed

label ‘Origin.’ (on pink paper);

e 3 with labels: handwritten ‘Saisan | Hbhr. [Haberhauer]’ (on

brown paper), printed ‘Origin.’ (on pink paper);

e 24 with circles of brown locality label paper and printed label

‘Origin.’ (on pink paper);

284

e 3 with labels: handwritten ‘Lepsa | Hbhr. [Haberhauer]’ (on

brown paper), printed ‘Origin. (on pink paper);

e 3 with labels: handwritten ‘Margelan | Hbhr. [Haberhauer]’ (on

brown paper), printed ‘Origin. (on pink paper);

e 34,32 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper); some specimens have handwrit-

ten date labels, not always clearly legible. |

mirza Staudinger, 1874

‘Lycaena Mirza n. sp.’ Staudinger, 1874: 90. Type locality: ‘Diese neue Art wurde

gleichfalls von Herrn Christoph entdeckt, der aber nur wenig Stücke davon fing. Ich

vom 28. Mai bis 1. Jumi bei Tasch, 2 dd am 16. Juni bei Schakuh, also auch im

nördlichen Persien, gefangen wurden.’ — Junior subjective synonym of Lycaena sieversi

Christoph, 1873 (Staudinger, 1901: 80).

Syntypes 44, 9:

e 44, 2 syntypes of sieversii Christoph, 1873.

The specimens are placed under a handwritten drawer label

‘Sieversii | Chr. | Mirza Stgr.’ (on white paper, in printed box); each

provided now with a printed label ‘sieversii Christoph, 1873 | mirza

Staudinger, 1874 | SYNTYPUS | Y. Nekrutenko det. 1.12.1998’ (on

yellow paper) — see sieversii below.

mithridates Staudinger, 1878

‘Lyc.[aena] Mithridates Stgr. n. sp.’ Staudinger, 1878: 247. Type locality: ‘Das eine

Stiick ist am 17 Juli, wahrscheinlich von mir im Kerasdere gefangen, da aber am selben

Tage Emil nach der Jenikeui-Hochebene, Zach und Johann nach dem Kyschlatschyk-

Wege waren, so bin ich meiner Sache nicht sicher’.

Syntypes 6d , 9:

e JS, pinned underside up, with labels: handwritten (Wagener)

‘Lycaena | mithridates | Stgr. 1878’ (on white paper), ‘Amasia

m.[ihi]’ (on yellow paper), printed ‘Origin. (on pink paper) and

‘Zool. Mus. | Berlin’ (on pale yellow paper), printed, with hand-

written (Wagener) insertions ‘Abgebildet in Hesselbarth | van

Oorschot & Wagener: | Tagfalter der Türkei. | Tafel //6

Figur 55’ (on white paper) — Hauser & Eckweiler (1997: 90);

285

® with labels: ‘Amasia | Joh.[ann]’ (on yellow paper), printed

‘Origin.’ (on pink paper);

e 3 with circle of yellow locality label paper and printed

“Origin. (on pink paper); |

e d with labels: ‘Amasia | Joh.[ann]’ (on yellow paper), printed

‘Origin.’ (on pink paper);

e 3d with circles of yellow locality label paper and printed

‘Origin.’ (on pink paper).

monticola Staudinger, 1901

‘Lycaena F. Alcon F. v. Monticola Stgr. Staudinger, 1901: 90. Type locality:

‘Cauc.[asus]; Helv.[etia]; Pont.[us]; ? Lyd.[ia] mont.[es]’.

Syntypes 48,2%:

e d with labels: handwritten ‘Achalzich | Habhr. [Haberhauer]

[18]82’ (on green paper) printed ‘Zool. Mus. | Berlin’ (on pale

yellow paper), handwritten “Lycaena alcon | monticola’ (on

white paper) and new printed “Typus’ (recto red, verso white);

e ? with circle of green locality label paper, handwritten

‘Lycaena alcon | monticola | Achalzich Hbhr. | [18]82’ (on white

paper) and new printed “Typus’ (recto red, verso white);

e d with labels: handwritten ‘Macugnaga | m.[ihi]’ (on brownish

paper) and new printed “Typus’ (recto red, verso white);

“Typus’ (recto red, verso white);

e ¢ (abdomen missing, pinned here to in a piece of packing paper

wrapper) with labels: handwritten ‘Valesia | Ag.[?]’ (on yellow-

ish paper) and new printed ‘Typus’ (recto red, verso white);

e G (abdomen missing) with labels: handwritten ‘Armenia |

Radde’ (on yellow paper), handwritten ‘Istissu’ (on white paper

faded yellow-brown, in pencil) and new printed “Typus’ (recto

red, verso white);

Likewise most of the Staudinger species-group taxa becoming

available after his death in 1900 (i. e. in Staudinger, 1901), the type

specimens were not designated with characteristic ‘Origin.’ labels.

286

As may be seen from the original description, the type locality

statement was pretty broad, so that all forementioned specimens are

labelled such as to fit Staudinger’s meaning of the ‘terra typica’ to

constitute a syntype series. The author of the syntype designation

with “Typus’ labels is unknown (cf. Hesselbarth er al., 1995: 576).

The specimens are placed under a handwritten drawer label

“v. Monticola | Stgr.’.

myrrha Herrich-Schäffer, [1851]

‘[Lycaena] Myrrha’ Herrich-Schäffer, [1851] 1(49): Tab. 106, Figs 508-511; [1852]

6(55): 26. Type locality: ‘Aus Kleinasien’ (cf. Staudinger, 1878: 242).

?-Syntype:

e JS with printed labels: ‘Origin. ? | ?’ (on pink paper, with hand-

written Staudinger insertions), ‘Coll Led.’ (on white paper),

‘Zool. Mus. | Berlin’ (on pale yellow paper), with handwritten

insertion ‘ZSM | Negativ | No. 107/108’ (on white paper).

One of 5 (4d, 2) specimens of the collection was designated by

Staudinger (1878: 241-242) as an apparent syntype: “Dieser ist in

Herrich-Schäffer’s Figur beim d gar nicht angegeben und auch bei

dem alten Lederer’schen Stück (vielleicht dem Original ?) nur sehr

matt erkennbar. Sonst ist die Figur gut. Die ächte Myrrha ist nur bei

Amasia und Tokat (Kindermann) gefunden. (see also Hesselbarth

et al., 1995: 666).

myrrhina Staudinger, 1901

‘Lycaena F. Myrrha HS. v. Myrrhina Stgr. Staudinger, 1901: 85. Type locality:

‘Pont.[us] or.[ientalis] Giimiischane)’.

Syntypes 4 :

e 6 with labels: handwritten (Staudinger) ‘v. Myrrhina | Stgr.’ (on

white paper), “Giimiischchane | Pont [18]94 Sint.[enis]’ (on yel-

low paper), printed “Zool. Mus. | Berlin’ (on pale yellow paper)

and ‘Origin.’ (on pink paper);

e 6 with labels: circle of yellow locality label paper, handwritten

(O. Bang-Haas?) ‘Gümüschchane d | Pont 94 Sint.[enis]’,

‘Lycaena myrrha H. S. | var. myrrhina Stgr.’ (the first was added

287

subsequently as an extension of the circle reference), printed

Bang-Haas label “Type” (on red paper), a square piece of pur-

ple-violet paper, handwritten (de Lesse) ‘de Lesse | 1946’ (on

white paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow

paper), handwritten (Wagener) ‘myrrhina | Berlin’, printed,

with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türkei. |

Tafel 711 Figur 29° (on white paper) — Hesselbarth et al.

COS Tate Dini eZ):

e JS with labels: handwritten ‘Egin | [18]84 Man.[issadjian]’ (on

yellow paper), small date label ‘4/7’, printed ‘Zool. Mus. |

Berlin’ (on pale yellow paper);

e JS with labels: circle of yellow locality label paper, handwritten

(unknown hand) ‘myrrha | myrrhina Stgr. | Egin, Türker (on

white paper); printed ‘Origin.’ (on pink paper), printed, with

handwritten insertion ‘ZSM | Negativ | No. 105/106’ (on white

paper), printed ‘Zool. Mus. | Berlin’ (on pale-yellow paper),

printed, with handwritten (Wagener) insertions ‘Abgebildet in

Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türke:. |

Tafel 771 Figur 30° (on white paper) — cf. Hesselbarth et al.

(1995: Tat: 111, Fig. 34):

myrtale Klug, 1834

‘Lycaena Myrtale’ Klug, 1834: Decas Quarta: Signature /(i), Tab. 40, Figs 15, 16. Type

locality: ‘Habitat in Syria’.

Syntypes 3d:

e 3 with labels: handwritten ‘Myrtale | Kl.[ug] Symb.[olae]

t.[abula] 40 f.[igurae] 15, 16 | Syria Ehrbg. [Ehrenberg]’ (recto

yellow, in printed box, verso white) and printed ‘4341’ (on

white paper);

e 24 without any label.

Catalogus Generalis: 4341 — see Appendix.

The specimens are provided now with a printed label ‘myrtale

Klug, 1834 | SYNTYPUS | A. Olivier & Y. Nekrutenko det.

30.11.1998’ (on yellow paper).

288

nahizerica Eckweiler, 1978

‘Aricia teberdina nahizerica ssp. n. Eckweiler, 1978: 115, Abb. 14. Type locality:

‘NO Türkei: Prov. Erzurum: Ispir: vic. Nahizer.’

Paratypes d , 49:

e 5, 49 with printed labels: ‘Türkei/Erzurum | Ispir/Nahizer |

1700-1900 m | 4.8.-8.8.[19]77 | leg. ECK WEILER’ (on white

paper), ‘Paratypus d (9) | Aricia teberdina ssp. | nahizerica |

des. ECKWEILER 1978’ (on deep red paper).

nogelii Herrich-Schäffer, [1851]

‘[Thecla] Nogelii’ Herrich-Schäffer, [1851] 1(51): Tab. 110, Figs 529-532; [1852] 6(55):

33-34. Type locality: ‘Aus Amasia’.

Types: not found (for lectotype designation see van Oorschot &

Wagener, 2000: 97-98).

oberthueri Staudinger, 1887

‘Thecla Oberthüri Stgr. Staudinger, 1887b: 138, pl. 6, fig. 4, a, b. Type locality:

“Dörries fing sie zuerst am Ussuri, später zog er sie auf Askold oder am Bykin; im vorigen

Jahr fand er sie auch am Suifun...’

Syntypes:

e 3 with labels: handwritten (Staudinger) ‘Butleri Waterh.?

<crossed out, corrected:> Fenton | Obertüri | Stgr. (on white

paper), handwritten ‘Askold | [18]83 Dörr.[ies]’ (on bluish-

violet paper), printed ‘Origin.’ (on pink paper);

e 5? with same printed labels ‘Origin’, two of these with hand-

written locality labels ‘Suifun | [18]85 Dörr.[ies]’ and ‘Ussuri |

Dörr.[ies] [18]82’ (both on bluish-violet paper), two with circles

of bluish-violet paper, one has a small handwritten label ‘13’.

oberthueri Staudinger, 1901

‘Lycaena F. Orbitulus Prun. subv.[arietas] Oberthiiri Stgr.’ Staudinger, 1901: 81. Type

locality: Pyr.[enees] c.[entrales]. — Junior primary homonym of Lycaena elvira

oberthueri Grumm-Grshimailo, 1887; replacement name: magnaglandon Verity, 1949:

134 (Kudrna, 1983: 30).

289

Syntypes 46 , 2:

e 24 with labels: printed ‘Hautes Pyrénées | Cauterets | R.

Oberthiir | Juillet 1881’ (on white paper, in box), printed

‘Origin.’ (on pink paper) and a circle of blue paper; one d with

handwritten (Staudinger) ‘subv. /Oberthüri | Stgr.’ (on white

paper);

e 6 with labels: handwritten “Pyrenais <sic> | Morel’ and print-

ed ‘Origin.’ (on pink paper);

e 6, 2 with circles of blue locality label paper and printed

‘Origin.’ (on pink paper).

obscurata Staudinger, 1892

‘Lyc.[aena] Euphemus Hb. var. Obscurata Stgr.’ Staudinger, 1892d: 320. Type locali-

ty: ‘Es liegen mir 46 4 und 59 2 vom Kentei vor...’

Syntypes 38,5%:

e 3 with labels: handwritten (Staudinger) ‘v. Obscurata’ (on

white paper), handwritten ‘Kentei | [18]89 Dörr.[ies]’ (on green

paper) and printed ‘Origin.’ (on pink paper);

e 6, 2? with circles of green locality label paper and printed

label ‘Origin.’ (on pink paper);

e d with labels: handwritten ‘Ala Tau | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

e 39% with circles of brown locality label paper and printed label

‘Origin.’ (on pink paper).

obscurata Staudinger, 1901

‘Cigaritis B. Acamas Klug v. Obscurata Stgr. Staudinger, 1901: 76. Type locality:

‘Taur.[us] s.[eptentrionalis] (Hadjin).’ — Junior subjective synonym of Lycaena acamas

acamas Klug, 1834.

Syntypes 36,29:

e 3 with labels: handwritten “Hadjin | [18]88 K. O.[?]’ (on

yellow paper), small handwritten ‘20/6’ and printed ‘Origin.

(on pink paper);

290

e 26,29 with same printed labels ‘Origin, with small hand-

written (date) labels and with circles of yellow locality label

paper.

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Obscurata | Stgr.’ (on white paper).

ochimus Herrich-Schaffer, [1851]

‘[Polyommatus| Ochimus’ Herrich-Schäffer, [1851] 1(49): Tab. 109, Figs 523-526;

[1852] 6(55): 31-32. Type locality: ‘Aus Kleinasien’.

Types: not found.

olympica Lederer, 1852

‘Lycaena Corydon Var. Olympica’ Lederer, 1852: 36, Nachtrag: 53. Type locality:

‘vom Olymp bei Brussa’.

Lectotype d:

e ¢ with labels: handwritten ‘Brussa’ (on white paper), printed

‘Coll. Led.’ (on white paper), printed “Zool. Mus. | Berlin’ (on

pale yellow paper), handwritten (Schurian) on printed lined

form ‘Polyommatus | (Lysandra) | ossmar olympica | Lederer

1852 | det. Schurian 1988’ (on white paper), handwritten

(Schurian) on printed form ‘Gen. Präparat | Nr. 2/6/1985’ (on

white paper), handwritten (Schurian) on printed form

‘Lysandra ossmar | ssp. olympica Led. stat. nov. | det. Schurian

1985’, handwritten (Schurian) ‘Lectotypus & <written over

printed ‘Paratypus’> | Lycaena | ossmar olympica | Lederer

1852 | des. Schurian, 1988’ (on red paper, in box) — Schurian

(1988: 136).

omphale Klug, 1834

‘Lycaena Omphale’ Klug, 1834: Decas Quarta: Signature k(iv), Tab. 40, Figs 12-14.

Type locality: ‘Habitat in Syria. — Junior subjective synonym of Lycaena thersamon

Esper, [1784].

Types: not found; Catalogus Generalis: 4554 — see Appendix;

only a drawer label “Var. | Omphale | Klug Symb. phys. | Syria |

Ehrbg’ (recto yellow, in dotted line box, verso white) was found.

291

orbitulinus Staudinger, 1892

‘Lyc.[aena] Orbitulus Esp. var. (Orbitulinus) Stgr.’ Staudinger, 1892d: 318. Type loca-

laity :y szimiKenter..

