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DEEL 115 1972

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

| Tijdschrift voor Entomologie, deel 115, 1972 |

Aflevering 1 verscheen 10 maart 1972

Afleveringen 2—3 verschenen 21 april 1972

Afleveringen 4—5 verschenen 28 juli 1972

Aflevering 6 verscheen 10 november 1972

Afleveringen 7—8 verschenen 12 december 1972

INHOUD VAN DEEL 115

BELLE, J. — Further studies on South American Gomphidae (Odonata) .

DIAKONOFF, A. — Remarks on Matsumuraeses Issiki, with descriptions of new

species (Lepidoptera, Tortricidae, Laspeyresiini) .

Jong, R. DE. — Systematics and geographic history of the genus Pyrgus in the

Palaearctic region (Lepidoptera, Hesperiidae) .

HOLLANDER, J. DEN & H. Lor. — Differential use of the habitat by Pardosa pul-

lata (Clerck) and Pardosa ea = iù; in a mixed ns Sai

neae, Lycosidae) ‘ :

LIEFTINCK, M. A. — Further studies on Old World Melectine bees, with stray

notes on their distribution and host relationships (Hymenoptera, Anthophori-

dae)

LiTH, J. P. van. — Contribution to the knowledge of Oriental Psenzlus (Hyme-

noptera, Sphecidae, Psenini)

LukoscHus, F. S., A. Fain & F. M. DRIESSEN. — Life cycle of sn

apodemi (Fain, 1965) (Glycyphagidae: Sarcoptiformes)

SIMON THOMAS, R. T. & A. M. J. SIMON THOMAS. — Some observations on the

behaviour of females of Philanthus triangulum (F.) (Hymenoptera, Spheci-

dae) TER u En et ee te Hes Magne sc

SIMON THOMAS, R. T. & E. P. R. PoORTER. — Notes on the behaviour of males

of Philanthus triangulum (F.) (Hymenoptera, Sphecidae) .

217

241

205

293

153

125

141

pt IT We CTO

‘

va e

tri radars entra) A manch bre afha steal aoa

„al | i A gi u f i ai u 4 A

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L u ol nn Î

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1 { i

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Le. (ANT

DEEL 115 AFLEVERING 1 1972

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

R. DE JONG. — Systematics and geographic history of the genus Pyrgus in the

Palaearctic region (Lepidoptera, Hesperiidae), p. 1—121, tekstfig. 1—60, pl.

16.

Tijdschrift voor Entomologie; deel 115. afl. 1 Gepubliceerd 10 3-1972

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SYSTEMATICS AND GEOGRAPHIC HISTORY OF THE GENUS

PYRGUS IN THE PALAEARCTIC REGION

(LEPIDOPTERA, HESPERIIDAE)

R. DE JONG

Rijksmuseum van Natuurlijke Historie, Leiden

ABSTRACT

A systematic and zoogeographic study on the Palaearctic species of the genus Pyrgus, comprising

a taxonomic review, a description of the geographic variation and a reconstruction of the geographic

history. Some zoogeographic problems are discussed and the Pyrgus species are classified as faunal

types.

CONTENTS

1. Introduction 2

a. Scope O: 2

b. Taxonomic renee a: SRO Ende HF ee al 2

c. Wing markings and genital armature . 6

d. Nomenclature bm Mer oe ANR E 6

e. Material 7

fi. ea Arda a: 7

g. Methods, measurements and abbreviations . . . . . . . . . . . . . . . 8

2. The genus Pyrgus Hübner, 1819 . 8

a. Delimitation . . DONNE ET A oe ee eo oe 8

bir Glassificationi © 0 4 ws 0 nn ow ee nn a ee 10

SubpenuseScelowrix ee IR

Subpenuseb year en ee Rem eo e I

ihevalversecomplexiey nee Se M Pa u ae we TG

Species of uncertainsaffinities … = 9% 6 2 ons mon «© = 5 à à à 23

he American Pyrg25 species, … … nen ve mn ee SO

c. Distribution . . ee ee ss de de en a a Ua a ‚24

d. Geographic and Eloi kon on Pao soar Sei, Swe eh at ds sal 25

Geographic history of the subgenus Sceloirix SR MATRA) Sires tp chalk CCT en 277,

Geographic history of the alveus species complex. . . . . . . . . . . . . 30

2 l;hesspeciessofs PyrgussinsthenPalaearctien. tana to Gos oo. se Cu, NU a 31

4. Discussion of some zoogeographic problems . . . EN Gos waldo IE" ae 99

a. The geographic origin of species and groups of species. . . . . . . . . . . 99

b. The geographic history of the Palaearctic Pyrgus species, in relation to the general

history of the Palaearctic fauna . . . . ‘ Malas lw Ge Dee se de un 100

5. Zoogeographic classification of the Palaearctic Pyrgus species. . . . . . . . . . 105

(kt. TER ONE ee trata SR cts af het eds. et te SE 107

Jen Eiterature MENEN APV ley Mere Waites a ea Gs Gr dhl e ee ue II

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

1. INTRODUCTION

1.a. Scope

The present study is an attempt to reconstruct the geographic and, in a few cases,

the evolutionary history of the Palaearctic species of the genus Pyrgus, on the basis

of their recent ranges and geographic variation patterns. The author believes that the

high morphologic and ecologic diversity of Pyrgus can be understood as, at least

partially, the result of geographic isolation and subsequent differentiation of former

geographically connected populations.

The history of Pyrgus, reconstructed in this way, clearly shows the enormous impact

of the alternation of Glacial and Interglacial Periods during the Pleistocene, on the

distribution and differentiation of the members of Pyrgus. Details of the history, how-

ever, are often vague.

The history of Pyrgus is discussed in relation to the general history of the Palaearctic

fauna. This leads towards a zoogeographic classification of the Pyrgus species into

faunal types; a comparison is made with a classification into distributional types.

As taxonomic research must provide the basic facts for the reconstruction of the

history, only taxonomically well-known groups can be selected for a historic-zoogeo-

graphic study. The Palaearctic members of the genus Pyrgus are such a group. This is

mainly due to the work of Rambur (1837-1842), who discovered the great importance

of the male genitalia for the identification of the species, of Reverdin (from 1910

onwards), of Warren (1926, 1953) and of Evans (1949). Moreover, Alberti, Kauffmann,

Opheim, Picard, and others have contributed to our knowledge, in dealing with detail

problems. Nevertheless, there still exists much disagreement about the taxonomy of

Pyrgus, especially with regard to the delimitation of species and subspecies. Therefore,

it was necessary to review the taxonomy of Pyrews, before a zoogeographic study of

the genus could be made. It must be emphasized, however, that this review is not the

aim of the present study, but only a means to permit a reconstruction of the geographic

and evolutionary history of this interesting group, which has fascinated the author for

many years.

1.b. Taxonomic categories

The influence of the application of the biological species concept on the number of

species within the genus Pyrgzs in the Palaearctic becomes apparent, when we compare

the 32 species recognized by Warren (1926) on the base of a typological species concept,

with the 19 species recognized by Evans (1949) who accepted a biological approach.