Syntypes 3d, 2:

e 5 with labels: handwritten (Staudinger) ‘var. Orbitulinus |

Stgr.’ (on white paper), handwritten ‘Kentei | [18]89 Dörr.[ies]’

(on green paper) and printed ‘Origin. (on pink paper);

e 23, 2 with circles of green locality label paper; one male

specimen has also a handwritten label ‘Kentei’ (on white, faded

yellow, paper).

orientalis Staudinger, 1881

‘Polyomm.{atus| Dorilis Hufn. var. orientalis Stgr.’ Staudinger, 1881: 281. Type locali-

ty: ‘...aus dem Caucasus und Kleinasien... Ala Tau... Brussa...” — Junior subjective

synonym of Papilio tityrus tityrus Poda, 1761 (Hesselbarth et al., 1995: 475).

Lectotype à:

e 6 with labels: handwritten ‘Ala Tau | Hbhr. [Haberhauer]’ (on

brown paper), printed ‘Origin.’ (on pink paper), circle of blue

paper, handwritten (Nekrutenko) on printed form ‘LECTOT Y-

PUS 4 | orientalis Stgr. | Y. Nekrutenko design.’ (on yellow

paper, in box) — Nekrutenko in Hesselbarth et al. (1995: 475).

Paralectotypes 34 , 39:

the lectotype and printed label ‘Origin, handwritten

(Nekrutenko) on printed form ‘PARALECTOTYPUS 2 | ori-

entalis Stgr. | Y. Nekrutenko design. (on yellow paper, in box);

e 6 with labels: handwritten ‘Amasia | Sch.’ (on yellow paper),

handwritten ‘10/5’ and printed ‘Origin.’ (on pink paper);

e 6, ? with labels: handwritten ‘Brussa’ (on white paper), print-

ed ‘Coll. Led. (on white paper) and printed ‘Origin. (on pink

paper);

e G, 2 with labels: handwritten ‘Caucasus’ (on white paper),

printed ‘Coll. Led.’ (on white paper) and printed ‘Origin.’ (on

pink paper).

292

orientalis Staudinger, 1901

‘Lycaena F. Amandus Schn. v. Orientalis Stgr. Staudinger, 1901: 85. Type locality:

‘Pont.[us]; Arm.[enia]; Hyrc.[ania]; Tarb.[agatai]; Ala Tau’; restricted by lectotype desig-

nation: ‘Amasia’ — Junior subjective synonym of Papilio amandus amandus Schneider,

1792:

Lectotype d:

e G (head and abdomen missing), with labels: handwritten

‘Amasia | m.[ihi?]’ (on yellow paper), handwritten ‘orientalis |

Stgr. (on white paper, in pencil), handwritten (Bernardi) on

printed form ‘L. amandus | orientalis Stgr. | lectotype ¢

Bernardi det. | 1964 | cf. Alexanor, III: 273 | 1964 (on white

paper) and combined printed/handwritten designation label

‘LECTOTYPE’ (on red paper) — Bernardi (1964: 273).

Paralectotypes 12(15?)d, 10(97) 9:

paper), small date label ‘10/6’;

‘4/6’;

e 6 with labels: handwritten “Taurus | Hbhr. [Haberhauer]’ (on

yellow paper), handwritten (Bernardi) ‘v. orientalis | Stgr. (on

white paper, in pencil), printed ‘Paratypus’ (on red paper, in

box);

e ¢ with labels: handwritten ‘Hadjin | [18]88 K. O. (on yellow

paper), small date label ‘28/5’, printed ‘Paratypus’ (on red

paper, in box);

2 with handwritten labels: ‘Eibes | [18]89 Man.[issadjian].’ (on

yellow paper), small date label ‘6/5’;

e 3, 2 with handwritten labels: ‘Tekke | [18]94 Wild.[?]. (on

yellow paper);

e d with circle of yellow locality label paper, small date label

‘11/6’ and printed ‘Paratypus’ (on red paper, in box);

3d with circle of yellow locality label paper;

6 with handwritten labels: ‘Achalzich | Hbhr. [Haberhauer]

[18]82’ (on green paper) and small date label ‘3/8’;

293

e d with circle of green locality label paper, small date label

e 26 with labels: handwritten ‘Caucasus’ (on white paper), circle

of green locality label paper (one 4 only), printed ‘Coll. Led.’

(on white paper) and printed ‘Paratypus’ (on red paper, in box);

e 3 with handwritten label: ‘Schakuh | Chr.[istoph]’ (on yellow

paper) and small date label 7/6’ (on brown paper);

e 2% with circle of yellow locality label paper;

yellow paper);

e 6 with handwritten label ‘Tarbagatai | Hbhr. [Haberhauer]’ (on

brown paper);

e 6 with handwritten label ‘Ala Tau | Hbhr. [Haberhauer]’ (on

brown paper);

e ?-5 with handwritten label ‘Margelan | Hbhr. [Haberhauer]’ (on

brown paper);

e ?-G with handwritten labels: ‘Margelan | Maur.[er] [18]84’ (on

brown paper), date label ‘17/5’ ‘Scheichm[?]’ (folded);

e ?-5 with handwritten label ‘Namangan | Hbh. [Haberhauer]

[18]84’ (on brown paper);

e ?-2 with circle of brown locality label paper and handwritten

date label ‘28/5’.

Likewise most of the Staudinger species-group taxa becoming

available after his death in 1900 (i. e. in Staudinger, 1901), the type

specimens were not designated with characteristic ‘Origin. labels.

As may be seen from the original description, the type locality

statement was pretty broad, so that all forementioned specimens are

labelled such as to fit Staudinger’s meaning of the ‘terra typica’ to

constitute a syntype series. Since ‘Grâce à l’amabilite du Dr

Hannemann j’ai sous les yeux certains des exemplaires de

Staudinger qui représentent donc, en langage moderne, les syn-

types d’orientalis’, Bernardi (1964: 273) received on loan only five

294

specimens (by selection of Hannemann), one of these “Je choisis

comme lectotype d un exemplaire étiqueté “Amasia”, the rest were

labelled by him (or by Hannemann on return of the loan?) with

printed labels ‘Paratypus’. In fact, the entire series mentioned

above represents the type series (syntypes and, subsequently, para-

lectotypes).

ornata Staudinger, 1892

‘Lycaena Orion Pall. und var. Ornata Stgr.’ Staudinger, 1892b: 159. Type locality:

‘Maack fand sie am Amur-Ufer, Radde am Onon und im Bur.[eja] G.[ebirge], Christoph

bei Radd.[efka] and Wlad.[iwostok], Dörries auf Ask.[old], am Sutschan und am Bik.[in],

Graeser bei Pokr.[ofka] häufig, bei Wlad.[iwostok] nur an einer stelle... ... alle mir vor-

liegenden Amur-Stücke der ersten Generation...’; replacement name: Scolitantides orion

athenae Hemming, 1934: 98 (the replacement is not justified since it is a senior primary

homonym of Lycaena astrarche ab. ornata Staudinger, 1892 below: for the dates

unknown to Hemming see Reference section).

Syntypes 58,22:

e JS with labels: handwritten (Staudinger) ‘v. Ornata | Stgr. | typ-

isch’ (on white paper), ‘Raddefskaja | Chr.[istoph]’ (on violet

paper), small handwritten date label ‘18/5’ and printed ‘Origin.’

(on pink paper);

e 3, 2 with circles of violet locality label paper and printed

“Origin. (on pink paper);

e d, with labels: handwritten ‘Wladiwostock | Chr.[istoph]’ and

printed ‘Origin. (on pink paper);

e 3, with labels: handwritten ‘Askold | Dörr.[ies]’ (¢ with date

‘[18]83’ and printed ‘Origin.’) (on pink paper);

e d with labels: handwritten ‘Bykin | 18[87] Dorr.[ies]’ and print-

ed ‘Origin.’ (on pink paper).

ornata Staudinger, 1892

‘Lyc.[aena] Astrarche Bersstr. ab. Ornata Stgr.’ Staudinger, 1892c: 280. Type locality:

‘...von Tunis... Ganz ähnliche (gleiche) Stücke fing ich in Anzahl auf der Insel Sardinien,

Corsica’ — Infrasubspecific (cf. Hemming, 1934: 98).

295

Syntypes 98, 9:

e JS with labels: handwritten (Staudinger) ‘v. (ab.) | Ornata Stgr.’

(on white paper), “Sardinia | m.[ihi?]’ (on pale pink paper) and

printed ‘Origin. (on pink paper);

e 56, with circles of pale pink locality label paper and printed

label ‘Origin.’ (on pink paper);

‘Coll. Led.’ (on white paper) and printed ‘Origin.’ (on pink

paper);

e 6 with labels: handwritten ‘Chiclana | m.[ihi?] [18]80’ (on

green paper) and printed ‘Origin. (on pink paper);

e d with labels: handwritten “Tunis | [18]91 Vaulog.’ (on white

paper) and printed ‘Origin. (on pink paper);

e d with labels: handwritten “Teneriffa | Boecker’ (on white

paper) and printed ‘Origin.’ (on pink paper).

osiris O. Bang-Haas, 1927

‘Lycaena osiris O. B.-Haas, spec. nov.’ Bang-Haas, O., 1927: 52, Taf. 7, Figs 26, 27. Type

locality: ‘Armenien; Kagymann (recte: Kagyzman) mont., Dorf Tschangla unteres Ufer

des Araratflusses. Gebirgswiesen 2300-2500 m.’ — Junior primary homonym of Lycaena

osiris Hopffer, 1855; replacement name: Aricia kagizmanensis Kogak, 1980a: 142; junior

subjective synonym of Papilio eumedon eumedon Esper, [1780] (Hesselbarth er al., 1995:

625).

Lectotype à:

e ¢ with labels: handwritten (O. Bang-Haas) ‘Lyc.[aena] osiris |

O B Haas’ (on white paper), printed ‘Kagyzman | 3000m Juli |

Russ. Armen.’ (on white paper), printed, wıth handwritten

insertions ‘Horae Macrolep. | Vol. I abgebildet | t. 7, f. 26 |

beschrieb. p. 52’ (on white paper), printed “Type | O. Bang-

Haas’ (on pink paper), printed ‘Zool. Mus. | Berlin’ (on yellow

paper), handwritten (Nekrutenko) on printed form ‘LECTOTY-

PUS ¢ | osiris O. Bang-Haas | 1927. | Y. Nekrutenko design.’

(on yellow paper) — Nekrutenko in Hesselbarth et al. (1995:

625).

296

3000m Juli | Russ. Armen. (on white paper), printed “Type |

O. Bang-Haas’ (on pink paper), printed “Zool. Mus. | Berlin’ (on

yellow paper), handwritten (Nekrutenko) on printed form

‘PARALECTOTYPUS 2 | osiris O. Bang-Haas | 1927. |

Y. Nekrutenko design. (on yellow paper).

ossmar Gerhard, [1851]

‘Lycaena Corydon Hbr. [Hübner] Var. Ossmar. Bischoff Gerhard, [1851]: 17, Pl. 31,

Fig. 4 a, b, c. Type locality: “Türkei. ... ‘...empfing ich durch Herrn Bischoff, der

dieselbe in der Türkei fing.’

Neotype d:

e 3 with labels: [handwritten ‘Amasia’ (on yellow paper)] — indi-

cated by Schurian (1988: 135) but not found, handwritten

‘corydonius | H. S.’ (on white paper, in pencil), printed ‘Zool.

Mus. | Berlin’ (on pale yellow paper), handwritten (Schurian)

on printed lined form ‘Polyommatus | (Lysandra) | ossmar |

GERHARD 1851 | det. Schurian 1988’ (on white paper), hand-

written (Schurian) on printed form ‘Gen. Präparat |

Nr. 2/4/1985 | Schurian’, handwritten (Schurian) on printed

form ‘Neotypus d | Lysandra ossmar | GERHARD (1851) |

teste SCHURIAN 1984’ (on red paper) and ‘Neotypus d <writ-

ten over printed ‘Paratypus’> | Lycaena corydon | ossmar |

GERHARD 1851 | det. Schurian, 1988’ (on red paper, in box)

— Schurian (1988: 135, Taf. 1, 2, Fig. 5).

palandoekis Schurian & Häuser, 1981

‘Meleageria daphnis palandökis n. subsp. Schurian & Häuser, 1981: 105. Type locali-

ty (holotype): ‘Anatolia/Erzurum Palandöken dagh 2200 m vic. Palandöken, 2 km

südlich’. — Junior subjective synonym of Lycaena meleager versicolor Heyne, [1895]

(Hesselbarth et al., 1995: 687).

Paratypes 24 , 9:

e 23, with printed labels: ‘ANATOLIA/Erzurum | Palandöken

dagh/2200 m | vic. Palandöken/2 km südl. | 5S.VIIT u. 13. -

15.vIll <sic> [19]77 | leg. Schurian’ (on white paper) and

297

‘Paratypus | Meleageria daphnis | palandoekis n. ssp. |

Schurian/Häuser 1981’ (on white paper, in red box).

panaegides Staudinger, 1886

‘Lyc.[aena] Panaegides Stgr. (Panagaea var.?)’ Staudinger, 1886: 206. Type locality:

‘im Juli mit typischen Cytis zusammen in den Gebirgen bei Samarkand gefangen haben.’

Syntypes 66,29:

e 6 with labels: handwritten (Staudinger) ‘Cytis v. Panaegides |

Stgr. | <in different ink> Alaica Stgr. | synonym.’ (on white

paper), “Prov. Samarkand | Hbhr. [Haberhauer] [18]81’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 54,29 with printed label ‘Origin. (on pink paper) and circles

of brown locality label paper; one of the male specimens also

has a small handwritten date label ‘6/7’ (on white paper).

panagaea Herrich-Schäffer, [1851]

‘[Lycaena] Panagaea’ Herrich-Schäffer, [1851] 1(47): Tab. 103, Figs 490-493;

‘Panagaea Kad. ... (früher Endymion Kind.)’ [1852] 6(55): 30. Type locality: ‘Beide

Geschlechter aus Kleinasien, von Herr Kindermann’. — Junior subjective synonym of

Lycaena endymion Gerhard, [1851]. The priority of Gerhard’s name over that of Herrich-

Schäffer’s [1851] is established here after Kocak, 1980c: 17 (cf. ICZN Art. 21.3.1,

22):

Types: not found.

parnassia Staudinger, 1870

‘Lycaena Semiargus Rott. (Acis S. V.) v. Parnassia’ Staudinger, 1870b: 55. Type locali-

ty: ‘Nur auf dem Parnass... gefangen und zwar in grosser Anzahl.’

Syntypes 78,6%:

e 3 with labels: handwritten ‘Parnass | Kr.[iiper]’ (on yellow

paper), printed, with handwritten insertion “Parnass | 20/4

[18]66’ (on white paper), printed ‘Origin.’ (on pink paper);

e 54,59 with circles of yellow locality label paper and printed

label ‘Origin.’ (on pink paper);

298

e G, 2 with labels: printed, with handwritten (Püngeler) inser-

tion ‘Graecia | sept.[entrionalis] mont.[es] (recto) | parnassia &

(2) | vom Stgr. | Püngeler (verso) (on white paper) and hand-

written (Püngeler) on printed form ‘Cotype <underlined> |

v. parnassia Stgr. 3 (2) | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Semiargus Rott. | v. Parnassia Stgr. (on white paper, in

printed box).

persephatta Alpheraky, 1881

‘Lycaena persephatta Alph. nov. sp.’ Alpheraky, 1881: 395, Tab. 14, Fig. 11. Type

locality: ‘L’espece assez rare, volait en Juin le long du Kounguesse entre 4 et

6000 pieds.

Syntypes 6, 3:

e 6 with labels: handwritten ‘Tian Schan | Alph.[eraky]’ (on

brown paper), handwritten (Alphéraky) date label *5/VI [1879]’

(on white paper) and printed ‘Origin. (on pink paper);

e 3% with circles of brown locality label paper, handwritten

(Alpheraky) date labels ‘3/VI [1879] and ‘7/VI [1879]’ (on

white paper) and printed labels ‘Origin.’ (on pink paper); one

female specimen has a handwritten (Alphéraky) label ‘3 VI

1879. | Tianchan. | Persephatta’ (on white paper).

Under a handwritten drawer label ‘Persephatta | Stgr. (on white

paper, in box) there are placed 7 more specimens designated with

Staudinger’s printed labels ‘Origin.’ (on pink paper): 26 , 49 from

‘Margelan | Hbhr. [Haberhauer]’ and a 2 from ‘Ala Tau | Hbhr.

[Haberhauer]’. Along with forementioned specimens, they consti-

tute the type series of a taxon Staudinger intended to describe, but

he abandoned this intention after Alphéraky’s publication (cf.

Staudinger, 1901: 89). The specimens collected by Alphéraky dur-

ing his 1879 trip to ‘district de Kouldja et des montagnes environ-

nantes’ (inclusive Tien-Shien Mts.) and which he sent to Staudinger

undoubtedly are an integral part of his type series, and hence are

provided now with a printed label ‘persephatta | Alphéraky, 1881 |

SYNTYPUS | Y. Nekrutenko det. 11.11.1998’ (on yellow paper).

299

persica Bienert, [1869]

‘P.[olyommatus] Thersamon var. persica m.’ Bienert, [1869]: 28. Type locality: ‘In

subalpiner Region des Gebirges im NW. von Meschet...’

Types: not found (see also Hesselbarth et al., 1995: 485).

persica Bienert, [1869]

‘Lyc.[aena] Icarus var. persica m.’ Bienert, [1869]: 29. Type locality: ‘Mit dem

Stammform bei Nischapur, Ssäbsewar, Chanlung und Meschet...” — Junior secondary

homonym of Lycaena thersamon persica Bienert, [1869]; replacement name:

Polyommatus icarus bienerti Bälint, 1992: 410, 415. Junior subjective synonym of Papilio

icarus Rottemburg, 1775.

Lectotype d:

e d with labels: handwritten ‘Mesch.[et] 170’ (on white paper), a

circle of yellow paper, printed ‘Origin.’ (on pink paper), hand-

written (Nekrutenko) on printed form ‘LECTOTYPUS 4 | per-

sica Bien. | Y. Nekrutenko design. (on yellow paper) —

Nekrutenko in Hesselbarth er al. (1995: 676).

Paralectotype d :

e d with labels: handwritten ‘Persia | Bnt. [Bienert]’ (on yellow

paper), printed ‘Origin.’ (on pink paper), handwritten

(Nekrutenko) on printed form ‘PARALECTOTYPUS 4 | persi-

ca Bien. | Y. Nekrutenko design.’ (on yellow paper) —

Nekrutenko in Hesselbarth et al. (1995: 676).

pfeifferi Brandt, 1938

‘Lycaena Pfeifferi sp. n.’ Brandt, 1938: 671, Taf. 4, Figs 1-3. Type locality: Iran. ‘Die

Typenserie von Barm i Firus (3750 m), Juli doch kommt diese Art auch im Elbursgebirge

vor, so z. B. im Juni auf den Bergen um Nissa’.

Paratypes à , 2:

e 3d, $ with labels: printed ‘IRAN FARS, Straße | Ardekan -

Talochsroe | Comé m 1937 | coll. Brandt’ (on pale yellow

paper), printed ‘62. (on white faded paper), a small square

piece of pale pink paper, printed ‘Paratypus’ (on red paper) and

handwritten ‘Lycaena | pfeifferi B.[randt]’ (on white paper) —

cf. Hauser & Eckweiler (1997: 94).

300

pharaonis Staudinger, 1895

‘Lycaena Pharaonis Stgr. n. sp. Staudinger, 1895a: 243. Type locality: ‘...beschriebe

December bei Cairo, ein anderes am 18. Januar bei Ipsambul (?) in Aegypten gefangen,

3 Stiicke habe ich, als aus Nubien stammend, erhalten.’ — Junior subjective synonym

of Lycaena eleusis Demaison, 1888 and Lycaena podorina Mabille, 1890 (Staudinger,

OUT)

Syntypes 48, 9:

e G with labels: handwritten (Staudinger) ‘Eleusis Demaison |

Podorina Mab. | Pharaonis Stgr.’ (on yellowish paper), hand-

written ‘Cairo’, handwritten ‘Ipsamb. | 18/I [18]80’ (on white

paper) and printed ‘Origin.’ (on pink paper);

e JS with labels: handwritten ‘Cairo | 5/12 [18]79 | Eibek.’ (on

white paper), circle of white locality label paper and printed

‘Origin.’ (on pink paper);

e 26, ¢ with circles of white locality label paper and printed

‘Origin.’ (on pink paper), one male specimen has a handwritten

label ‘Nubien’ (on white paper, in pencil).

phellodendri Staudinger, 1892

Nomen nudum. ‘Thecla Herzi Fxsn. ... Phellodendri... Staudinger, 1892b: 150. Type

locality: ‘Christoph fing diese Art anfang Juli bei Wlad.[iwostok] auf Phellodendron

amurense und hatte ich sie bereits als Phellodendri beschrieben und in einigen Stücken

so versandt, zog aber dann die Beschreibung zurück, da Dr. Fixsen sie nach Stücken von

Korea, die Herz sammelte, diesem zu Ehren benennen wollte.’

Syntypes 2¢, 39:

e 6 with labels: handwritten (Staudinger) ‘Herzi Fixsn |

Phellodendri | Stgr.’ (on white paper), handwritten ‘Baranowsky

| Dörr.[ies]’ (on blue-violet paper) and printed ‘Origin.’ (on pink

paper);

e 6 with labels: handwritten ‘Wladiwostock | Chr.[istoph]’ (on

blue-violet paper), handwritten (in pencil) °6.7.[18]77’ (on

white paper), and printed ‘Origin.’ (on pink paper);

paper), and printed ‘Origin. (on pink paper);

301

small handwritten ‘2/7’, handwritten ‘475 (on white paper)

and printed ‘Origin.’ (on pink paper);

label ‘Origin.’ (on pink paper). —

pheres Staudinger, 1886

‘Lyc.[aena] Pheretides Ev. ... var. pheres Stgr.’ Staudinger, 1886: 209. Type locality:

‘Aus dem Gebirgen bei Namangan und vom siidlichen Alai...’

Syntypes 108, 7%:

e d with labels: handwritten (Staudinger) ‘v. Pheres | Stgr.’ (on

white paper), and ‘Namangan | Hbh. [Haberhauer] [18]84’ (on

brown paper) and printed “Origin. (on pink paper);

e 73,52 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper);

e 3 with labels: handwritten (Staudinger) ‘v. Pheres’ (on white

paper), and ‘Alai | [18]85 Hbh. [Haberhauer]’ (on brown paper)

and printed ‘Origin.’ (on pink paper);

e 6, 22 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper).

pheretimus Staudinger, 1892

‘Lyc.[aena] Pheres Hb. Var. Pheretimus Stgr.’ Staudinger, 1892d: 317. Type locality:

‘...23 Stücke vom Kentei (146 d und 9@ 2)...

Syntypes 2d, 29:

e G with labels: handwritten (Staudinger) ‘Pheretes var.

/Pheretimus | Stgr.” (on white paper), and ‘Kentei | [18189

Dörr.[ies]’ (on green paper), printed ‘Origin. (on pink paper);

e 6, 22 with circles of green locality label paper and printed

label ‘Origin.’ (on pink paper).

pheretulus Staudinger, 1886

‘Lyc.[aena] Pheretiades Ev. var. Pheretulus Stgr. Staudinger, 1886: 209. Type locali-

ty: ‘Aus den Gebirgen bei Osch und später vom Alai...’

302

Syntypes 94,79:

e 6 with labels: handwritten (Staudinger) ‘v. Pheretulus | Stgr.’

(on white paper), and ‘Osch | Hbh. [Haberhauer] [18]82’ (on

brown paper), printed ‘Origin.’ (on pink paper) — Tshikolovets

(19972: 155, 238, pl. 40, figs 31(33));

e 34,29 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper); one of the male specimens also

has a small date label ‘19/7’ (on white paper) — Tshikolovets

(1997a: pl. 40, figs. 34(36)); |

e 6 with labels: handwritten ‘Alai | [18]85 Hbh. [Haberhauer]’

(on brown paper), printed ‘Origin.’ (on pink paper);

e 43,52 with circles of brown locality label paper and printed

label ‘Origin. (on pink paper).

phiala Groum-Grshimaïlo, 1890

‘Lycaena Phiala Gr. Gr.’ Groum-Grshimailo, 1890: 366, pl. 21, fig. 4. Type locality:

‘J'ai trouvé la L. Phiala, à un nombre fort restreint d’exemplaires, dans le Kabadian’.

Paralectotype 3:

e 6 with labels: handwritten (Groum-Grshimailo, in Russian)

‘Kaba- /dian’, ‘12 | 5.[18]85’ (both on white paper), handwrit-

ten (Groum-Grshimailo) ‘Kabadian | Buchara’ (on pink paper),

printed ‘Orig.’ (on green paper), printed ‘Coll. Gr.-Gr (on

white paper), handwritten (Staudinger) ‘Phiala | Gr. Gr. | [18]97

<sic>’ (on white paper).

Lectotype with the same original author’s labels is deposited in

Zoological Institute, Russian Academy of Sciences, St.-Petersburg

(Nekrutenko, 1984a: 31, fig. 2, 2).

phlaeoides Staudinger, 1901

“Chrysophanus Hb. Phlaeas L. v. Phlaeoides (Knaggs in I. ?) Stgr. Staudinger, 1901: 74.

Type locality: ‘Mad.[eira]’.

Syntypes 364, @:

e 6 with handwritten (Staudinger) label ‘Madeira’ (on yellowish

paper);

303

e 2 with circle of yellow locality label paper;

e ¢ with handwritten (Staudinger) label ‘Madera <sic> | 4000’

Bauer’ (on yellowish paper);

e 3 with circle of yellowish locality label paper.

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Phlaeoides Knags? | Stgr’ (on white paper);

each provided now with a printed label ‘phlaeas phlaeoides |

Staudinger, 1901 | SYNTYPUS | Y. Nekrutenko det. 1998’ (on

white paper).

phoenicurus Lederer, 1869

‘Polyomm.[atus] Phoenicurus Led. Lederer, 1869: 8, Taf. 1, Figs 4, 5. Type locality:

‘bei Astrabad in Persien... Ich erhielt nur 13 Exemplaren’ (op. cit.: 3).

Lectotype à:

e 6 with labels: handwritten ‘Hadschyabad’ (on white paper),

printed ‘Origin. (on pink paper), printed ‘Coll. Led. (on

white paper), handwritten (Nekrutenko) on printed form

‘LECTOTYPUS à | phoenicurus Led. | Y. Nekrutenko design.’

(on yellow paper) — Nekrutenko in Hesselbarth et al. (1995:

499).

Paralectotypes 6, 29:

e G, 22 with labels: handwritten ‘Hadschyabad’ (on white

paper), printed ‘Origin.’ (on pink paper), printed “Coll. Led.’ (on

white paper), handwritten (Nekrutenko) on printed form

‘PARALECTOTYPUS 6 (9) | phoenicurus Led. | Y.

Nekrutenko design. (on yellow paper) — Nekrutenko in

Hesselbarth et al. (1995: 499).

phryxis Staudinger, 1886

‘Lyc.[aena] Phryxis Stgr. Staudinger, 1886: 212. Type locality: “Bei Margelan und

Namangan wurde Phryxis Ende Mai gefunden, bei Osch Ende Juni und bei Samarkand

Anfang Juli. Wahrscheinlich sind sie an den letzteren beiden Lokalitäten bedeutend höher

gefanden.’ — Junior primary homonym and junior objective synonym of Lycaena phryx-

is Lang, 1884: 372; junior subjective synonym of Lycaena sarta Alpheraky, 1881

(Staudinger, 1901: 85).

304

Syntypes 74,79:

e d with labels: handwritten (Staudinger) ‘Phryxis | Stgr. (on

white paper), and ‘Margelan | Hbhr. [Haberhauer]’ (on brown

paper), printed ‘Origin.’ (on pink paper);

e 34,39 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper), one male specimen also has a

handwritten date label ‘25/5 [18]80’;

e 6 with labels: handwritten (Staudinger) ‘Sarta Alph.[éraky] |

Phryxis Stgr. (on white paper), handwritten ‘Osch | Hbh.

[Haberhauer]’ (on brown paper), handwritten date label ‘26/6’

(on white paper) and printed ‘Origin. (on pink paper);

label ‘26/6’ (on white paper) and printed label ‘Origin.’ (on pink

paper);

[18]84’ (on brown paper), handwritten date label ‘23/5’ (on

white paper) and printed ‘Origin.’ (on pink paper);

e 6 with labels: handwritten ‘Prov. Samark.[and] | Hbh.

[Haberhauer] [18]81’ (on brown paper), handwritten date label

‘6/6’ (on white paper) and printed ‘Origin.’ (on pink paper);

e 3,2% with circles of brown locality label paper, handwritten

date label *26/6’ (on white paper) and printed label ‘Origin.’

(on pink paper); the male specimen has a handwritten date

label ‘6/9’ (on white paper).

phyllides Staudinger, 1886

‘Lyc.[aena] Phyllis Chr. var. Phyllides Stgr. Staudinger, 1886: 225. Type locality:

“...erhielt ich in geringer Anzahl aus dem Gebirgen von Namangan, Osch, Margelan und

Samarkand’.

Lectotype d :

e 6 with labels: handwritten ‘Namangan | Hbhr. [Haberhauer]

[18184 (on brown paper), printed ‘Origin.’ (on pink paper),

handwritten (unknown hand) ‘24/5 Lyc. sp. n. | Rogenhoferii’

(on white paper), handwritten (Forster) ‘Lectotypus d |

Lycaena phyllis | var. phyllides Stgr. | W. Forster 1947’ (on pink

305

paper) — Forster (1960: 138); cf. Häuser & Eckweïler (1997:

95).

Paralectotypes 10d, 39:

3 with labels: handwritten (Staudinger) ‘v. Phyllides | Stgr. (on

white paper), handwritten ‘Osch | Hbh. [Haberhauer]’ (on

brown paper), small date label ‘6/7’ (on white paper), printed

‘Origin.’ (on pink paper);

3 with circle of brown locality label paper and printed label

‘Origin.’ (on pink paper);

3, 22 with circles of brown locality label paper of the lecto-

type (‘Namangan | Hbhr. [Haberhauer] [18]84’) and printed

label ‘Origin.’ (on pink paper);

6 with labels: handwritten ‘Alai | [18]85 Hbh. [Haberhauer]’

(on brown paper) and printed ‘Origin.’ (on pink paper);

3S with circle of brown locality label paper and printed label

‘Origin.’ (on pink paper);

6 with labels: handwritten ‘Prov. Samark.[and] | Hbhr.

[Haberhauer] [18]81’ (on brown paper) and printed ‘Origin.

(on pink paper);

48, 2 with circles of brown locality label paper, small date

labels ‘14/5’ ‘16/5’, ‘23/5’, ‘24/5’, ‘15/5° (2) and printed label

‘Origin. (on pink paper) — cf. Häuser & Eckweiler (1997: 95).

phyllis Christoph, 1877

‘Lycaena Damon var. Phyllis Stgr. in litt Christoph, 1877: 237, Tab. 5, Figs 9, 10. Type

locality: ‘...bei Schahkuh den Juli hindurch im Thale und auf den Hügeln bei etwa

7000’ hinauf recht häufig.’