The typological and biological species concepts have extensively been discussed by Mayr

(1942, 1963). It would suffice here to refer to these publications, if there were not

many authors who in recent times have still applied a typological species concept

in entomology. Three examples concerning Hesperiidae are:

(a) Heydemann (1954: 426): “Carcharodus Hb. swinhoei Watson ist gute Art und

von der äusserlich ausserordentlich ähnlichen fritillarius Poda (= alceae Esp.) durch

verschiedenen Genitalapparat des Männchens getrennt’.

(b) Warren (1957: 375), having examined the female genitalia of the N. American

Pyrgus freija and the Scandinavian Pyrgus centaureae, concludes: “These mounts proved

R. DE JONG: Pyrgus in the Palaearctic region 3

beyond doubt that P. freija and P. centaureae were distinct species, ...”

data that Pyrgus malvae, P. malvoides, P. melotis and P. ponticus were distinct species.

Palaeo-zoogeographic data can never prove conspecificity, nor the reverse; morphologic

data, on the other hand, sometimes can, in combination with chorologic data, viz. when

the study concerns contact zones of morphologically different forms. Sichel only showed

that there are morphologic differences between the four forms just mentioned and that

these forms are allopatric.

In these and in many other cases, morphologic differences between specimens and not

non-interbreeding of populations are considered decisive for specific distinction. As

long as the typological species concept will be applied, there will be much disagreement

about classification. It may be useful to quote here the definition of the (biological)

species of Mayr (1942: 120): “Species are groups of actually or potentially interbreeding

natural populations, which are reproductively isolated from other such groups.” This

concept will be adhered to in the present study.

The dynamic character of the species is apparent in those cases where neighbouring

populations interbreed only to a limited extent, showing that these populations have

attained a certain degree of specificity with regard to each other. Such cases are by no

means rare; in Pyrgus strikingly limited interbreeding is known in groups which in the

present work have been called superspecies malvae, superspecies carlinae and the alvens

species complex. It does not seem relevant to the author at what percentage of hybrids

we can consider the populations, which come into contact with each other, specifically

distinct. In the superspecies malvae and carlinae I term the component species, which are

only partially reproductively isolated, semispecies (see Lorkovic, 1953; Mayr, 1963,

1969). Further research in the a/vews species complex may make it preferable to apply the

term semispecies to what has been called here accretus subspecies group and to P. war-

renensis (see also Chapter 2).

The term semispecies was proposed by Mayr for the designation of the component

species of a superspecies (see Mayr, 1963). Mayr (1969: 52) defined the superspecies

as follows: “A superspecies is a monophyletic group of closely related and largely or

entirely allopatric species’. By broadening the term semispecies in order to include all

border-line cases in speciation (see Mayr, l.c.) it may be useful to broaden also the term

superspecies, to include not only semispecies which are entirely reproductively isolated,

but also semispecies which are partially reproductively isolated. In this sense I apply

the term superspecies, to malvae and carlinae. By extending the term superspecies, it

becomes clear that in specific cases there is not an essential difference between species

and superspecies, but only a difference in degree of speciation.

Similarly, superspecies and species groups do not differ essentially but only in degree

of distributional overlap. The present author prefers to use the term species group

instead of the term subgenus, as the former has no nomenclatural consequences. In some

cases it may have practical value to unite species groups into a subgenus, e.g. the sub-

genus Scelotrix of Pyrgus. The advantage of recognizing a subgenus for a single

superspecies, as subgenus Pyrgus for superspecies malvae, is at least disputable. Of

course, any decision with regard to the problem ‘genus versus subgenus’’ is entirely

subjective. This applies also to the problem “subgenus versus species group” and even

“genus versus species group”.

Lepidopterists are notorious for their treatment of infraspecific variation. The immense

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

number of names for all kinds of variations has little contributed to the understanding

of the variation. Only the geographic variation is important for the present study. The

treatment of this geographic variation by various authors does not always show a clear

understanding of the subject. Most, if not all, difficulties arise from different views

(if there is any view at all) about the subspecies concept. Some authors apply the term

subspecies to any geographic variation, hereditary or not; others confuse subspecies with

the occurrence of certain characters and describe then even sympatric “subspecies” of the

same species. Kauffmann (1951) succeeded in combining both not current conceptions

of the term subspecies. He observed e.g. that the “subspecies” caecus and serratulae of

Pyrgus serratulae fly together in Switzerland and that the ratio between them changed

annually and depended on weather conditions. The extensive literature concerning the

subspecies (apparently overlooked by many lepidopterists) makes further extensive

discussion of the problems here senseless; it may suffice to refer to Mayr (1969) for a

recent treatment of the subject and for many references. Mayr (l.c.: 41) defined the

subspecies as follows: “A subspecies is an aggregate of phenotypically similar populations

of a species, inhabiting a geographic subdivision of the range of a species, and differing

taxonomically from other populations of the species.”

We should be aware of the fact that this definition may easily lead to the recognition

of subspecies which are evidently taxonomic artifacts and therefore of no use in tax-

onomy. For example, Mayr (1963: 349) states: “Nor should the subspecies be confused

with phenomena of a very different nature, such as character gradients (clines).” How-

ever, his definition is not at all a guarantee against a confusion. Furthermore, misunder-

standing of Mayr’s definition could easily lead to the recognition of nongenetic

ecophenotypic varieties as subspecies. For example, a common nongenetic ecophenotypic

variation in Pyrgus at high altitudes concerns the reduction of white spots on the upper-

side of the wings. Most specimens of P. serratulae in the Alps at altitudes above 2000 m

have very small spots; they are known under the name “forma caecws’’. Evidently, the

recognition of a “subspecies” caecus is of no use in taxonomy, as this “‘subspecies’’

would be a taxonomic artifact.

Virtually Mayr's subspecies concept would be entirely typological, if it were not

based on a biological species concept. Mayr has never stated that a subspecies has to be

phenotypically uniform under all conditions, but I am of the opinion that especially the

typological concept has caused many difficulties. If we abandon a typological idea

altogether and admit that a subspecies is not only a taxonomic but also a zoogeographic

concept, it seems possible to avoid any confusion and to define a subspecies representing

a natural unit, as follows:

A subspecies is an, at present or formerly, geographically isolated group of populations

of a species, which can be distinguished by one or more characters from other such

groups of populations of the species.

The existence of such subspecies can be concluded from the idea of speciation through

geographic isolation. The subspecies thus defined is a useful tool for a historic-

zoogeographic study. The fact that it is often very difficult to recognize a subspecies,

cannot be a serious objection. However, we must be aware that not all geographically

isolated populations differentiate into recognizable subspecies (for example, see Chapter

3, P. andromedae). For the subjective nature of the decision whether a geographic

isolate differs sufficiently from other populations to warrant subspecific separation, see

Mayr (l.c.). Some attention will be paid to the question, how groups of populations,

R. DE JONG: Pyrgus in the Palaearctic region 5

5 È Di

gen. pl.

antevag. pl.

Fig. 1—3. Wing markings, male and female genital armatures in Pyrgus. 1. Wing markings: bas.