Lectotype à:

306

d with labels: circle of yellow paper, Christoph’s date label

‘11/7’ (11 on white, ‘7’ on pink background, recto white/pink,

verso pink), printed ‘327. (on white paper faded yellow), print-

ed ‘Zool. Mus. | Berlin’ (on pale yellow paper), handwritten

(Forster) ‘Cotypus 4 | Lycaena phyllis | Christoph’ (on pink

paper), new handwritten (Hauser) “Hadschyabad’ (on white

paper), printed, with handwritten (Wagener, in pencil) insertion

‘Abgebildet in Hesselbarth, | van Oorschot & Wagener: |

Tagfalter der Türkei | Tafel 727 Figur 33° (on white paper),

printed ‘Lectotypus | Polyommatus (Agrodiaetus) | phyllis

(Christoph, 1877) | des. ECKWEILER & TEN HAGEN 1998’ (on red

paper, in box) — Eckweiler & ten Hagen (1998: 113, Abb.

7 N#errHesselbarth'erial. (1995: 752, Taf. 12T, Fig. 33, Taf.

128, Figs 66, 72); Häuser & Eckweiler (1997: 95); Eckweiler &

Häuser (1997: Pl. 9).

Paralectotypes 54 , 39:

e ¢ with labels: handwritten ‘Schakuh | Chr.[istoph]’ (on yellow

paper), printed ‘326.’ (on white paper faded yellow) and hand-

written (Forster) ‘Cotypus d | Lycaena phyllis | Christoph’ (on

pink paper) — Forster (1960: 135);

e 34,39 with circles of yellow locality label paper;

e d with labels: handwritten ‘Hadschyabad’ (on white paper),

printed, with handwritten insertion ‘Damon | Coll. Led. (on

white paper) — cf. Eckweiler & ten Hagen (1998: 113).

e 23, 22 designated with printed labels ‘Paralectotypus |

Polyommatus (Agrodiaetus) | phyllis (Christoph, 1877) | des.

ECKWEILER & TEN HAGEN 1998’ (on bluish paper, in box).

poseidon Herrich-Schäffer, [1851]

‘[Lycaena] Poseidon Kad. Herrich-Schäffer, [1851] 6(48): 24. Type locality: ‘Aus

Kleinasien’.

Types: not found (cf. Häuser & Eckweiler, 1997: 95).

poseidon Lederer, 1852

‘Lycaena * Poseidon Kdm. [Kindermann] in lit.’ Lederer, 1852: 20 (nomen nudum; *

designates ‘was nicht echt europdisch’ — op. cit.: 15); ‘Lycaena Poseidon Kinderm.|ann]

(nicht Kaden) in lit” Lederer, 1852: 37. Type locality: [‘asiatische Tiirkei’?]. — Junior

primary homonym of Lycaena poseidon Herrich-Schäffer, [1851].

Types: not found (cf. Häuser & Eckweiler, 1997: 96).

307

poseidonides Staudinger, 1886

‘Lyc.

[aena] Poseidon var. Poseidonides Stgr. Staudinger, 1886: 225. Type locality:

‘Vier d vom südlichen Alai sowie eins von den südlich von Samarkand gelegenden

Gebi

gen...

Syntypes 5d:

3 with labels: handwritten (Staudinger) ‘v. Poseidonides | Stgr.’

(on white paper), handwritten ‘Prov. Samark.[and] | Hbhr.

[Haberhauer] [18]81’ (on brown paper), small date label ‘12/7’

(on white paper) printed ‘Origin.’ (on pink paper) and printed

‘Zool. Mus. | Berlin’ (on pale yellow paper);

6 with labels: handwritten ‘Alai | [18]89 Hbh. [Haberhauer]’

(on brown paper), printed ‘Origin.’ (on pink paper) and printed

‘Zool. Mus. | Berlin’ (on pale yellow paper);

d with circle of locality label brown paper, handwritten

(Staudinger) label ‘v. Poseidonides | Stgr. | Alai’ (on white

paper) and printed ‘Origin.’ (on pink paper);

28 with circles of brown locality label paper, printed label

‘Origin.’ (on pink paper), one of them with printed label “Zool.

Mus. | Berlin’ (on pale yellow paper) and two new handwritten

labels ‘poseidonides’ (in ball pencil) and ‘Alai’ (in pencil), both

on white paper (cf. Häuser & Eckweiler, 1997: 96).

posthumus Christoph, 1877

‘Lycaena Damon var. Posthumus Chr. Christoph, 1877: 238, Tab. 5, Fig. 11. Type

locality: ‘Ich fand den Schmetterling nur von 9—11,000’ auf dem Schahkuh-Gebirge, wo

er vereinzelt im Juli flog’.

Lectotype d:

308

d with labels: handwritten ‘Schahkuh | Chr.[istoph]’ small date

label ‘26/V’ (‘26° on white, ‘V’ on pink background, recto

white/pink, verso pink), printed ‘329.’ (on white paper faded

yellow), handwritten (Forster) ‘Cotypus 4 | Lycaena damon |

var. posthumus | Christ.’ (on pink paper), printed ‘Lectotypus |

Polyommatus (Agrodiaetus) | posthumus (Christoph, 1877) |

des. ECKWEILER & TEN HAGEN 1998’ (on red paper, in box) —

Forster (1960: 141); Häuser & Eckweiler (1997: 96); Eckweiler

& Häuser (1997: Pl. 9); Eckweiler & ten Hagen (1998: 113,

Abb. 10-12).

Paralectotypes 34 , 39:

e 6 with labels: handwritten (Christoph?) ‘vom 9000’ <illegi-

ble>‘, printed ‘330. (on white paper faded yellow), printed

‘Zool. Mus. | Berlin’ (on pale yellow paper), handwritten

(Forster) ‘Cotypus d | Lycaena damon | var. posthumus |

Christ.’ (on pink paper) — Forster (1960: 141);

e 26 with circles of yellow locality label paper and small date

labels ‘8/7’ and ‘26/7’;

date label ‘2/7’, printed label ‘331. (on white faded yellow

paper), printed ‘Zool. Mus. | Berlin’ (on pale yellow paper),

handwritten (Forster) ‘Cotypus 2 | Lycaena damon | var.

posthumus | Christ.’ (on pink paper) and new ‘Astrabad | coll.

Stgr.’ (on white paper) — Forster (1960: 141);

date label ‘18/7’, printed label ‘328. (on white faded yellow

paper), handwritten (Forster) ‘Cotypus 2 | Lycaena damon |

var. posthumus | Christ.’ (on pink paper) — Forster (1960: 141);

cf. Eckweiler & ten Hagen (1998: 113).

3d and 39 designated with labels ‘Paralectotypus |

Polyommatus (Agrodiaetus) | posthumus (Christoph, 1877) | des.

ECKWEILER & TEN HAGEN 1998’ (on bluish paper, in box).

praemerisina Lorkovié, 1943

‘Everes argiades merisina n. Frühjahrsform praemerisina n. Lorkovié, 1943: 451,

Taf. 25, Fig. 12, Taf. 26, Fig. 11. Type locality: ‘China, Kwang-tung, Gao-fung.’ —

Infrasubspecific.

Holotype à :

e 6 with labels: printed, with handwritten insertion ‘China

Kwantung | Gao fung | 8. 5.[19]77 IR. Mell’ (on white paper),

handwritten (Forster <sic>) on printed form ‘Holo-Type

309

<corrected to Typus> d | Everes argiades | praemerisina

Lorkovic’ (on pink paper), handwritten original field label

‘GF 8. V. 1713 u.<?> 4 (recto) | <text in Chinese> (verso)’ (on

white paper, in pencil, folded);

Paratypes 69, 39:

e 54,39 with labels: printed, with handwritten insertion ‘China

Kwantung | Gao fung | <different dates late April and early

May> .[19]77 IR. Mell’ (on white paper), handwritten (Forster

<sic>) on printed form ‘Para-Typus 4 (2) | Everes argiades |

praemerisina Lorkovic’ (on pink paper);

e 6 with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao shan | 18. 4.[19]77 IR. Mell’ (on white

paper), handwritten (Forster <sic>) on printed form ‘Para-

Typus do | Everes argiades | praemerisina Lorkovic’ (on pink

paper).

Some specimens have also original field labels (folded).

pretiosa Staudinger, 1886

‘Lyc.[aena] Pretiosa Stgr. Staudinger, 1886: 209. Type locality: ‘Diese Art erhielt ich

in Anzahl der Umgegend von Margelan und Namangan.’ — Junior primary homonym and

junior objective synonym of Lycaena pretiosa Lang, 1884: 370.

STADE PO 70

e 3 with labels: handwritten (Staudinger) ‘Pretiosa | Stgr (on

white paper), handwritten ‘Margelan | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 53,32 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper);

e 3 with labels handwritten ‘Namangan | Hbhr. [Haberhauer]

[18]84’ (on brown paper) and printed ‘Origin.’ (on pink

paper);

e 39 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

310

privata Staudinger, 1896

‘Lyc.[aena] Chiron Hufn. v. Privata Stgr. Staudinger, 1896: 300. Type locality:

‘...wurden mir von Herrn R Tancré eingesandt, dessen Sammler E. Rückbeil sie in den

Jahren 1892 uns 1893 im Innern Tibets, zwischen dem Lob Noor und Kuku Noor, fand.

Syntypes 46, 2°:

e 3 with labels: handwritten (Staudinger) ‘Chiron var. | Privata |

Ster. (on white paper), handwritten ‘Kuku Noor | [18]94 <sic!

— lapsus calami? — see above under Type locality> Rckbl.

[Riickbeil]’ (on brown paper) and printed ‘Origin.’ (on pink

paper);

e 2% with circles of brown locality label paper, one specimen

also has a small square piece of pink paper.

prunoides Staudinger, 1887

‘Thecla Prunoides Stgr. Staudinger, 1887b: 129, pl. 6, fig. 1, a, b. Type locality:

“...besitze ich 1 & und 3 @ 9, die Christoph Mitte Juli bei Wladiwostok fing, so wie ein

(altes) 2, das Kindermann 1852 bei Ust-Kamenogorsk im Altai-Gebiet fand und das in

Lederer’s Sammlung steckte.’

Syntypes d,4®:

e ¢ with labels: handwritten (Staudinger) ‘Prunoides | Stgr. in

litt” (on white paper), handwritten ‘Wladiwostock |

Chr.[istoph]’ (on blue paper), small handwritten ‘16/7’ (recto

white, verso red), printed ‘Origin.’ (on pink paper);

e 32 with circles of blue locality label paper, small handwritten

labels ‘16 | 7’, ‘10 | 7’, ‘16° and printed ‘Origin. (on pink

paper);

‘Ustkam.[enogorsk] | 1852’ (on white paper), printed, with

handwritten insertion ‘W-album’ <sic> | Coll. Led.’ (on white

paper), printed ‘Origin.’ (on pink paper).

pseudocyanea Forster, 1956

‘Agrodiaetus xerxes pseudocyanea ssp. nov.’ Forster, 1956: 120, Taf. 12, Figs 6, 7, Taf. 13,

Figs 6, 7. Type locality: ‘Achalzich’. — Junior subjective synonym of Lycaena

damone cyanea Staudinger, 1899 (Carbonell, 2000: 213, 215).

311

Holotype: d :

e 6 with labels: handwritten ‘Achalzich | Hbhr. [Haberhauer]

[18]82’ (on green paper), small handwritten ‘24/5’ (‘24° on

white background, ‘5’ on pink background; recto white/pink,

verso pink), printed ‘319.’ (on white paper faded yellow), print-

ed ‘Origin. (on pink paper), printed ‘Zool. Mus. | Berlin’ (on

pale yellow paper), handwritten (Forster) on printed form

‘Cotypus | Lycaena damone | cyanea Stgr. (on pink paper),

handwritten (Forster) on printed form ‘Holo-Typus <overwrit-

ten to correct “Type’> à | Polyommatus xerxes | pseudocyanea

Forster | W. Forster Miinchen’ (on pink paper), printed, with

handwritten (Wagener, in pencil) insertions ‘Abgebildet in

Hesselbarth, | van Oorschot & Wagener: | Tagfalter der Türkei |

Tafel /23 Figur 37’ (on white paper) — Forster (1956: Taf. 12,

Fig. 6, Taf. 13, Fig. °6); Hesselbarth ef ara Ze

Fig. 37); Hauser & Eckweiler (1997: 97); see also cyanea

Lederer paralectotype above.

Allotype 9:

printed ‘320. (on white paper faded yellow), printed ‘Origin.

(on pink paper), printed ‘Zool. Mus. | Berlin (on pale

yellow paper), handwritten (Forster) on printed form ‘Co-Typus

2 | Lycaena damone | cyanea Stgr (on pink paper), hand-

written (Forster) on printed form ‘Allo-Typus <overwritten to

correct ‘Type’> 2 | Polyommatus xerxes | pseudocyanea

Forster | W. Forster Miinchen’ (on pink paper), printed, with

handwritten (Wagener, in pencil) insertions ‘Abgebildet in

Hesselbarth, | van Oorschot & Wagener: | Tagfalter der Türkei |

Tafel 124 Figur 50° (on white paper) — Forster (1956: Taf. 12,

Fig. .7, Taf. 13, Fig. 7); Hesselbarth et. al... (1: TEE,

Fig. 50).

pseudoxerxes Forster, 1956

‘Agrodiaetus carmon pseudoxerxes ssp. nov.’ Forster, 1956: 118, Taf. 9, Figs 3-5,

Taf. 13, Figs 3-5. Type locality: ‘Persia s.[eptentrionale] Elburs mts., Schahkuh’ (holo-

type).

312

Paratypes 89, 129:

e ¢ with labels: handwritten ‘Schakuh | Chr.[istoph]’ (on yellow

paper), small date label ‘9/7’, a small square piece of paper;

printed ‘Zool. Mus. | Berlin’ (on pale yellow paper), handwrit-

ten (Forster) on printed form ‘Para-Typus 6 | Agrodiaetus car-

mon | pseudoxerxes Forster | W. Forster München’ (on pink

paper, in box);

2 with labels: circle of yellow paper, printed ‘290. (on white to

yellow paper), handwritten (Forster) on printed form ‘Para-

Typus 2 | Agrodiaetus carmon | pseudoxerxes Forster |

W. Forster München’ (on pink paper, in box) and handwritten

(Wagener) ‘Schahkuh | Chr.[istoph]’ (on white paper);

3d with labels: circle of yellow paper, small date label ‘15/7’,

printed ‘298. (on white to yellow paper) and handwritten

(Forster) on printed form ‘Para-Typus d | Agrodiaetus carmon

| pseudoxerxes Forster | W. Forster München’ (on pink paper, in

box):

38, 2 with same circles of yellow paper and small date labels

that apparently belong to the type series;

d with labels: printed, with handwritten insertions 'HYRCA-

NIA, (Schahkuh) | Funke 1898’ (on white paper, in box), hand-

written (Püngeler) ‘carmon’ (on white cross-lined notebook

paper) and handwritten (Forster) on printed form ‘Para-Typus

3 (2) | Agrodiaetus carmon | pseudoxerxes Forster | W. Forster

München’ (on pink paper);

28,9% with labels: printed, with handwritten insertions ‘HYR-

CANIA, (Schahkuh) | <some with different dates>’ (on white

paper, in box), handwritten (Püngeler) and handwritten

(Forster) on printed form ‘Para-Typus 4 (9%) | Agrodiaetus

carmon | pseudoxerxes Forster | W. Forster München’ (on pink

paper);

2 with labels: handwritten ‘Hyrcania’ (on white paper, in box)

and handwritten (Forster) on printed form ‘Para-Typus ¢ (8)

Agrodiaetus carmon | pseudoxerxes Forster | W. Forster

München’ (on pink paper) — see also Häuser & Eckweiler

(19971291:

313

psittacus Herrich-Schäffer, [1844]

‘Lycaena Psittacus’ Herrich-Schäffer, [1844] 1(7): Tab. 48, Figs 220-223; [1845] 1(10):

129. Type locality: ‘Aus der Türkei, auch von der asiatischen Seite.’ — Junior subjec-

tive synonym of Lycaena balkanica Freyer, [1843] (Staudinger, 1871a: 9; 1901: 76).