= basal spots; discoid. = discoidal series; med. = median series; subcost. = subcostal spots;

submarg. = submarginal series. 2. Male genital armature, lateral view, with left valve removed:

aed. = aedeagus; an. sh. = anal sheath; antist. = antistyle; cuill. = cuiller; ha. — harpe; lam.

tr. = lamina transversa; lat. ap. — lateral apophyses; st. = style; stf. = stylifer; subh. pl.

= subharpal plate; teg. = tegumen; unc. = uncus. 3. Female genital armature, ventral view:

antevag. pl. = antevaginal plate; b.c. = bursa copulatrix; d.b. = ductus bursae; gen. pl. = genital

plate; ost. = ostium; pap. an. = papilla analis; VIII abd. segm. = eighth abdominal segment

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

which were formerly geographically isolated, can be recognized. This can only be done

by tracing the existence of zones of secondary intergradation. Of course, our decision

whether an intergradation zone is primary or secondary, may be incorrect, but this is not

due to an artificial subspecies concept, but to a wrong interpretation of the facts. A

discussion on zones of intergradation can be found in Mayr (1963: 368-384).

The subspecies thus recognized may be very variable, even geographically. Un-

doubtedly, this geographic variation within the subspecies is partly clinal, partly ecophe-

notypic (as defined by Mayr, 1969: 402): “A nongenetic modification of the phenotype

by specific ecological conditions, particularly those of a habitat”). It has often been

emphasized that it is undesirable to give scientific names to these variations. Only the

most striking ones of the described forms have been mentioned in this paper (as

“forma’’). Due to lack of material or data it is at the moment often impossible to

decide whether a variation is ecophenotypic or clinal. In the present study I am

attempting to separate subspecific, clinal and ecophenotypic variations. I am aware of

the fact that the results will not always be convincing, but recognition of these types of

variation may be a stimulus to fill the many gaps in our knowledge.

1.c. Wing markings and genital armature

In the present paper a combination has been adopted of the terms used by Warren

(1926) and Evans (1949) to denominate the wing markings and the various parts of

the male genitalia. For the female genitalia I have followed Opheim (1956) and Guil-

laumin (1966).

(a) Wing markings (Fig. 1).

Fore wings — from termen to base there are three series of spots: a submarginal, a

median, and a discoidal series. In addition, there are subcostal spots, a cell spot (in the

cell) and a discal spot (closing the cell). The separate spots are indicated by their

intervenal space, e.g. median spot in space 6, submarginal spot in space 2, etc. (for the

numbering of the veins and the intervenal spaces, see Evans, 1949). The submarginal

spots are often absent or vague.

Hind wings — from termen to base there are three series of spots: a submarginal,

a median, and a basal series. The separate spots are numbered as on the fore wings.

The median spots in spaces 4 and 5 are confluent and indicated as the central spot.

The spots on the upperside of the hind wing are often very faint.

(b) Male genitalia: Fig. 2.

The bursa copulatrix lacks a signum, a structure often important as a diagnostic,

specific character in other groups of Lepidoptera.

1.d. Nomenclature

Little attention has been drawn to matters of nomenclature and it has not been at-

tempted to list all names relating to Pyrgus and its species. A full account of these

names can be found in Evans (1949). Names proposed after 1949 have only been in-

cluded here when they refer to species or subspecies or when they relate to forms which

I think to be worth mentioning.

In one respect I have intentionally neglected the International Code of Zoological

R. DE JONG: Pyrgus in the Palaearctic region 7

Nomenclature, viz. by omitting the parentheses enclosing the author's name in new

combinations, as meant in Article 51d of the Code. The use of parentheses does not add

necessary, important or even interesting information. It only indicates that the author

of the species-group name (in nomenclatural sense) listed the taxon represented by this

name in another genus, as can be expected when names are rather old. If one wishes to

use parentheses, the names of the authors of all specific and most subspecific names of

Pyrgus can be enclosed in parentheses.

1.e. Material

The material examined for this study comprises more than 10.000 specimens. Though

this number does not appear very small, much more material is needed to solve some of

the taxonomic problems. I must, however, stress the fact that many problems can only

be solved by a study on the spot and not by examining large numbers of specimens in

collections. In the present work such problems are apparent a.o. in the alveus species

complex and the superspecies malvae.

Special mention must be made of the large collections of Hesperiidae in the British

Museum (Natural History). These collections, containing e.g. the very important Ch.

Oberthiir collection, have been arranged by the great specialist of Hesperiidae, the late

Brigadier W. H. Evans, whose extensive work on the Hesperiidae of the world was

based on this material. I am glad to have had the opportunity of examining this

material during a visit to the British Museum in the spring of 1971.

1.f. Acknowledgements

For the loan and gifts of material, for valuable information and for help in various

ways I am greatly indebted to the following persons and institutions: Dr. B. Alberti

(Waren-Müritz, DDR) (abbreviated: BA), G. Bernardi (Muséum National d'Histoire

Naturelle, Paris) (MP), F. Bink (Amsterdam), H. Coene (Amsterdam) (HC), Dr.

W. Forster (Zoologische Sammlung des Bayerischen Staates, Munich) (ZSM), M. Guil-

laumin (Paris) (MG), Dr. H. J. Hannemann (Zoologisches Museum der Humboldt-

Universitat, Berlin) (ZMHB), F. C. Howarth (British Museum, Natural History,

London) (BM), Instituut voor Taxonomische Zoologie (Zodlogisch Museum, Am-

sterdam) (ZMA), Dr. F. Kasy (Naturhistorisches Museum, Vienna) (MW), A. Lille-

hammer (Zoologisk Museum, Oslo) (MO), J. H. Lourens (Amsterdam), Dr. E.

Niculescu (Bucuresti), M. Opheim (Oslo), H. van Oorschot (Amsterdam) (HO), R.

van Oorschot (Amsterdam), Rijksmuseum van Natuurlijke Historie (Leiden) (ML, RJ),

Mrs. M. A. da Silva Cruz (Candal-Gaya, Portugal), H. Stammeshaus (Amsterdam)

(HS), J. Thurner (Klagenfurt, Austria), B. C. S. Warren (Folkestone, England),

J. Wolschrijn (Muiderberg, Netherlands), C. W. Wyatt (Farnham, England) (CW).

I wish to take the opportunity to express my gratitude to Prof. Dr. K. H. Voous for

his interest, stimulating suggestions and highly appreciated criticism. I am also much

indebted to Dr. A. Diakonoff for valuable help in various ways, and to Dr. Ir. L. P. S.

van der Geest for his aid in correcting the English text.

There is one person who has helped me more than I can realize at the present or

than she will ever admit: I am profoundly grateful to my wife.

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

1.g. Methods, measurements and abbreviations

(1). There are various methods of measuring the size of a butterfly, from such a

vague term as “wing span” to the exact but rather odd method of Warren (1926) of

measuring the length from the centre of the thorax to the tip of the wing and multi-

plying it by two. I have applied a simple but exact method by measuring the length

of the fore wing from base to apex (without fringes). Thus all measurements given in

this paper refer to the length of the fore wing.

(2). To avoid confusion between specific and subspecific names, a species (and

semispecies) is indicated by the combination Pyrgus a-us or P. a-us, a subspecies by the

combination 4-5 b-us or ssp. b-us, whatever may be appropriate. Names referring to the

infrasubspecific variation can be recognized by the word “forma” or the abbreviation

“f.” preceding the name of the variety.

(3). In the drawings of the male genitalia the aedeagus and the left valve have been

omitted or only the right valve has been pictured.

The drawings of the female genitalia only show the ventral side of the eighth

abdominal segment.

(4). Much care has been given to the drawing of the distribution maps; their scale,

however, does not permit the reproduction of small disjunctions. The maps serve to

facilitate a rapid recognition of a particular distributional area and a rapid comparison of

different distributional areas.

Although as many as possible faunal lists have been examined (the most important

ones have been listed in Chapter 7), it must be stressed that the distribution limits,

especially in Asia, are often very poorly known.

Abbreviations used

upf = upperside of fore wing

f. = forma uph = upperside of hind wing

ssp. = subspecies unf = underside of fore wing

unh

underside of hind wing

2. THE GENUS Pyrgus HUBNER, 1819

2.a. Delimitation

The delimitation of the genus Pyrgus accepted in this study is that proposed by

Warren (1926) (under the name Hesperia). This delimitation has been generally

accepted, but in recent times some authors (e.g. Alberti, 1969), have used the name

Hesperia again in the sense of most authors prior to Warren, i.e., to include also the

species which have been classified by Evans (1949) in the genera Spialia and Muscham-

pia (recte Syrichtus, see Hemming, 1967). Therefore, it may be useful to summarize the

distinctive characters of Pyrgus and to call attention to a few, hitherto overlooked.

The type species of Hesperia Fabricius, 1793, H. comma Linnaeus, and the type species

of Pyrgus Hübner, 1819, P. malvae Linnaeus, are usually regarded as non-congeneric

(they are classified in different subfamilies). Therefore, the use of the name Hesperia

instead of Pyrgus, should be avoided (see Hemming, 1934a).

R. DE JoNG: Pyrgus in the Palaearctic region 9

The characters of Pyrgus mentioned below refer to Palaearctic species only. Most

(and perhaps all) American Pyrgus species share these characters, but for the delimi-

tation in America (only interesting with regard to the genus Heliopetes) I may refer to

Evans (1935). Pyrgus may be characterized as follows.

Fore wing median spots in spaces 4 and 5 completely detached from the median

spots in spaces 6 to 8, nearer the termen; submarginal spots absent or faint and in-

complete.

Male hind tibiae with a hair tuft fitting into a double thoracic pouch.

Male fore wing with a costal fold, enclosing specialized, androconial scales (absent

in superspecies alpinus). The fact that the androconial scales offer a distinctive character

seems to have been overlooked. These scales are more or less fusiform, very narrow,

hair-like at the apex (Fig. 4-6). The variation in width (6-9 y) and in length (180-300

p) does not seem to be of great specific value, as there exists much overlap in the

100 u

4 5 6 ti 8 9 10 11 12

Fig. 4—12. Androconial scales in Pyrgus, Carcharodus and Syrichtus. 4, Pyrgus sidae (Turkey);

5, P. andromedae (Abisko, Sweden); 6, P. alveus (Bavaria); 7, Carcharodus alceae (Tarragona,

Spain); 8, C. boeticus (C Spain); 9, C. floccifer (Italian Alps); 10, Syrichtus proto (Marash,

Turkey); 11, S. tessellum (Aksu, C Asia); 12, S. cribrellum (N Tibet)

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

various species. P. sidae stands somewhat apart by its very long androconial scales:

280-300 y, in the other species the length varies mainly between 180-220 y.

Male genitalia with well-defined, strongly sclerotized, highly specialized lateral

apophyses. In the Palaearctic species the distal end of the harpe is differentiated into a

style, a stylifer and an antistyle.

Female genitalia have not been used in the search for distinctive generic characters in

Hesperiidae. I wish to call attention to a character which does not concern the genitalia

proper, but the general sclerotization of the eighth abdominal segment. In Pyrgus this

sclerotization consists of a single part of variable shape at both sides of the segment.

The left-hand and right-hand sclerites are connected ventrally and dorsally by a mem-

brane; in some species the sclerites are connected ventrally, by a narrow sclerotized

band.

By these characters Pyrgus stands taxonomically somewhat apart. The genera supposed

to be most closely related to Pyrgus, viz. Carcharodus, Syrichtus, and Spialia (cf. Evans,

1949, 1953), have more characters in common with each other (particularly Carcharo-

dus and Syrichtus) than with Pyrgus. The fact that there is some disagreement about the

classification of Carcharodus and Syrichtus, does not need to bother us here. On the

fore wing the median spots in spaces 4 and 5 (if present) are contiguous to the median

spots in spaces 6 to 8 in Syrichtus and Spialia. The median spots in spaces 4 and 5 are

absent in Carcharodus, but very rarely a minute spot is visible in space 5; this spot is

contiguous to the median spots in spaces 6 to 8.

A hair pencil on the hind tibia of the males is wanting in Carcharodus, Syrichtus and

Spialia. A costal fold is present in the males of Carcharodus and Syrichtus (except S.

antonia) and of a few Ethiopean species of Spialia.

In Carcharodus and Syrichtus the costal fold encloses androconial scales which are

quite different from the type found in Pyrgws: they are constricted at varying intervals

(Fig. 7-12). I did not examine the androconial scales in Sp/alia.

In the male genitalia the lateral apophyses are absent or rather well-developed (in

some species of Syrichtus and Spialia); they are less specialized than in Pyrgus. The

distal end of the harpe is very variable, but never with a style.

The sclerotization of the eighth abdominal segment in the females usually consists of

two sclerites on both sides, the four parts connected by a membrane. In some Spialia

species (e.g. S. sertorius) there is a single sclerite on both sides, dorsally connected by

a membrane, ventrally by a narrow band of continuous sclerotization.

In conclusion, it appears highly undesirable to consider the species of the genera

Syrichtus, Spialia and Pyrgus congeneric.

2.b. Classification

One of the characters of the genus Pyrgus is the strong sclerotization of the lateral

apophyses. Their specialized nature makes these structures very useful for the classification

of the genus.

The classification of the genus by Warren (1926) is mainly based on the develop-

ment of the lateral apophyses:

Subgenus Teleomorpha — lateral apophyses ventrally broadly connected.

R. DE Jong: Pyrgus in the Palaearctic region 11

19 Imm

Fig. 13—20. Male genitalia, 13, Pyrgus maculatus (Amdo); 14, P. bieti (Ta Tsien Lou); 25;

P. oberthuri (Ta Tsien Lou); 16, P. dejeani (Nan-Shan); 17, P. alpinus (C Asia); 18, P. carthami

(Digne, SE France); 19, P. sidae (Aksehir, Turkey); 20, P. badachschanus (Panjshir V., Hindu-

Kush)

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Subgenus Hemiteleomorpha — lateral apophyses with a narrow ventral connection.

Subgenus Ateleomorpha — lateral apophyses ventrally not connected.