Types: not found.

psylorita Freyer, [1845]

‘Lycaena Pap.{ilio| Psylorita’ Freyer, [1845] 5(79): 146, Tab. 469, Figs 3, 4. Type

locality: ‘Von Dr. Frivaldzky <sic> habe ich zwei Exemplare von diesem Tagfalter unter

obigen Namen erhalten... Nach der mir von Herrn Dr. Frivaldzky <sic> gegebenen

Nachricht fliegt dieser Falter im Juni auf dem Berge Ida auf der Insel Creta’.

Syntype d:

e 3 with labels: handwritten ‘Psylorita | Friv. H Sch. f. 328-31 |

Frey. n. B. t. 469. | Creta Friv.’ (on white paper, in printed box),

printed ‘4713’ (on white paper).

Catalogus Generalis: 4713 — see Appendix.

The specimen is provided now with a printed label ‘psylorita

Freyer, [1845] | SYNTYPUS | Y. Nekrutenko det. 30.11.1998’ (on

yellow paper).

pusillargus Strand, 1919

‘Lycaena semiargus Rott. f. pusillargus m. Strand, 1919: 18. Type locality: ‘Marburg’.

— Infrasubspecific.

E. Strand, [190]8. | 7.07’ (on white paper), handwritten (Strand)

on printed form Lycaena | semiargus | Rott. f. pu | silargus

<sic> m. & | Strand det.’ (on white paper) and printed ‘Type’

(on dirty-pink paper).

quercivora Staudinger, 1887

‘Thecla Quercivora Stgr. Staudinger, 1887b: 137, pl. 6, fig. 2, a, b. Type locality:

[Amur-Gebiet]: ‘...erhielt ich nur drei erzogene Stücke durch Dörries, die mir alle

Weibchen zu sein scheinen’.

314

Syntypes 39:

e 2 with labels: handwritten (Staudinger) ‘Signata | Butl. |

Quercivora | Stgr. (on white paper), handwritten ‘Askold |

[18]83 Dorr.[ies]’ (on violet-blue paper), small handwritten

‘11’ and printed ‘Origin. (on pink paper);

let-blue paper) and printed ‘Origin. (on pink paper);

(on pink paper).

quercus Kardakoff, 1928

‘Zephyrus taxila Brem. f.. 2 quercus (nom. coll.)’ Kardakoff, 1928: 271, Taf. 7, Fig. 16.

Type locality: ‘...aus dem Ussurigebiet...’ — Infrasubspecific.

Syntype ©:

‘Sidemi Ussurigebiet | 5.8. 1927. N. Kardakoff’ (on white

paper), handwritten (Kardakoff?) ‘Zephyrus taxila Brem L. |

f. 2 quercus Kardakoff’ (on white paper) and printed “Typus’

(on red paper, in box).

quercus’ <underlined with red pencil> (on a stripe of white paper,

in blue type ribbon).

rebeli Hirschke, 1905

‘Lycaena Alcon F. v. Rebeli’ Hirschke, 1905: 110, Taf. 2, Figs 1,2. Type locality: ‘Den

ganzen Sommer 1904 verbrachte ich in Steiermark und erbeutete dort im zirka 1700 m

Höhe eine auffallende, in beiden Geschlechtern äußerst constante Lycaena-Form... Von

den in Anzahl erbeuteten Exemplaren, welche untereinender keine Abweichungen zeigen,

Syntypes 24, 9:

e 26, 2 with labels: printed, with handwritten (Hirschke?) inser-

tion ‘Hirschke. Wien | Styria | VII. 1904 | Co=Typ (recto) |

alcon var. | Rebeli 8 (2) | Cotype <double underlined> |

1A19]06 v.[om] A. Hirschke (verso, written by Püngeler)’ (on

white paper, recto in printed box), handwritten (Püngeler) on

315

printed form ‘Cotype <underlined> | v. Rebeli Hirschke Salz)

| vom Autor | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Alcon F. | v. Rebeli Hirschke’ (on white paper, in printed

box).

rezniceki Bartel, 1904

‘Lycaena coridon var. rezniceki nov. var.’ Bartel, 1904: 117. Type locality: “Eine weit-

ere, noch unbeschriebene Lokalform von Lyc. coridon kommt an der Riviera vor.’

‘Fluggebiet: Rapallo (Riviera di Levante 1.-10. Juni 1904), Monte Carlo (10.-13. Juni

1904, E. v. Reznicek). — Auch von Bordhigera und Nizza bekannt.’

Syntype d:

e 3 with labels: printed ‘Gall.[ia] mer.[idionalis] or.[ientalis] |

Riviera V à | coll. M. Bartel’ (on white paper, in box), hand-

written (Püngeler) on printed form ‘Cotype <underlined> |

v. Rezniceki Bartel |. 3 | vom Autor | 2.1905 | Püngeler’ (on pink

paper).

rileyi Forster, 1940

‘Zephyrus rileyi spec. nov.’ Forster, 1940: 871, Taf. 22, Figs 1, 2, Taf. 23, Figs 1, 2. Type

locality (holotype): ‘China, Kwangtung, Lin ping.’

Holotype 3:

e 3 with labels: printed, with handwritten insertion ‘China

Kwantung | Lin ping | 25.5.[19]21 | R. Mell’ (on white paper),

handwritten (Mell) ‘Lp [Ling ping] 25.V.[19]21’ (on white

paper, in pencil, folded), printed ‘Zool. Mus. | Berlin’ (on pale

yellow paper), handwritten (Forster) on printed form ‘Holo-

Type 4 | Thecla (Zephyrus) | rileyi Forst. |W. Forster München’

(on pink paper, in box), printed, with handwritten (Howarth)

insertion ‘T.G.H. | 1955 | 152’ (on white paper), two printed,

with handwritten (Howarth) insertion ‘Neg. No. 17767 and

‘Neg. No. 17768’ (both on white paper).

Allotype ©:

e 2 (abdomen missing), with labels: printed, with handwritten

insertion ‘China Kwantung | Lung tao shan | 30.6.[19]77 |

316

R. Mell’ (on white paper), handwritten (Mell) field label ‘Dr

30.VI.[19]17 (on white paper, in pencil, verso with text in

Chinese in Indian ink, folded), printed ‘Zool. Mus. | Berlin’ (on

pale yellow paper), handwritten (Forster) on printed form ‘Allo-

(on pink paper, in box), printed, with handwritten (Howarth)

insertion ‘Neg. No. 17764’ (on white paper).

Paratypes 2 9 :

2 with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao shan | 50.6.[19]77 | R. Mell’ (on white

paper), handwritten (Mell) field label ‘Dr 30.VI.17’ (on white

paper, in pencil, verso with text in Chinese in Indian ink and

‘Zeph. paraiberica’ <?>, folded), printed “Zool. Mus. | Berlin’

(on pale yellow paper), handwritten (Forster) on printed form

‘Para-Type 2 | Thecla (Zephyrus) | rileyi Forst. | Dr. W. Forster

München’ (on pink paper, in box);

? with labels: printed, with handwritten insertion ‘China

Kwantung | Lung tao shan | 12.6.[19]77 | R. Mell’ (on white

paper), handwritten (Mell) field label ‘Dr 12.VI.17’ (on white

paper cut from a newsletter, in pencil, verso with text in

Chinese in Indian ink and <illegible> <?>, folded), printed

‘Zool. Mus. | Berlin’ (on pale yellow paper), handwritten

(Forster) on printed form ‘Para-Type 2 | Thecla (Zephyrus) |

rileyi Forst. | Dr. W. Forster München’ (on pink paper, in

box).

The holotype and allotype are figured by Forster (op. cit.) on

Tafel 22 and 23, Abb. 1, 2; when compared to the present state of

the specimens, it becomes clear that they were exposed to high

humidity and an apparently inaccurate treatment, resulting in

changes in the position of the wings and antennae and the loss of

the abdomen.

rippertü Boisduval, 1832

‘Argus Rippertii’ Boisduval, 1832: 68, pl. 16, figs 4-6. Type locality: ‘...aux environs

de Digne, ou il a été découvert en 1829 par M. Rippert de Beaugency...’ — Junior sub-

jective synonym of Lycaena ripartii Freyer, 1830.

2117)

Syntypes 2d, 2:

e 6 with labels: handwritten ‘Rippertii | Boisd.[uval] Ic. Hb. f.

958 | Frey. H Sch. | Alp.[ae] Gallopr.[ovinciales] Boisd’ (on

white paper, in printed box), printed ‘4715’ (on white paper);

e 3, 2 without any label.

Catalogus Generalis: 4715 — see Appendix; cf. Häuser &

Eckweiler (1997: 99).

The specimens are provided now with a printed label ‘rippertii

Boisduval, [1832] | SYNTYPUS | Y. Nekrutenko det. 30.11.1998’

(on yellow paper). For a lectotype designation see Balint (1999:

54-55): the present specimens thus become paralectotypes.

romanovi Christoph, 1882

‘Thestor Romanovi n. sp. Christoph, 1882: 106. Type locality: ‘Hier folgt die

Beschreibung einiger als neu erkannter, fast nur von Ordubat stammenden Arten’ (op. cit.:

106).

Paralectotypes 4d , 29:

e 6 with handwritten label ‘Ordubat | Cauc.[asus] m.[eridionalis]

or.[1entalis] | Chr.[istoph]’ (on green paper);

e Ö with handwritten label ‘Ordubat | | Chr.[istoph]’ (on green

paper) and small elongated handwritten (Christoph) label

‘Ordub.[at] 32.5.[18]81’ (on white paper, in much faded ink);

e 24,29 with circles of green locality label paper; of these, two

male specimens with small elongated handwritten (Christoph)

label ‘Ordub.[at] 32.5.[18]81° (on white paper, in much faded

ink), two female specimens with small handwritten (Christoph)

date labels ‘15/5’ and ‘24/5’ (recto yellowish white, verso deep

blue, almost black).

Lectotype designated by Nekrutenko in Hesselbarth et al., 1995:

506 (deposited in Zoological Institute, Russian Academy of

Sciences, St.-Petersburg, Russia). Additional evidence in support to

the label date of forementioned specimens as belonging, as an inte-

gral part, to Christoph’s type series who shared it with Staudinger,

a quotation from Romanoff (1884: 48) is given: “Romanovi Chr.

318

(Pl. I, fig. 1 et 2). — C’est au printemps de 1881 que M. Christoph

a découvert ce nouveau Thestor dans les environs d’Ordoubad. The

specimens are provided now with labels ‘romanovi | Christoph,

1882 | PARALECTOTYPUS | Y. Nekrutenko det. 13.11.1998’ (on

yellow paper) (ICZN Art. 74.1.3, Recommendation 74F).

rosacea Austaut, 1885

‘Lycaena Theophrastus Fabr. v. Rosacea’ Austaut, 1885: 141. Type locality: ‘...est

fondée sur une douzaine d’exemplaires d que j’ai reçus des hauteurs situées entre El-

Arricha et le Maroc.

Syntypes 4d:

e 3 with labels: handwritten (Staudinger) ‘v. Rosacea |

Aust.[aut]’ (on white paper), handwritten (Staudinger)

‘Marocco | Aust.[aut]’ (on white paper) and printed ‘Origin.’ (on

pink paper); |

e 3d with circles of white locality label paper and printed

‘Origin.’ (on pink paper).

rueckbeili Forster, 1960

‘Agrodiaetus iphigenia rückbeili ssp. (f. alt?) nov.’ Forster, 1960: 126, Taf. 10, Figs 21,

22, Taf. 11, Figs 21, 22. Type locality (holotype): ‘Ost-Turkestan, Aksu 1900 leg.

Riickbeil’.

Paratypes 36:

e JS with labels: printed, with handwritten (Püngeler) insertions

“Ost-Turkestan | (Aksu) | Rückbeil 1900° and handwritten

(Forster) on printed form ‘Para-Typus & | Polyommatus

(Agrodiaetus) | iphigenia rückbeili Forst. | W. Forster München’

(on pink paper, ın box);

e 26 with labels: printed, with handwritten (Püngeler) insertions

“Ost-Turkestan | (Aksu) | Rückbeil 1900° and handwritten

(Forster) on printed form ‘Para-Typus 6 | Polyommatus

(Agrodiaetus) | iphigenia riickbeili Forst. | W. Forster München’

(on pink paper, in box) — cf. Forster op. cit.: 126 under

Variabilität entry; Hauser & Eckweiler (1997: 100).

319

rufopunctatus Neuburger, 1907

‘Lycaena icarus Rott. ab. rufopunctatus’ Neuburger, 1907: 180. Type locality:

‘Beschrieben nach einem ¢ aus Beirut in Syrien...” — Infrasubspecific.

Holotype 4 (by monotypy):

e 3 with labels: printed ‘Syria | Beirut | Neuburger’ (on white

paper, in box), large handwritten (Neuburger?) ‘Lycaena icarus

Rott. | ab. rubropunctatus | Neubgr.’ (on white faded yellowish

lined notebook paper), printed ‘Type’ (on pink-brownish

paper).

rutilans Staudinger, 1886

‘Lyc.[aena] Rutilans Stgr. Staudinger, 1886: 208. Type locality: ‘Diese interessante

neue Art erhielt ich erst kürzlich in 12 meist nicht ganz reinen Stiicken vom südlichen

Alai, wo sie im Juni (oder Juli) von Haberhauer und Maurer gefunden wurden.’

Syntypes 66 , 29:

e d with labels: handwritten (Staudinger) ‘Rutilans | Stgr (on

white paper), handwritten ‘Alai | [18]85 Hbh. [Haberhauer]’ (on

brown paper) and printed ‘Origin.’ (on pink paper) — figured in

Tshikolovets (1997b: 36, figs 5, 6);

e 5d, 22 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

saphirina Staudinger, 1887

‘Thecla Saphirina Stgr. Staudinger, 1887b: 135, pl. 16, figs 3-5. Type locality: ‘...aus

dem Amur-Gebiet erhielt ich nur ein d und 2 2 2 von der Insel Askold durch Dörries...’

Syntypes 6, 2°:

e 3 (abdomen missing), with labels: handwritten (Staudinger)

‘Saphirina | Stgr. (on white paper), handwritten ‘Askold |

Dör.[ries]’ (on bluish paper), printed ‘Origin.’ (on pink paper),

two handwritten (Dantchenko) in red ink ‘d saphirinus |

8.03.96 | Dantchenko | abdomen’ and ‘1’ (both on white paper);

e 2° with circles of bluish locality label paper and printed label

‘Origin.’ (on pink paper).

320

sarthus Staudinger, 1886

‘Pol.[yommatus] Sarthus Stgr. n. sp.’ Staudinger, 1886: 202. Type locality: ‘Von dieser

neuen Art erhielt ich im vorigen Jahre nur 4 Stücke (3 4 1 2) von den Herren Haberhauer

und Maurer, die sie im südlichen Alai Gebirge, wohl beim Kara Kasuk Paß gefangen

haben.’

Syntypes 38, 9:

e ¢ with labels: handwritten (Staudinger) ‘Sarthus | Stgr. (on

white paper), handwritten ‘Alai | [18]85 Hbh. [Haberhauer]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 26, 2 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

sassanides Kollar, 1849

‘Thecla Sassanides Kllr.’ Kollar, 1849: 10. Type locality: not specified; ‘Siidpersien’.

Syntypes 2 d :

e <6 with labels: handwritten ‘Sassanides | Koll.[ar] Verh. Wien.

Ac. | TI p. 10 | Persia Kollar’ (recto yellow, in printed box, verso

white), printed ‘4117’ (on white paper);

e JS, pinned underside up, with a printed label ‘4340’ (on white

paper).