The name Teleomorpha is a junior objective synonym of Scelotrix Rambur, 1857,

and the name Hemiteleomorpha is a junior objective synonym of Pyrgus Hübner, 1819;

see Hemming (1967).

This classification has generally been accepted, but Guillaumin (1966) stated that the

structure of the female genitalia does not support (nor contradict) this concept (he

only dealt with the species of France). This statement is only partly true and apart

from this I do not see why it should devaluate the classification. There is no obvious

reason to assume that the male and female genitalia evolved at the same speed.

Because of the large number of species it is practical to divide the genus into sub-

genera. However, any biological meaning disappears when the division is artificial, in

other words: when the subgenera are not monophyletic. Of the three subgenera proposed

by Warren, Scelotrix and Pyrgus can be considered monophyletic. The latter comprises

only one superspecies (malvae); at least two of the four species recognized by Warren

in this subgenus interbreed to some extent (malvae and malvoides). In the subgenus

Scelotrix the presence in all species of the ventral connection of the lateral apophyses

and of the structure of the genital plate can only be attributed to an evolution from a

common ancestral situation.

The common feature of the species of Warren’s subgenus Ateleomorpha is the absence

of a ventral connection of the lateral apophyses. The absence of a structure is much more

difficult to interpret phylogenetically than its presence. Besides, there is much variation

in the shape of the lateral apophyses and this variation runs to some extent parallel to

that found in the female genitalia, in the sense that in the species that have most similar

male genitalia, also the female genitalia are most alike. Although it is not impossible

that the species united by Warren into the subgenus Ateleomorpha are monophyletic,

I believe the conception of monophyly in this case to be less well-founded than in the

subgenus Scelotrix. Therefore, I prefer to restrict the subgeneric distinction to the

species of the alvews and cinarae groups of Warren, thus excluding P. serratulae and

P. onopordi and the superspecies carlinae (comprising the species carlinae and fritillum

= cirsii, recognized by Warren) which I prefer to consider species of uncertain affinity

(in relation to each other and to other species). However, the type species of Ateleo-

morpha is P. onopordi (Hemming, 1934); therefore this name cannot be used as a

subgeneric name for the a/vews and cinarae groups. As I have no intention of proposing

new names in this paper I am referring to the a/veus and cinarae groups as “the

alveus species complex’.

Subgenus Scelotrix

Male genitalia. — Lateral apophyses connected ventrally by a broad, spined, sclerotized

plate. Sichel (1962) named the narrow plate in the subgenus Pyrgus, lamina transversa;

this name can also very well be used in the subgenus Scelotrix. Uncus short, more or less

curved, usually with a blunt apex.

Female genitalia — An obvious genital plate, free (not embedded in a membrane),

variable in size, usually heart-shaped and always with an incision at the top. In the

sclerotized or membranous sternum of the eighth abdominal segment there is a corrugant

sclerotization in most species.

Facies — Median spot 5 unh usually projecting towards the base of the wing. In

R. DE JONG: Pyrgus in the Palaearctic region 13

some species a pale spot at the base of space 2 upf, always absent in members of the

other Palaearctic subgenera, but present in all American species (see below).

For the division into species I have followed Evans (1949), except in the case of

alpinus which was given specific rank by that author, but which I consider a super-

species (for reasons, see Chapter 3).

Some species resemble each other so closely with regard of the male and sometimes

also of the female genitalia, that we may assume a close relationship. This is the case in

the following species.

P. maculatus and P. bieti resemble each other by the shape of the valvae, the in-

distinct subharpal plate, the formation of style and antistyle, the uncus, the lateral

apophyses and the aedeagus (Fig. 13 and 14). It should be noted that the unh of

P. maculatus deviates from all known species of Pyrgus.

P. oberthuri and P. dejeani resemble each other particularly by the strong dorsal

development of the lateral apophyses, the uncus, the aedeagus and the general formation

of the valvae (Fig. 15 and 16). Externally they are also very similar, but so is the

case with many Pyrgus species.

P. alpinus, P. cashmirensis and P. darwazicus. The male genitalia are almost alike,

although the species differ very obviously in pattern (see Chapter 3 and Fig. 17, 58-60).

A further common feature is the absence of a costal fold in the males, a unique

character in the Palaearctic Pyrgus species. I unite these species in a superspecies.

P. andromedae, P. cacaliae, and P. centaureae. The male genitalia differ mainly in the

shape of the cuiller and of the antistyle (Fig. 51-56). The female genitalia chiefly differ

in the corrugant sclerotization of the sternum of the eighth abdominal segment, but, as

is shown in Fig. 21-29, the difference between these species is less than that from

other species.

The other species of the subgenus (P. carthami, P. sidae and P. badachschanus) seem

to stand more apart, or at least, their relationship to other species (and to each other)

is rather obscure.

The little developed cuiller and style-antistyle of P. carthami (Fig. 18) suggest a

relationship with the Chinese species (P. maculatus, P. bieti, P. oberthuri, P. dejeani).

The development of the sternum of the eighth abdominal segment in the female

(Fig. 27) supports this, although the genital plate is larger. Externally P. carthami is

not at all related to Chinese species, but more related to the European species; however,

most Pyrgus species look very much alike.

Warren (1926) thought P. sidae closely related to the species P. centaureae and

P. alpinus, but he only examined the male genitalia (Fig. 19) and even then the

relationship is debatable; as to the female genitalia (Fig. 28), P. sidae does not closely

resemble any other species and the same applies to the colour and design unh (alter-

nating white and orange bands).

Special mention must be made of P. badachschanus. This species, confined to N

Afghanistan, does not resemble any other Scelotrix species with regard to the male

genitalia (Fig. 20). The extremely dense hairing of the uncus is a unique character

and the very large cuiller only recalls the structure found in the a/vews species complex.

As far as I know, the female genitalia have remained unknown, but I could examine a

female from the captures made by Wyatt & Omoto (1966) (Fig. 29). Although clearly

of the Scelotrix type, it lacks the corrugant sclerotization of the sternum of the eighth

abdominal segment.

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

1 mm

Fig. 21—29. Female genitalia. 21, Pyrgus maculatus (Amur); 22, P. alpinus (Fort Naryn, C Asia);

23, P. andromedae (Abisko, Sweden); 24, P. cacaliae (Swiss Alps); 25—26, P. centaureae (Lap-

land); 27, P. carthami (Pyrenees); 28, P. sidae (Aksehir, Turkey); 29, P. badachschanus (Panjshir

V., Hindu-Kush)

Subgenus Pyrgus

Male genitalia. — Lateral apophyses ventrally connected by a narrow, either spined or

unspined lamina transversa. The main difference with the subgenus Scelotrix, apart

from the narrowness of the lamina transversa, is the development of small to very large

processes of the lateral apophyses. Uncus short, entire, incised or longitudinally split

into two parts, the strongly sclerotized apex rather blunt.

Female genitalia. — Genital plate very large, entire or split into two parts. No cor-

rugant sclerotization.

Facies. — Above spots usually well-developed, even the submarginal spots uph; unh

spots reduced, median spot 5 usually more or less projecting towards the base of the

wing.

There are three types of genitalia (Fig. 30-35).

The malvae type. — Uncus bipartite; lamina transversa spined; lateral apophyses with

long, strongly spined processes; cuiller pointed; style-antistyle short and broad. Genital

plate bipartite.

The melotis type. — Uncus entire or partly bipartite; lamina transversa smooth,

R. DE JoNG: Pyrgus in the Palaearctic region 15

laterally with a spine; lateral apophyses with long, unspined processes; cuiller rounded;

style-antistyle slender. Genital plate entire, but apically incised.

The malvoides type. — Uncus narrow, entire, usually apically more or less incised;

lamina transversa smooth; short spined processes comparable with the long processes in

the melotis and malvae types, but much less developed; valvae resembling the melotis

type. Genital plate entire, apically incised as the melotis type.

Presumably this subgenus comprises the most discussed forms of Hesperiidae. Warren

(1926) recognized four allopatric species in this subgenus: malvae, malvoides, melotis

and ponticus. Picard (1948-a, 1950-a) demonstrated the occurrence of hybridization

between malvae and malvoides in C. France. Hybridization was also reported from N

Tyrol (Kauffmann, 1955) and from Triest (Alberti, 1956). However, in Switzerland,

where malvae occurs in the north and malvoides in the south, hybridization has not been

reported (Kauffmann, 1951). The two forms are possibly geographically separated by a

very narrow zone (Wiltshire & de Bros, 1966).

The situation in C France has been re-examined by Guillaumin (1962), who found

a lower rate of hybridization than stated by Picard; like Picard he did not mention a

percentage of hybrid specimens, but at least the hybridization seems to occur more than

occasionally,

For the evaluation of the hybridization it is useful to mention the fact that malvae is

usually monovoltine and malvoides bivoltine.

In France the single generation of malvae flies from the end of April until the

beginning of June and the first generation of malvoides from the middle of May until

the middle of June (Guillaumin, 1964). Thus, only late specimens of malvae and early

specimens of malvoides have a chance to interbreed. Besides, in Switzerland malvae and

malvoides may differ in habitat preference, malvae being commoner in the valleys and

malvoides in the mountains (Wiltshire & de Bros, 1966).

The situation can be summarized as follows. The allopatric forms malvae and malvoi-

des interbreed where they come into contact. The proportion of hybrids depends on the

flight period and habitat preference of both forms and is, therefore, locally different.

The absence of transitional populations between malvae and malvoides has led to

various theoretical reflections (e.g. Pröse, 1955a; Sichel, 1962a; Niculescu, 1967).

Generally the two forms are considered to be separate species or at least semispecies of a

superspecies. Only Evans (1949) united them as subspecies (together with me/otis and

ponticus) into a single species. I fail to see why the choice of a term for a particular

situation would be so important, that much time and trouble is spent in order to show,

on merely theoretical grounds, that other authors have used a wrong term for this,

instead of recognizing the fact that the situation is much more important than attaching

a name to it. Therefore the remark of Guillaumin (1970, in litt.), that after a study of

many years of the situation in France, he cannot say whether malvae and malvoides are

species or subspecies, is much more important. It shows that in this case the decision

“species against subspecies’ should be justified by practice and cannot be evaluated

theoretically. As both malvae and malvoides can be subdivided into subspecies it seems

convenient to call them semispecies of a single superspecies. As hybridization seems to

occur more than occasionally, at least in France, I do not see a practical value in declaring

them species as this would obscure the situation.

The melotis type of genitalia is found in the externally different forms melotis and

ponticus. Unlike Sichel (1962) I could not find constant differences of the male

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Fig. 30—32. Male genitalia. 30, Pyrgus malvae malvae (Netherlands); 31, P. melotis ponticus

(Aksehir, Turkey); 32, P. malvoides malvoides (Pyrenees). Fig. 33—35. Female genitalia. 33, P.

malvae malvae (Netherlands); 34, P. melotis ponticus (Aksehir, Turkey); 35, P. malvoides mal-

voides (Pyrenees)

genitalia in melotis as against those in ponticus and as they also intergrade externally

more or less I do not see any reason for considering them specifically distinct (see

further Chapter 3). The zone of contact between malvae and ponticus is presumably

situated in the west of Turkey, but at least as far to the west as Aksehir, Egridir and

Antalya, all specimens, of which I could examine the genitalia, are ponticus without

any trace of malvae.

The fact that transitional specimens between malvae and ponticus are unknown does

not mean that these do not occur, as the western part of Turkey has not been carefully

investigated. As compared with the situation malvae - malvoides, we can only state that

ponticus and malvae potentially interbreed and therefore I place melotis (with the

subspecies melotis and ponticus) as a semispecies in the superspecies malvae.

The alveus complex

Male genitalia. — Lateral apophyses not connected by a lamina transversa, only

connected by a membrane. The ventral part of the lateral apophyses is strongly

sclerotized, more or less triangular and with a convex, straight or concave, strongly

spined dorso-distal edge. Uncus slender, with a hooked apex (that may be split lon-

gitudinally). Cuiller well-developed, at least about a semi-circle (P. armoricanus). Valvae,

Fig. 36—45.

Female genitalia. — Without a distinct genital plate, at most two sclerotized, post-

vaginal spots.

R. DE JONG: Pyrgus in the Palaearctic region 17

Facies. — Variable. Median spot 5 unh usually without a projection towards the

base of the wing.

There is much confusion in literature about the taxonomy of this species complex.

The following forms have been given specific rank on a certain day (arranged alphabe-

tically) : accretus, alvens, armoricanus, bellieri, cinarae, iliensis, jupei, kansuensis, numidus,

reverdini, scandinavicus, schansiensis, sifanicus, speyeri, and warrenensis. It seems

desirable to review all these forms critically in order to arrive at an acceptable taxonomic

division.

No problems have been caused by the forms armoricanus, bellieri, and cinarae. They

have always been regarded as separate species, at least since the work of Reverdin at the

beginning of this century.

P. cinarae deviates from the other types with regard to the male genitalia by a very

long uncus, and a broad, flat style which is folded over, a character only found in a

number of species of the subgenus Scelotrix. Also stylifer and antistyle are broad. The

female genitalia are characterized by the undeveloped postvaginal region.

The regular, semicircular type of cuiller found in P. armoricanus cannot be mistaken.

The ventral part of the lateral apophyses has a convex dorso-distal edge, distinct from

the usually concave edge in the other forms of the subgenus. The female genitalia lack

a distinct antevaginal plate which is present in the other forms. Very rarely P. ar-

moricanus and P. alveus fly together. This may be the case in the southern

Alps where Kauffmann & de Bros (1952) found a few specimens of P. armoricanus

with some features of P. alveus in the shape of the cuiller and of the lateral apophyses.

It is not certain whether these are hybrids, but if so, they must be the results of very

incidental cases of hybridization that still demonstrate the close relationship between

P. armoricanus and P. alveus but that do not devaluate the specific rank of P. armori-

canus.

In P. bellieri we find the most developed type of cuiller, amply reaching more than

half of the total length of the valve, with a large ventro-distal bulge.