Catalogus Generalis: 4340 — see Appendix.

The specimens are provided now with a printed label ‘sassanides

Kollar, 1849 | SYNTYPUS | Y. Nekrutenko det. 29.11.1998’ (on

yellow paper).

satraps Staudinger, 1878

‘Pol.[yommatus] Satraps (Ld. coll.) Stgr. n. sp.’ Staudinger, 1878: 230. Type locality:

‘Unter diesem Namen hatte Lederer ein abgeflogenes & aus Külek, von ihm selbst gefan-

gen, in seiner Sammlung stecken. Haberhauer fing 1875 eine grössere Anzahl bei Tersus

(sic; recte Tarsus) und Dorak (auch Taurus)...” — Junior subjective synonym of

Polyommatus asabinus Herrich-Schäffer, [1851] (Staudinger, 1881: 65).

321

Lectotype d :

e 3d with labels: handwritten ‘Taurus | Hbhr. [Haberhauer]’ (on

yellow paper), printed ‘Origin. (on pink paper), printed ‘Zool.

Mus. | Berlin’ (on pale yellow paper), handwritten ‘28°, hand-

written (Nekrutenko) on printed form ‘LECTOTYPUS <& |

satraps Stgr. | Y. Nekrutenko design. (on yellow paper) —

Nekrutenko in Hesselbarth et al. (1995: 494).

Paralectotypes 10d, 69:

e 94,59 with circles of yellow locality label paper and printed

labels “Origin. (on pink paper); one male has handwritten

labels ‘Külek’ (on white paper) and (Lederer) ‘Satraps m.[ihi]’

(on white paper, in printed box), printed ‘Coll. Led.’ (on white

paper) and ‘Origin. (on pink paper);

e 6, 2 with labels: handwritten (Püngeler) ‘Taurus’ (on white

paper, in printed box) and handwritten (Piingeler) on printed

form ‘Cotype <underlined> | v. satraps Stgr. & (2) | vom Autor

| Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Asabinus HS. | v. Satraps Stgr.’ (on white paper, in printed

box).

schmidti Kardakoff, 1928

‘Lycaena orion Pall. ab. schmidti (nov.)’ Kardakoff, 1928: 272, Taf. 7, Fig. 18. Type

locality: ‘Gefangene dd am 9.V. in der Umg.[egend] v.[on] Wladiwostok.’ —

Infrasubspecific.

Syntype d:

e 3 with labels: printed, with handwritten (Kardakoff?) insertion

“Wladiwostok | S. Ussurigebiet | 9.V. 1927. N. Kardakoff’ (on

white paper), handwritten (Kardakoff?) ‘Lycaena orion Pall. |

ab. schmidti Kardakoff’ (on white paper) and printed “Typus’

(on red paper, in box).

The specimen is placed under a typewritten drawer label ‘ab.

schmidti’ <underlined with red pencil> (on a stripe of white paper,

in blue type ribbon).

322

scudderi Möschler, 1874

‘Lycaena Scudderi Edw. Proc. Ac. Nat. Sc. Phil. 1861, p. 164° Möschler, 1874: 155. Type

locality: ‘in einigen Exemplaren von Labrador erhielt.... — Misidentified; junior

primary homonym of Lycaena scudderi Edwards, 1861; oldest available name: Lycaena

aster Edwards, 1882 (Staudinger, 1901: 79).

Syntype d :

e 3 with labels: handwritten (Püngeler) ‘Labrador m.[eridio-

nalis] | Scudderi Möschl. 3, | Cotype, <underlined> 12. [18]75

v.[om] Möschler’ (on white paper), handwritten (Püngeler) on

printed form ‘Cotype <underlined> | Scudderi Möschl. ? |

Püngeler’ (on pink paper).

The specimen is placed under a handwritten (Püngeler) drawer

label ‘Aster Edw.’ (on white paper, in printed box). As may be seen

from the label, the specimen was received by Püngeler from

Möschler in December 1875 (he was then 18!) and was later on

designated as a ‘Cotype’. The question mark on the pink label

apparently related not to the specimen status as a syntype, but

rather to its identity with aster Edwards (cf. drawer label).

scylla Staudinger, 1887

‘Lycaena Scylla Stgr. Staudinger, 1887b: 139, pl. 16, fig. 7. Type locality: ‘...erhielt

ich in grösserer Anzahl durch Christoph, die sie von Ende Juni bis mitte Juli bei Raddefka

fand.— Junior primary homonym of Lycaena scylla Oberthür, 1880, junior primary

homonym and junior objective synonym of Lycaena scylla Lang, 1884; junior subjective

synonym of Lycaena lycormas Butler, 1866 (Staudinger, 1901: 88).

Syntypes 94,39:

e d with labels: handwritten (Staudinger) ‘Scylla | Stgr. (on

white paper), handwritten (Staudinger) ‘lycormas | Butl. (on

cross-lined notebook white paper), handwritten (Staudinger)

‘Ob Lycormas Butl.?’ (on white paper) handwritten

‘Raddefskaja | Chr.[istoph]’ (on violet-blue paper) and printed

‘Origin.’ (on pink paper);

e 73, 3% with circles of violet-blue locality label paper and

printed label ‘Origin.’ (on pink paper);

e à with labels: handwritten ‘Ussuri | Dörr.[ies]’ (on violet-blue

paper) and printed ‘Origin. (on pink paper).

323

septentrionalis Grum-Grshimaïlo

Unpublished.

Syntypes CPE

e 6 with labels: handwritten (Grum-Grshimailo?) ‘Ural mer. |

[18]88 Gr. Gr’ (on green paper), ‘Ural | Donz.[elii] | v.

Septentrio- | nalis G-G’ (on white paper, in red ink), handwrit-

ten (Grum-Grshimailo, in Russian) ‘Salda | VII [18]88’ (on

white paper);

(Grum-Grshimailo, in Russian) ‘Salda | VI [18]88’ (on white

paper).

As evidence in support that this material was collected by Grum-

Grshimailo during his expedition to the Ural, a quotation from his

itinerary description follows: ‘Next 1888 Grigoriy Efimovich (first

and patronymic names of Grum-Grshimailo), on his own funds,

with just a limited support of the Academy of Sciences of the

Zoological Museum, has undertaken an expedition to the Central

Ural. The aim of the voyage remained the same: mostly zoogeo-

graphical.

G. E. started a survey of the Central Ural from the South, from

Zlatoust, and concluded at the Irbit latitude. On his arrival back, he

forwarded all his entomological collections to the Zoological

- Museum.’ (Grumm-Grshimailo, 1947: 26).

Salda is a populated place in Sverdlovskaya Oblast’ (Russian

Federation), on a river with the same name (ca. 170 km N of

Yekaterinburs, 58° 21777 INGOs" 9):

sibirica Staudinger, 1892

‘Lyc.[aena] Optilete Fab. var. Cyparissus Hb. (var. Sibirica Stgr.)’ Staudinger, 1892d: 317.

Type locality: ‘...in Anzahl vom Kentei...’

Syitypes SC oe :

e 3 with labels: handwritten (Staudinger) ‘var. Sibirica | Stgr.’

(on white paper), handwritten ‘Kentei | [18]89 Dorr.[ies]’ (on

green paper) and printed ‘Origin.’ (on pink paper);

324

e 45,3% with circles of the green locality label paper and print-

ed label ‘Origin.’ (on pink paper).

sibirica Staudinger, 1899

‘[Lycaena] Damone Ev. var. Sibirica’ Staudinger, 1899: 139. Type locality: ‘...vom

Altai und Tarbagatai..” — Junior primary homonym of Lycaena optilete sibirica

Staudinger, 1892; oldest available name: Lycaena altaica Elwes, 1899.

Syntypes 4(6?) d:

e ¢ with labels: handwritten (Staudinger) ‘var. | Sibirica | Stgr.’

(on white paper), ‘Saisan | Hbhr. [Haberhauer]’ (on brown

paper), handwritten date label ‘1/8’, handwritten ‘66’ and print-

ed ‘Origin. (on pink paper);

e 36 with circles of brown locality label paper, two of them with

printed ‘Origin. (on pink paper);

e ?-¢ with labels: handwritten (O. Bang-Haas) large ‘Damone’

with glued handwritten (Staudinger) ‘Sibirica | Stgr’ (on white

paper, Bang-Haas in pencil, Staudinger in black ink), printed

‘Ongodai, | Altai Mts., | Berezowsky, | 1898’ and printed

‘Origin.’ (on pink paper) (apparently a subsequent designation

by Bang-Haas);

e ?-G with labels: handwritten ‘Altai alp.[es] (recto) | Damone

var. | Sibirica d | von | A. Bang-Haas 1/[19]02 (verso)’ (on

white paper, recto in printed box), handwritten (Piingeler) on

printed form ‘Cotype <underlined> | v. sibirica Stgr. 3 | v.[on]

B.-Haas | Piingeler’ (on pink paper) and handwritten (Forster)

on printed form ‘Polyommatus | (Agrodiaetus) & | damone |

sibirica Stgr. | det. W. Forster 1944 (or 47?)’ (on white paper)

— see also (Hauser & Eckweiler, 1997: 102).

In the Staudinger collection there are d and 29 with printed

labels ‘Ongodai | Altai Mts. | 3-5000 feet | June [18]98 </2 July

overwritten in red ink> A. Jacobson’, one male and one female

specimens have printed labels ‘380° and ‘381’ (on white paper

faded yellow) and handwritten (Forster) labels on printed form

‘Polyommatus | (Agrodiaetus) | damone & (®)l sibirica Stgr. | det

W. Forster 1947’ (on white paper), all have printed labels ‘Zool.

Mus. | Berlin’ (on pale yellow paper). Male specimen has a

325

handwritten (Balint) label on printed (in red ink) form ‘Lectotypus

| P (Agrodiaetus) | sibiricus Stgr. (recto) | des. by | Zs. Balint 1989.1

(verso)’ (on white paper, recto in box); female (without printed

number label) ‘Paralectotypus | P. (Agrodiaetus) | sibiricus Stgr.

(recto) | des. by | Zs. Balint 1989.I (verso)’ (on white paper, recto in

box).

The designation above is unavailable for two reasons: (1) it has

never been published; (2) the specimens designated do not belong

to the type series (not designated with Staudinger’s “Origin. labels,

apparently received by Staudinger after the original description was

published);

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Damone Ev. | v. Sibirica Stgr.’ (on white paper, in printed

box); provided with printed labels: ‘Designatio per schedam | IRRI-

TA: | non syntypus, non edita | Y. Nekrutenko det. 3.12.1998’ (on

violet paper).

sidemina Kardakoff, 1928

‘Zephyrus taxila Brem. ab. sidemina (nov.)’ Kardakoff, 1928: 271, Taf. 7, Fig. 17. Type

locality: ‘Ich habe nur ein d gefangen. ... Gefangen am 16.VII in Sidemi’. —

Infrasubspecific.

Holotype 4 (by monotypy):

e 3 with labels: printed, with handwritten insertion ‘Sidemi

Ussurigebiet | 29.7. 1921. N. Kardakoff’ (on white paper),

handwritten (Kardakoff?) ‘Zephyrus taxila Brem L. | ab sidemi-

na Kardakoff’ (on white paper) and printed “Typus’ (on red

paper, in box).

The specimen is placed under a typewritten drawer label ‘ab.

sidemina’ <underlined with red pencil> (on a stripe of white paper,

in blue type ribbon).

sieversii Christoph, 1873

‘[Lycaena] Sieversii Chr.’ Christoph, 1873: 23, Tab. 1, Fig. 1, 2. Type locality: ‘war

sowohl bei Tasch, als auch Schahkuh, von 4-10,000’, stets vereinzelt zu finden’.

326

Syntypes 46 , 9:

e 3 with labels: handwritten (Christoph) ‘Fasch <sic> |

Chr.[istoph]’ (on yellow paper), printed ‘Origin. (on pink

paper);

e 6, & with circles of yellow locality label paper and with small

date labels ‘18/7’ and ‘1/6’;

e G with labels: handwritten (Christoph) ‘Schahkuh |

Chr.[istoph]’ (on yellow paper), printed ‘Origin. (on pink

paper);

e JS with circles of yellow locality label paper and with small date

label ‘16/6’.

e 39 not designated with Staudinger’s ‘Origin. label, also have

Christoph’s small date labels.

The specimens are placed under a handwritten drawer label

“"Sıeversi | Chr. | Mirza Stgr.’ (on white paper, in printed box), each

provided now with a printed label ‘sieversii Christoph, 1873 | mirza

Staudinger, 1874 |SYNTYPUS | Y. Nekrutenko det. 1.12.1998’ (on

yellow paper) — see mirza above.

sinensis Alpheraky, 1881

‘L.[ycaena] Sinensis Alph. n. sp.’ Alphéraky, 1881: 383, tab. 14, fig. 7. Type locality:

‘district de Kouldja... sur l’un des promontoires du Tian-Chan’.

Syntypes 2 :

e d (hindwings and abdomen missing) with labels: handwritten

(Alphéraky) ‘Th.[ecla] Sinensis | Alph.[éraky] /Tian-Chan’ (on

white paper), circle of brown paper, small date label ‘14/V

[1879] and printed ‘Origin.’ (on pink paper);

e G with labels: handwritten ‘Tian Schan | Alph.[éraky]’ (on

brown paper), and a small date label ‘14/V [1879]’.

The specimens are provided now with a printed label ‘sinensis |

Alphéraky, 1881 | SYNTYPUS | Y. Nekrutenko det. 3.12.1998’ (on

yellow paper).

splendens Staudinger, 1881

‘Polyomm.atus] Splendens Stgr. n. sp.’ Staudinger, 1881: 280. Type locality: ‘Von

dieser wohl sicher hoch gefangenen Art sandte Haberhauer 20 Stücke ein...’ (I. c.: 280),

‘Ala Tau; Thian. [Schan] oc.[cidentalis] et m.[eridionalis], Iss.[yk] K.[ul] oc.[cidentalis]’

(Staudinger, 1901: 74).

Syntypes 48,49:

e J with labels: handwritten ‘Ala Tau | Hbhr. [Haberhauer]’ (on

brown paper) and printed ‘Origin. (on pink paper);

e 35,4% with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper).

standfussi Grum-Grshimailo, [1891]

‘Polyommatus Standfussi’ Grum-Grshimailo, [1891]: 450. Type locality: ‘In montibus

ad Sinin, in regione Amdo dicta, detectus’.

Syulypessery

e 4 with handwritten labels: “Thibet | Sinin Alp.[es] | [18]90

[legit] Gr. Gr.’ (on brown paper) and ‘Pol.[yommatus] |

Standfussi | G.G. | Sinin Mont.[es]’ (A. Bang-Haas, on white

faded paper);

e ° with circle of brown locality label paper.

The specimens, undoubtedly received by Staudinger from Grum-

Grshimailo, are provided now with a printed label ‘standfussi |

Grum-Grshimailo [1891] | SYNTYPUS | Y. Nekrutenko det.

26.11.1998’ (on yellow paper).

staudingeri Christoph, 1873

‘Lycaena Staudingeri Chr.’ Christoph, 1873: 26, Tab. 1, Fig. 5, 6. Type locality: ‘Ich

Syntype d:

e 3 with labels: handwritten “Tasch | Chr.[istoph]’ (on yellow

paper), small date label ‘31/5’ (on paper faded brown) and

printed “Origin. (on pink paper).

The specimen is placed under a handwritten drawer label

‘Staudingeri | Chr.’ (on white paper, in box).

328

suaveola Staudinger, 1881

‘Thecla Rubi L. und v.? Suaveola’ Staudinger, 1881: 279. Type locality: ‘...von Lepsa

und Saisan... Vier d (davon eins aus Saisan)...’