The ventral part of the lateral apophyses is short, with a more or less concave

dorso-distal edge. In the female genitalia there are three postvaginal lobes, instead of

two, as in the other forms (except in P. cinarae which lacks postvaginal lobes).

There is only one form which closely approaches P. bellieri, viz. accretus. They come

into contact in the SW Alps, both at the French and the Italian side. Occasionally

specimens have been found with transitional characters. Warren sent me a photograph

of such a specimen from Cesane (Piedmont) and I saw two specimens from the

Basses Alpes. This suggests that (very rarely) hybridization occurs.

The taxonomic position of warrenensis is still rather uncertain, but I am inclined

to assign to it specific rank and, I shall deal with it below, as a species. In the original

description by Verity it was thought to be an extreme variation of P. alveus, confined

to the high altitudes of the Alps. Warren (1953), however, showed that it is quite

distinct from P. alvews, externally as well as in the male genitalia and that it may fly

with P. alveus, evidently without producing transitional forms. He even states that in

the localities in which warrenensis and alveus occur together the former emerges at least

ten days before alvezs, and is quite worn when the latter is fresh. Some specimens from

the Gross Glockner region, however, are difficult to identify and I think hybridization

occasionally occurs there. Besides, if we consider the matter from other side, the

statement of Warren that warrenensis emerges earlier than alveus, doesn't settle the

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

matter: warrenensis is adapted to a high altitude environment and therefore emerges as

early as alveus in spite of this higher altitude; in the course of the flight period the

weather becomes warmer and alveus ascends into the regions of warrenensis, but only

individuals of alveus that have emerged late in the season may reach the warrenensis

habitat. Of course, there is no proof of this, but it shows that the position of warrenensis

is still uncertain and that the interrelation between warrenensis and alveus can only be

made clear by a study on the spot.

Warren (l.c.) placed warrenensis as a subspecies in the species s/fanicus (see below),

an incomprehensible action, if one does not take into account the fact that Warren

misjudged the position of the latter form.

The form sifanicus has been surrounded by confusion since Reverdin (1915)

identified alveus-like specimens from the Caucasus and Transcaucasia as szfanicus which

originally was described from the Kuku-Nor region (N Tibet) by Groum-Grshimailo

(1891). Only in 1967 Alberti explained the situation in the Caucasus and Transcaucasia

by showing the occurrence of three forms in that region: one form like the N Tibetan

sifanicus, another form indistinguishable from the C European alveus and a third, that

he named jupei. Warren (1968, in litt.) is of the opinion that jupe? is only an extreme

variation of a variable s/fanicus. It is interesting to note that the photograph of the

male genital armature of ‘‘s/fanicus’ from Borshom in Reverdin (1915, pl. 6 fig. 6)

and evidently of the same microscopical slide, in Warren (1926, pl. 9 fig. 6) shows

the typical structure of jupei, while, as far as I know, the male genital armature of

sifanicus-like specimens from Transcaucasia has been figured by Alberti only (1967, pl.

1, figs. 1-c, 2a-c). However, the warrenensis type of genitalia resembles the jupe? type

and not the s/fanicus type, especially with respect to the raised apex of the cuiller

(compare Fig. 44 and 45). Furthermore, jupei is characterized by the very small style-

stylifer-antistyle, the height of antistyle + stylifer being about half of the total depth

from the base of the style to the ventral edge of the valve. Also in warrenensis it may be

rather small, but it is larger than in jupe? and the style is longer.

Externally jupei and warrenensis are quite distinct jupe? being a larger and usually

better marked form. Although the similarity between jupe? and warrenensis may be due

to a common ancestor, I do not see any reason to unite them into a single species, as

Warren did with warrenensis and sifanicus. Whereas warrenensis is confined to the high

altitudes of the Alps, jupe? only occurs in the high altitudes of the Caucasus region.

Specimens transitional to alvezs are not known, but, as a matter of fact, the Caucasus

region is much less explored than the Alps. Like warrenensis, jupei will be treated here

as a separate species.

There is another form that has been regarded by Warren (1953) as conspecific with

sifanicus, viz. numidus. This form represents the alveus complex in NW Africa where

it flies in the mountains from 1700 to 2700 m. Indeed, the male genitalia of numidus

with the highly raised apex of the cuiller, are very suggestive of warrenensis. In size

and design numidus and warrenensis represent about the extremes of variation in the

alvens complex, numidus being one of the largest and most extensively spotted forms,

warrenensis the smallest and least spotted form. Of all forms of the alvezs complex

numidus can be distinguished by the anvil-shaped central spot, unh, being in this respect

very similar to the large N African Pyrgus onopordi.

As in the case of jupei I do not see any reason for considering warrenensis and

numidus conspecific. Moreover, a form occurs in the Sierra Nevada, externally similar

R. DE JONG: Pyrgus in the Palaearctic region 19

to numidus but with the male genitalia intermediate between »umidns and central-

hispaniae (belonging to the form accretus): it possesses the narrow cuiller of numidus,

but apically the cuiller bends over to the harpe as in centralhispaniae. Possibly this form

represents a hybrid population of accretus and numidus. Therefore, I regard numidus

and centralhispaniae (and for that matter accretws) conspecific.

The form accretus is characterized by the short free proximal edge of the cuiller which

slopes towards the harpe and even may nearly touch it. Besides that, the cuiller has a

ventro-distal expansion, not as marked, as in bellieri, but usually obvious. Another

character of accretzs is the short ventral part of the lateral apophyses and the long shaft

that connects it with the dorsal part. With some geographic variation this form is

distributed throughout Spain and France and penetrates a little into NW Italy. According

to Warren (1953) it has been found together with alveus and bellieri near St. Martin

Vesubie (Alpes Maritimes), together with bellzeri at Limone (Piedmont, Ligurian Alps)

and probably together with a/vews at Oulx (Piedmont, Cottian Alps). He also supposes

a sympatric occurrence with alvezs in the Swiss Jura. However, his figures of an alveus

type specimen and an accrefus specimen (Pl. 7, Fig. 3 and 4) taken flying together at St.

Martin Vésubie do not indicate any more than that both types occur at the same spot.

It does not prove that a/vews and accretus are unable to interbreed. Indeed, examining

the genitalia of eight males from St. Martin Vésubie, I found four specimens of the a/vews

and two specimens of the accretus type of genitalia and two specimens with intermediate

characters. I also saw some transitional specimens from Haute Savoie. It is interesting to

note that Kauffmann (1953) recorded the occurrence of accretus in SW Germany

where also alvezs flies, but where it is found later in the season. Evidently, alveus and

accretus interbreed and although the interbreeding doesn’t seem to have led to extensive

hybrid populations, I prefer to regard accretzs only subspecifically distinct from alvews

until a detailed study of both forms in the western and southern Alps has revealed

their interrelations.

As accretus can be subdivided into geographic forms I will distinguish a subspecies

group accretus (see further below, Chapter 3). Of course, this decision is as subjective,

as that of Warren (1953), who raised accretus to specific rank. Evans (1956) says

(p. 749): “I have failed to separate accretus as a subspecies from alveus’”’, an incompre-

hensible observation, as they are easily separable.