Lectotype &:

e ¢ (antennae broken off), with labels: handwritten ‘Lepsa |

Hbhr. [Haberhauer]’ (on brown paper), printed ‘Origin.’ (on

pink paper), handwritten in pencil ‘Callophrys | rubi var. |

suaveola | Stgr.’ (on white paper), printed “Zool. Mus. | Berlin’

(on pale yellow paper), handwritten (Bernardi) on printed form

‘C. suaveola Stgr. | lectotype 3 | Alexanor III : 276 | Bernardi

dét. | 1964’ (on white paper) and combined printed/handwritten

designation label ‘LECTOTYPE’ (on red paper) — Bernardi

(1964: 276); figured in Hesselbarth et al. (1995: Taf. 127,

Res 31537).

Paralectotypes 36:

e 25 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper), one of them has a handwritten label in

pencil ‘suaveola | Stgr.’ (on white paper);

e 6 with labels: handwritten ‘Saisan | Hbhr. (on brown paper)

and printed ‘Origin.’ (on pink paper).

sultan Staudinger, 1886

‘Polyommatus Sultan Stgr. n. sp.’ Staudinger, 1886: 202. Type locality: *...im Juli in

den südlichen Gebirgen bei Samarkand gefangen, die Haberhauer mir als Hazret Sultan

Gebirge angab. — Junior primary homonym and junior objective synonym of

Polyommatus sultan Lang, 1884: 368 (Nekrutenko, 1983: 11).

Syntypes 44, 2°:

e 5 with labels: handwritten (Staudinger) ‘Sultan | Stgr.’

(on white paper), ‘Prov Samark.[and] | Hbh. [Haberhauer]

[18181 (on brown paper) and printed ‘Origin. (on pink

paper);

e 35,2% with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper).

329

superba Staudinger, 1887

‘Lycaena Superba Stgr.’ Staudinger, 1887a: 51. Type locality: ‘...erhielt ich jetzt 6 d

und 2 2 aus dem Transalai von Herrn Maurer... 2 ¢ von Haberhauer aus dem Alai erhal-

ten hatte.’

Syieypes Ono

e 6 with labels: handwritten (Staudinger) ‘Superba | Stgr.’ (on

white paper), handwritten ‘Alai | [18]85 Hbh. [Haberhauer]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

‘Superba 2 von | mir als <crossed> unter | Venus-? 2

beschrieben’ (on white paper), ‘Alai | [18]85 Hbh.

[Haberhauer]’ (on brown paper) and printed, with handwritten

(Staudinger) insertion ‘Origin. | Venus ® | false’ (on pink

paper);

e JS with labels: handwritten ‘Transalai | [18]86 Maur.[er]’ (on

brown paper) and printed ‘Origin.’ (on pink paper);

e 34,29 with circles of brown locality label paper and printed

label ‘Origin. (on pink paper), one male specimen also has a

handwritten (Staudinger) label ‘Superba Stgr.’ (on white paper)

— Hauser & Eckweiler (1997: 103).

taherides Eckweiler, 1998

‘Polyommatus (Agrodiaetus) erschoffii taherides subsp. nov. Eckweiler, 1998: 225,

Farbtafel 2. Type locality (holotypus): ‘Iran, Khorasan, Kuh-e-Sorkh (Ost), S

Kallehmenar, 45 km S Fariman, 2300-2400 m’.

Paratype d :

e ¢ with printed labels: ‘Iran, Khorasan | 45 km S Fariman |

Kallehmenar 2300-2400 m | 15.-17.VII.1998 | leg. Eckweiler

548’ (on white paper, in box) and ‘Paratypus | Polyommatus

(Agrodiaetus) | erschoffii taherides subsp. nov. | ECKWEILER

1998’ (on bluish paper, in box).

tancrei Graeser, 1888

‘Lycaena Tancrei Graeser.’ Graeser, 1888: 77. Type locality: ‘Diese neue Art wurde

von mir bei Nicol.[ajefsk], auf eine Bergwiese, am oberen Laufe der Kamera, in Mehrzahl

gefangen.’

330

Syntypes d, 9:

e 6, ? with labels: printed ‘Nicolajefsk’ (on dirty white paper, in

box), handwritten (Püngeler) “Tancrei 4 | Original <under-

lined> | 11/09 ex coll. | Dieckmann’ (on white paper), hand-

written (Püngeler) on printed form ‘Cotype <underlined> |

Tancrei Graesl[er] 8 (2) | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label “Tancrei Graes.[er]’ (on white paper, in printed box).

tartarus Staudinger, 1896

‘Lycaena Orbitulus Esp. var. Tartarus Stgr. Staudinger, 1896b: 300. Type locality:

‘wurden mir von Herrn R Tancré eingesandt, dessen Sammler E. Rückbeil sie in den

Jahren 1892 und 1893 im Innern Tibets, zwischen dem Lob Noor und Kuku Noor, fand.’

— Junior subjective synonym of Lycaena orbona Grum-Grshimailo, 1891: 452

(Staudinger, 1901: 81).

Syntypes 2d, 2°:

e 6 with labels: handwritten (Staudinger) ‘Orbitulus var. |

(verso) (on white paper), handwritten “Kuku Noor | [18]94

<sic! — lapsus calami? — see above under Type locality> Rckbl.

[Rückbeil]’ (on brown paper) and printed ‘Origin.’ (on pink

paper);

e 4,29 with circles of brown locality label paper, two female

specimens also with rectangular pieces of pink paper.

tekessana Alphéraky, 1897

‘Lycaena Pheretiades Ev. var. Tekessana Alph. nova’ Alphéraky, 1897: 234, pl. 12, fig. 3.

Type locality: ‘C’est le 4 Juillet 1893 que l’expédition Roborowsky-Kozlow rencontra

cette jolie Lycaena sur le fleuve Tékesse, dans le Thian-Chan. Nous avons devant nous 6

Gi euewunen onde) cette racé-c.,

‘Syntype’ d:

e 6 with labels: handwritten (Staudinger) ‘var. Tekessana |

Alph.[éraky]’ (on white paper, handwritten ‘Kuldja | Rkbl.

[Rückbeil] [18]84’ (on brown paper) and printed ‘Origin.’ with

handwritten (Staudinger) insertion (illegible) (on pink paper).

331

This specimen does not belong to the Alphéraky type series, and

its designation by Staudinger as an ‘Original’ (apparently for inter-

nal use) is not available.

thebana Staudinger, 1895

‘Lyc.[aena] Thebana B. (?) Staudinger, 1895: 244, Taf. 9, Fig. 3. Type locality: “Unter

diesem Namen steckte ein & aus Cairo in Lederer’s Sammlung, Boisduval scheint aber

diese Art nicht beschrieben zu haben.’

Holotype 4 (by monotypy):

e 6 with labels: handwritten (Lederer) ‘Thebana B.[oisduval] |

Cairo’ (on white paper, in printed box), handwritten

(Staudinger) ‘Thebana | Stgr. (on white paper), handwritten

‘Cairo’ (on white paper), printed ‘Coll. Led.’ (on white paper)

and ‘Origin. (on pink paper).

The specimen is provided with a printed label ‘thebana

Staudinger, 1895 | HOLOTYPUS (par monotypiam) | Y.

Nekrutenko det. 28.10.1998’ (on red paper).

thetis Klug, 1834

‘Lycaena Thetis’ Klug, 1834: Decas Quarta: Signature /(i), Tab. 40, Figs 17, 18. Type

locality: ‘Habitat in Syria’.

Types: not found; Catalogus Generalis: 4557 — see Appendix.

transcaspica Staudinger, 1899

‘[Lycaena] Damone Ev. var. Transcaspica’ Staudinger, 1899: 141. Type locality:

‘...aus Transcaspien (Achal Tekke), wo sie in den an Persien grenzenden Gebirgen (wohl

bei Nochur)...’.— Junior primary homonym and junior objective synonym of Lycaena

kindermanni transcaspica [Heyne], [1895].

Syntypes 58, 32 (Lectotype?):

e 3d with labels: handwritten (unknown hand; Heyne?)

‘Kindermani <sic> | v. Transcaspica | Transcauc <sic> | [18]94

U. S. (or U. 5.?)’ (on white faded paper), printed “Zool. Mus. |

Berlin’ (on pale yellow paper), handwritten (Forster) on printed

form ‘Para-Typus ¢ | Lycaena transcaspica | Stgr. (on pink

paper), printed, with handwritten (Wagener, in pencil) inser-

tions ‘Abgebildet in Hesselbarth, | van Oorschot & Wagener: |

332

Tagfalter der Türkei | Tafel 779 Figur 2’ (on white paper) — not

figured; |

d with labels: handwritten (Staudinger) ‘var. Transcaspica |

Tekke Stgr. | N. ed.<?-illegible> (recto) | Kindermanni | v.

Transcaspica | Stgr. i. 1. (verso), a circle of yellow paper and

printed ‘Origin.’ (on pink paper);

23, 2 with circles of yellow paper (which might be interpret-

ed as “Tekke [18]94 | Wild’) and printed label ‘Origin.’ (on pink

paper) — cf. Eckweiler & Häuser (1997: 121, Pl. 5);

2 with labels: handwritten (unknown hand; Heyne?)

“Kindermani <sic> | v. Transcaspica | Transcauc <sic> | [18]94

U. S. (or U. 5.?)’ (on white faded paper), printed ‘Sammlung |

Daumiller’ (on white paper), handwritten (Forster) on printed

form ‘Para-Typus ¢ | Lycaena transcaspica | Stgr. (on pink

paper);

[Lectotype] d with labels: handwritten “Tekke | 94 Wild,

printed ‘Origin.’ (on pink paper), printed ‘295.’ (on white paper

faded yellow), printed ‘Zool. Mus. | Berlin’ (on pale yellow

paper), handwritten ‘transcaspica’ (on white paper), handwrit-

ten (Forster) “Typus | Lycaena | transcaspica Stgr.’ (on pink

paper), printed, with handwritten (Wagener, in pencil) inser-

tions ‘Abgebildet in Hesselbarth, | van Oorschot & Wagener: |

Tagfalter der Türkei | Tafel 727 Figur 63’ (on white paper) —

Hesselbarth et al. (1995: Taf. 121, Fig. 63).

e [Paralectotype] 2 with circle of yellow locality label paper,

printed ‘Origin.’ (on pink paper), printed ‘296.’ (on white paper

faded yellow), printed “Zool. Mus. | Berlin’ (on pale yellow

| Stgr.. (on pink paper), handwritten (Wagener) “Tekke [18]94 |

Wild’, printed, with handwritten (Wagener, in pencil) insertions

‘Abgebildet in Hesselbarth, | van Oorschot & Wagener: |

Tagfalter der Türkei | Tafel 727 Figur 64° (on white paper) —

Hesselbarth et al. (1995: Taf. 121, Fig. 64).

The designation of the two last-mentioned specimens as “Types’

and its subsequent publication (‘1d 2 Transcaspien, Tekke 1894

(Typen in Slg. Staudinger)’ — Forster, 1956: 80) may be

333

considered as an available lectotype designation; only these two

specimens are declared to be the syntypes of ‘Lycaena damone Ev.

var. transcaspica’ Staudinger, 1899 (Hesselbarth et al., 1995: 735,

footnote 3; see also Hauser & Eckweiler, 1997: 105).

transcaspica Staudinger, 1901

‘Cigaritis B. Acamas Klug. v. Transcaspica Stgr. Staudinger, 1901: 76. Type locality:

‘Tura oc.[cidentalis] m.[eridionalis] mont; (Nuchur)’.

Syntypes 2d:

e d with labels: handwritten ‘Tekke | [18]94 Wild.[?]’ (on yellow

paper), and printed ‘Origin. (on pink paper);

e d with circles of yellow locality label paper and printed labels

‘Origin.’ (on pink paper).

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Transcaspica | Stgr.’ (on white paper).

transcaucasica Rebel, 1901

‘Lycaena Argyrognomon Brest. var. Transcaucasica n. var. d 2’ Rebel, 1901: 166. Type

locality: ‘...gehören doch die aus Siidarmenien von Eriwan und Kulp von Herrn Korb

1898 und 1901 mitgebrachten Argyrognomon-Stiicke...’

SyMiypesno ae).

e 3, & with labels: handwritten ‘Armenia (recto) | Lyc.[aena]

argus var d (2) | v.[om] Mkorb 11/[19]01 (verso) (on white

paper, recto in printed box), printed, with handwritten insertion

‘Russ.[isch] Armen.[ien] | Kulp | 1901 Korb’ (on white paper),

handwritten (Püngeler) on printed form ‘Cotype <underlined> |

v. transcaucasica | Rbl. & (2) | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Argyrognomon Begstr. | v. Transcaucasica Rbl.’ (on white

paper, in printed box).

transiens Staudinger, 1886

‘Pol.[yommatus] Caspius Led. var. Transiens Stgr. Staudinger, 1886: 201. Type locali-

ty: “Von Haberhauer erhielt ich 1881 eine größere Anzahl im Juli bei Samarkand (in den

südlich davon gelegenen Gebirgen)...’

334

Syntypes 68,42:

e 3 with labels: handwritten (Staudinger) ‘Caspius var. |

Transiens | Stgr. (on white paper), ‘Prov. Samark.[and] | Hbh.

[Haberhauer] [18]81’ (on brown paper) and printed ‘Origin.’

(on pink paper);

e 54,49 with circles of brown locality label paper and printed

‘Origin.’ (on pink paper).

triphysina Staudinger, 1892

‘Lycaena Triphysina Stgr. n. sp. Staudinger, 1892a: 235, Taf. 3, Fig. 2. Type locality:

*...erhielt ich durch Herrn Tancré; sie wurde von Rückbeil im Kaschgar-Gebiet gefangen’.

Syntypes 48,22:

e d with labels: handwritten (Staudinger) “Triphysina | Stgr.’ (on

white paper), “Kaschgar | [18]90 Rckbl [Rückbeil]’ (on brown

paper) and printed ‘Origin.’ (on pink paper);

e 23,3% with circles of brown locality label paper and printed

labels ‘Origin.’ (on pink paper).

uliginosa Dannehl, 1921

‘L.[ycaena] argus Schiff. var. uliginosa’ Dannehl, 1921: 39. Type locality: ‘Eine der

interessantesten Erscheinungen der Beuerberger Fauna.’

Syntypes 36,92:

e 38,5% with labels: printed ‘O. Bayern | Beuerberg | c.[oll.]

Dannehl [19]20’, small handwritten date labels (on white paper,

in pencil) handwritten (Püngeler) on printed form ‘Cotype

<underlined> | v. uliginosa Dannehl 3 (2) | von Dr. Frhr

v. Rosen | Püngeler’ (on pink paper).

The specimens are placed under a handwritten (Piingeler) draw-

er label ‘Argyrognomon Bgstr. | v. Uliginosa Dannehl’ (on white

paper, in printed box).

unicolor Kardakoff, 1928

‘Zephyrus taxila Brem. f. 2 unicolor (nom. coll.)’ Kardakoff, 1928: 271, Taf. 7, Fig. 13.

Type locality: ‘...aus dem Ussurigebiet...’ — Infrasubspecific.

339

Syntype ©:

with handwritten (Kardakoff?) insertion ‘Sidemi Ussurigebiet |

6.8. 1921. N. Kardakoff’? (on white paper), handwritten

(Kardakoff?) ‘Zephyrus taxila Brem L. | f. 2 unicolor

Kardakoff’ (on white paper) and printed “Typus’ (on red paper,

in box).

The specimen is placed under a typewritten drawer label ‘f. 2

unicolor’ <underlined with red pencil> (on a stripe of white paper,

in blue type ribbon).

vandalusica Staudinger, 1871

‘Thecla Spini Schiff. S. V. ab. 2 Lynceus Hb. 674-5 (Lynceus Esp. n. syn.); Vandalusica

Ld. z. b. 1852, p. 19°. — Unjustified emendation of vandulusica Lederer, 1852 (see

below); the name has been already emended by Lederer himself (1857: 32): ‘Lycaena Fab.

spini S. V. V.[arietas] vandalusica m. (Lynceus Hb.)’.

Types: not found.

vandulusica Lederer, 1852

‘Thecla Lynceus Hb. v. Vandulusica’ Lederer, 1852: 19. — A replacement name for junior

secondary homonym Thecla spini lynceus (Hiibner, [1813]), non Thecla lynceus (Esper

[1777]), junior subjective synonym of Papilio spini [Denis & Schiffermiiller], 1775 (cf.

Lederer, 1857: 32; Staudinger, 1871a: 7 as vandalusica) — see vandalusica above.

venus Staudinger, 1886

‘Lyc.[aena] Venus Stgr. Staudinger, 1886: 211. Type locality: ‘Diese prächtige neue

Lectotype 6:

e d with labels: circle of brown paper, printed ‘Origin.’ (on pink

paper), handwritten (Lukhtanov) on printed form ‘Lectotypus

3 | venus Stgr. design. V. Lukhtanov’ (on white paper, recto red,

verso white) — Lukhtanov & Lukhtanov (1994: 272).

Paralectotypes 58, 49:

e 3 with labels: handwritten (Staudinger) ‘Venus | Stgr (on

white paper), and ‘Alai | [18]85 Hbhr. [Haberhauer]’ (on brown

paper), printed ‘Origin. (on pink paper);

336

e 45,42 with circles of brown locality label paper and printed

label ‘Origin.’ (on pink paper); one & destroyed, head, thorax

and abdomen removed, forewings pinned to the specimen in a

piece of notebook paper; labelled with a new handwritten label

‘Alai | 85 Htsch’ <sic> and printed label ‘Zool. Mus. | Berlin’

(on pale yellow paper).

violaceus Staudinger, 1892

‘Pol.[yommatus] Dispar Haw. Var. Violaceus Stgr. (und var. Dahurica Graes.?)’

Staudinger, 1892d: 315. Type locality: ‘...Ende Juli, auf einer hoch-gelegenen

Waldwiese des Kentei gefangen.’

Syntypes 36, 2:

(on white paper), ‘Kentei | [18]89 Dörr.[ies]’ (on green paper)

and printed ‘Origin. (on pink paper):

e 3d with circles of green locality label paper and printed

‘Origin.’ (on pink paper).

virgaureola Staudinger, 1892

‘Polyommatus Virgaureae L. var. Virgaureola Stgr. Staudinger, 1892d: 314. Type

locality: ‘...auf höher gelegenen Wiesen im Kentei’.

Syntypes 54,79:

e 3 with labels: handwritten (Staudinger) ‘var. | Virgaureola |

Stoer. (on white paper), “Kentei | [18]89 Dörr.[ies]’ (on pale

green paper) and printed ‘Origin.’ (on pink paper);

e 34,69 with circles of pale green locality label paper and print-

ed ‘Origin.’ (on pink paper); one male specimen also has a

handwritten (Staudinger) label ‘var. | Virgaureola’ with an addi-

tion in pencil ‘Kentei’ (on white paper);

e G, 2 with labels: handwritten (Püngeler) “Transbaical |

(Kentei) | Stgr. (recto) | Polyo.[mmatus] virgaureae | v. virgau-

reola Stgr. | d (2) | ex coll. Heydenr.[eich] | 9.[18]98 (verso)’

(on white paper, recto in printed box) and handwritten

(Püngeler) on printed form ‘Cotype <underlined> | v. virgaure-

ola Stgr. 3 (2) | Püngeler’ (on pink paper).

331

The specimens are placed under a handwritten (Püngeler) draw-

er label ‘Virgaureae L. | v. Virgaureola Stgr.’ (on white paper, in

printed box).

xerxes Staudinger, 1899

‘[Lycaena] Damone Ev. var. Xerxes’ Staudinger, 1899: 143. Type locality: ‘...wurde

von Christoph bei Schahrud in Nord-Persien im Mai 1873 gefangen; er sandte mir 4d &

und 32 2 davon.’

Lectotype d:

e d with a circle of yellow locality label paper, printed ‘325’ (on

white paper faded yellow), printed ‘Origin.’ (on pink paper),

handwritten (Forster) on printed form ‘Lecto-Typus & |

Polyommatus xerxes | Stgr. | W. Forster München’ (on pink

paper, in box) — Forster (1956: 122); Hauser & Eckweiler

(1997: 108).

Paralectotypes 38, 32:

e 3 with labels: handwritten (Staudinger) ‘Kindermanni | v.

Xerxes | Stgr. (on white paper), handwritten ‘Schahrud |

Chr.[istoph]’ (on yellow paper), handwritten small (Christoph)

date label ‘10/5’ (recto white, verso green), printed ‘Origin.’ (on

pink paper), handwritten (Forster) “Cotypus | Lycaena damone

| xerxes Stgr.’ (on pink paper);

e 23,3% with circles of yellow locality label paper and printed

‘Origin.’ (on pink paper), one male and one female have

also a handwritten (Forster) label ‘Cotypus | Lycaena damone |

xerxes Stgr. (on pink paper), some of these also have

Christoph’s small date labels ‘20/5’, ‘22/5’, ‘26/5’ (recto white,

verso green). |

The specimens are placed under a handwritten (Staudinger)

drawer label ‘v. Xerxes.’ (on white paper).

zeituna Forster, 1960

‘Agrodiaetus phyllis zeituna (Stgr. i. 1.) ssp. nov.’ Forster, 1960: 129, Taf. 10, Fig. 23, Taf.

11, Fig. 23. Type locality: ‘Zeitun’ (Turkey, Maras, Süleymanlı).

338

Holotype à:

e d with labels: handwritten (Staudinger) ‘Iphigenia v. | Zeituna

i. 1. (on white paper), handwritten ‘Zeitun | [18]98 Har.<?>‘(on

white paper faded yellow), printed ‘306.’ (on white paper faded

yellow), printed ‘Origin.’ (on pink paper), printed ‘Zool. Mus. |

Berlin’ (on pale yellow paper), handwritten (Forster) on printed

form ‘Holo-Typus <overwritten to correct ‘Type’> à |

Polyommatus (Agrodiaetus) | phyllis zeituna Forster |

W. Forster München’ (on pink paper), printed, with handwritten

(Wagener, in pencil) insertions ‘Abgebildet in Hesselbarth, | van

Oorschot & Wagener: | Tagfalter der Türkei | Tafel /2/ Figur

27’ (on white paper) — Forster (1960: Taf. 10, Fig. 23, Taf. 11,

E22 9)Hesselbarth er al (1995: Taf: 121, Fis. 27); Häuser &

Eckweiler (1997: 108).

Paratype d:

e d with labels: a circle of yellow paper, printed ‘307. (on white

paper faded yellow), printed ‘Zool. Mus. | Berlin’ (on pale yel-

low paper), handwritten (Forster) on printed form ‘Para-Typus

6 | Agrodiaetus phyllis | zeituna Forster | W. Forster München’

(on pink paper).

zephyrinus Christoph, 1884

‘Lycaena Zephyrinus Stgr. Christoph, 1884: 102, pl. 4, fig. 3a, b. Type locality: ‘Im

Mai bei Askhabad... spater bei Nuchur gesammelten aus.’

Syntypes 6, ®

e d with handwritten (Staudinger): labels: ‘als Zephyrinus | von

Christoph’ (on white paper), ‘Nochur Teke | Christ.[oph]’ (on

yellow paper), and small label ‘75/6’;

[29 May 1882] (recto) | Chr.[istoph] (verso)’ (on white paper,

recto in violet box).

Other syntypes with Christoph’s identical handwritten labels are

deposited in the Zoological Institute, Russian Academy of

Sciences, St.-Petersburg.

339

The specimens are provided with a printed label ‘zephyrinus

Christoph, 1884 | SYNTYPUS | Y. Nekrutenko det., 29.10.1998’.

zephyrinus Staudinger, 1886

‘Lyc.[aena] Zephyrus HS. var. Zephyrinus Stgr.’ Staudinger, 1886: 204. Type locality:

“...erhielt ich besonders aus der Umgegend von Samarkand und Namangan, wo sie Ende

Mai flog, aber auch von Usgent und Osch, wo sie Mitte Juli gefangen wurde. — Junior

primary homonym of Lycaena zephyrinus Christoph, 1884 (Staudinger, 1901: 79);

replacement name: Plebeius pylaon usbeka Forster, 1939: 6.

Syntypes 78,98:

e d (abdomen removed), with printed labels ‘Origin. (on pink

paper) and ‘Zool. Mus. | Berlin’ (on yellow paper), handwritten

(Balint) on printed form ‘Plebejides | usbekus FORST. | gen.

prep. No. | Zs. Balint 37’, handwritten (Balint) on printed form

‘Lectotypus | Lycena zephyrinus | STGR. | Plebeius | usbeka

FORSt. <sic>‘(print in red ink, in box), “Lectotypus | Plebejides

| usbekus FORST. (recto) | det. BALINT, | 1986.XII (verso) (on

white paper, print in red ink, in box);

e 3 (right forewing missing), with labels: handwritten

(Staudinger) ‘Zephyrus var. | Zephyrinus | Stgr (on white

paper), handwritten ‘Prov. Samarkand | Hbhr. [Haberhauer]

[18]81’ (on brown paper), printed ‘Origin.’ (on pink paper);

e 34,49 with circles of brown locality label paper and printed

labels ‘Origin.’ (on pink paper); one male specimen also has a

handwritten (Staudinger) label ‘Zephyrus var. | Zephyrinus

Stgr.’ (on white paper);

e ? with circle of brown locality label paper, printed labels

‘Origin.’ (on pink paper) and ‘Zool. Mus. | Berlin’ (on yellow

paper), handwritten (Balint) labels on printed form

‘Paralectotypus | Lycena zephyrinus | STGR. = Plebeius | usbe-

ka FORST.’ (on white paper, print in red ink, in box),

‘Paralectotypus | Plebejides | usbekus FORST. (recto) | det.

BALINT, | 1986.XII (verso)’ (on white paper, print in red ink, in

box);

(on brown paper), handwritten small date label ‘13/6’ (on white

paper) and printed ‘Origin. (on pink paper);

340

e 3 with labels: handwritten ‘Namangan | Hbhr. [Haberhauer]

[18184 (on brown paper) and printed ‘Origin. (on pink paper);

e 3, 2? with circles of brown locality label paper and printed

‘Origin. (on pink paper); the male specimen also has a hand-

written small date label ‘28/5’, female specimens ‘27/5’ and

‘28/5’ (on white paper);

[18]83’ (on brown paper) and printed ‘Origin.’ (on pink paper).

No information related to these specimens in connection with the

lectotype designation has been published yet by Balint (‘I have

examined closely the syntypes of zephyrinus (Staudinger)... —

Balint & Kertész, 1990: 203-204).

Acknowledgements. I am much indebted to Dr. Wolfram Mey who

made my visit to the Museum für Naturkunde der Humboldt

Universität zu Berlin possible and who provided a most favourable

atmosphere that allowed me to focus entirely on the project imple-

mentation. I kiss the hand of Miss Viola Richter for her everyday

attendance, assistance, generous help and friendly encouragement

during my stay in Berlin. During the preparation of this text, I

kept in permanent e-mail touch with Mr. Alain Olivier (Antwerp)

who kindly helped to solve certain important problems with

literary sources and advice. Mr. Alain Olivier and Mr. & Mrs. Jos

and Jeannine Dils (Stabroek-Hoevenen, Belgium) visited me

during my stay in Berlin to participate, encourage and assist my

efforts. Dr. Mary E. Petersen, Zoological Museum, University of

Copenhagen (Denmark), kindly read through the introductory

section. The real joy of any work is much increased when one has

somebody to share the impressions and to feel unlimited under-

standing and perpetual counter-interest; in this respect, in addition

to the forementioned persons, I am greatly indebted to Mr. Manfred

Gertsberger, Mr. Matthias Nuß, Dr. Wolfgang Speidel and

Mr. Vadim V. Tshikolovets for fascinating and unforgettable

discussions that removed all doubts, ambiguities and uncertainties

that appeared during this study and who shared the delight of

discoveries.

341

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344

kol. Taf.: 1(1-16): 182 S., 96 Taf., ([1831]-1833); 2(17-32): 162 S., 96 Taf.,

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1-23, a-h, k, S. 1-184, Index [1]-34, Taf. Geometrides 1-91 (1843-1856); 4

(Hefte 27-30, 33-34, 39-40, 44, 46, 48, 53, 55, 57, 60, 63, 68): Sign. 1-36,

a-m, S. 1-288, Index [1]-48, Taf. Pyralides 1-23, Taf. Tortricides 1-59

(1847-1855); 5 (Hefte 22, 30, 34, 39, 42, 44, 46, 48, 53, 55, 57-68): Sign.

1-50, a-n, S. 1-394, Index [1]-52, Taf. Tineides 1-124, Taf. Pterophides 1-7,

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350

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(ST)

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Appendix

An extraction from Catalogus Generalis Musei Zoologici Berolinensis. Vol. 1 (inven-

tory nos 1-3753), Vol. 2 (inventory nos 3754-10098) showing recorded type speci-

mens of the Lycaenidae.

Inventory | Species Specimens Locality Collector/

no. no. Provider

Iolaus Ismenias Kl. Ambukol Ehrenberg

Aphnaeus Acamas Kl. 5 Syria Ehrenberg

Aphnaeus Acamas Kl. 1 Arabia fel. Ehrenberg

Deudorix Livia Kl. | Arabia fel. | Ehrenberg

4118 rege. sake [1] [Aegypt] [Ehrenberg]

4340 Thecla Sassanides Koll. 2 Persia

4341 | Thecla Myrtale Kl. 3 Syria

4353 Thecla Spini F. Var. 2 Syria Ehrenberg

Melantho

*4554 Chrysophanus Thersamon | 7 Syria Ehrenberg

F. Var. Omphale KI.

*4557 Chrysophanus Thetis Kl. Syria Ehrenberg

4679 Lycaena Astraea Frey. Amasia

4713 | Lycaena Psylorita Friv. 2 Creta

4715 Lycaena Rippertii Bsd. + Alp. Gallopr.

4721 Lycaena Hopfferi Kind. Amasia

4723 | Lycaena Actis HSch.

4724 Frivaldszky

Lycaena Iphigenia Friv.

Missing specimens are marked with an asterisk.

532

Nota lepid. 23 (3/4): 353; O1.XI1.2000 ISSN 0342-7536

Vol 23 — 2000

Dates of publication — Publikationsdaten — Dates de publication

23 (1): 01.IV.2000 pp. 1-84

23 (2): O1.VIL2000 pp. 85-190

23 (3/4): 01.XI.2000 pp. 191-353

Contents — Inhalt - Sommaire

New taxa described in Vol. 23

Neue Taxa in Band 23 beschrieben

Nouveaux taxa décrits dans le Vol. 23

PSYCHIDAE

Manneouarawysnica Rütjan, 2000 .........................................

NYMPHALIDAE

Brebiaslavejasciata juliana Cupedo, 2000 ...............................

LASIOCAMPIDAE

Euanosoma Zolotuhin & Kostjuk, 2000 ...............................--

Phantosoma witti Zolotuhin & Kostjuk, 2000 ..........................

181

141

142

353

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Trans.R.ent.Soc.Lond. 101: 435-489, figs. 1-44, 7 maps.

HIGGNS, L. G. & RILEY, N. D., 1980. A field guide to the butterflies of Britain and Europe. 4th ed. — Collins,

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