The form alveus is characterized by: (1) the free proximal edge of the cuiller is

about vertical to the ventral line of the valve, its length being about one third the total

depth of the proximal edge of the cuiller (in accretus less than one third); (2) the

apex of the cuiller is level with the dorsal ridge of the harpe or slightly above it (unlike

warrenensis, jupei and numidus); (3) the outline of the cuiller is nearly circular or

partly flattened, without a ventro-distal bulge; (4) the ventral part of the lateral

apophyses is larger than in accretus and consequently the shaft is shorter. With con-

siderable individual and slight geographic variation this form extends from Central

Europe to the Altai. Even some specimens from the Kuku-Nor region cannot be

distinguished from C European ones. The only real difference with s/fanicus is the

white border of the underside of the wings in the latter, while in a/veus such white

borders are very rare, particularly unh. Evans (1949) regarded all specimens from the

Caucasus to Kuku-Nor as s/fanicus but this is obviously wrong as the white borders of

the wings, below, only occur in specimens from the Kuku-Nor region and from Trans-

caucasia (and N Iran). From the Kuku-Nor region I examined nine males and one

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

u, Be

2 43

Fig. 36—45. Male genitalia (right valve). 36, Pyrgus cinarae (S Russia); 37, P. armoricanus (C

Italy); 38, P. bellieri (Digne, SE France); 39, P. alveus accretus (C Pyrenees); 40, P. alveus alveus

(Swiss Alps); 41, P. alveus scandinavicus (Beito, Norway); 42, P. alveus numidus (Morocco);

43, P. alveus speyeri (Kentei); 44, P. warrenensis (Gross Glockner); 45, P. jupei (Kasbegi, C

Caucasus)

R. DE JONG: Pyrgus in the Palaearctic region 21

female; only five males and the female showed obvious white borders, the other males

more closely resembled the C European a/veus. In the Caucasus region the s/fanicus

type seems to be mainly confined to Transcaucasia and even there many specimens can

hardly or not at all be separated from the C European alveus. The male genitalia do

not warrant a specific separation of sifanicus either, as was done by Warren (1953)

(see above, under warrenensis and jupei). I examined the genitalia of four males

from Kuku-Nor and four (more or less s/fanicus-like) males from Transcaucasia. They

show some variation (one male from Transcaucasia has a highly raised apex of the cuil-

ler but is otherwise quite different from jupe?) but the variation seems to fall within

the variation width of alveus.

Kauffmann (1954a) dealt with the variation (particularly in the male genital arma-

ture) of alveus in the southern Alps (Insubria); in a confused and confusing manner

he named these populations zliensis colurnus, thus considering them conspecific with the

form iliensis, described from the river Ili (N Turkestan). Undoubtedly this conspecificity

is right, but the specificity of zliensis is without any foundation; I can only regard it as

a subspecies of alveus. According to Warren (1953) zliensis belongs to the speyeri

group (see below) but I do not understand how he arrived at this conclusion.

In Scandinavia the alveus complex is represented by a form (scandinavicus)

characterized by the distad slant of the proximal edge of the cuiller, which often gives

the impression that the cuiller is bent backwards; the length of the free part of the

proximal edge of the cuiller is slightly more than in alvezs; the cuiller is dorsally

usually more developed than ventrally and the antistyle is usually broader than in alvens.

Warren (1953), examining 28 males from Scandinavia, found a single specimen

from Gjendestunga (Jotunheim), with genitalia of the alveus type. As also

the scandinavicus type was known to occur there, he promoted scandinavicus

to the specific rank. Opheim (1956), however, examined the male genitalia

of about one hundred specimens from Norway and a single one from Gotland

and observed a fairly continuous variation from the scandinavicus type to the

alveus type; consequently, he assigned to scandinavicus subspecific rank only. I could

examine the male genitalia of about 40 specimens from Scandinavia and can support

Opheim’s view.

It is interesting to note that the formation of the cuiller found in scandinavicus

resembles in some specimens the formation in the Chinese form reverdini. This has

led to very confusing remarks by Warren on the specificity of reverdini. Without know-

ing the scandinavicus type of genitalia, Warren (1952) dissected a male from Sylkyn-

jarvi (Finland) and found the genitalia similar to a photograph by Reverdin (the same

as Warren, 1926, Pl. 6, Fig. 2) of reverdini and as the specimen had been taken flying

with a “perfectly typical’ male of alveus (of which Warren did not dissect the genitalia),

he concluded that reverdini was specifically distinct from alvens. Besides, he compared the

habitus of the Finnish “reverdinz’’ with the figures 1216 and 1217 of Oberthür (1912)

(belonging to the original description) and as the Finnish specimen was somewhat

different, he erected for it a new subspecies scotti!

In 1953 he had still not seen more of the Chinese reverdini than the photograph

mentioned in his 1952 paper. But he had studied the genitalia of scandinavicus and of

the “perfectly typical’ male of alvews which had been taken flying with scotti, and which

turned out to have the genitalia of the scandinavicus type (i.e, according to Warren,

the reverdini type). The subsequent taxonomic change was drastic, scandinavicus be-

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

coming a species, reverdini a subspecies of scandinavicus and scotti a form of the

nominate subspecies (sic!).

Apart from the fact that such actions do not promote real systematics, the nature of

reverdini remained entirely obscure. Kauffmann (1952) even doubted its subspecific

value, though he never saw a specimen of it.

I have examined the male genitalia of four specimens of reverdini (apart from the

photograph in Warren, 1926). They show some variation and are as much suggestive of

alvens as of scandinavicus. This also means that they are difficult to distinguish from

the structure, found in s/fanicus of the Kuku-Nor region. In addition reverdini and

sifanicus share the white borders at the underside of the wings. There is, therefore,

no reason to give reverdini more than subspecific rank.

There are still three forms left to deal with, all three Asiatic ones. One of these,

kansuensis, described by Reverdin, is entirely obscure and I doubt whether anybody

except Reverdin ever saw this form. Its locality suggests a close relationship, if not

identity with sifanicus, but it is better to leave kansuvensis here out of consideration.

Another rare but somewhat better known form is schansiensis. Though only very

few specimens have been described, it seems to be rather constant over a large area from

Shan Si to north of the river Amur. With regard to external characters, it resembles

alveus, although it is much larger. As to the male genitalia, it differs from alveus by

the very long ventral, spined part of the lateral apophyses, a character I also found

developed to some extent in a number of speyeri specimens. The cuiller appears to be

better developed dorsally than ventrally. Apart from the fact that the ranges of schan-

siensis and speyeri seem to overlap to a certain extent, there is no reason to place

schansiensis otherwise than as a subspecies of alveus. The fact that schansiensis and

speyeri are very different, leads us to the assumption that both forms do not interbreed;

while considering schansiensis and alvens conspecific it is sensible to consider speyeri

specifically distinct.

The form speyer? may be the best known Asiatic form of the alveus species complex.

It is widely distributed in Transbaicalia and the Amur district. I examined the genitalia

of fifteen males and found very much variation. The proximal edge of the cuiller is

vertical to the ventral line of the valvae, as in alveus, or slopes backwards, as in

scandinavicus. The antistyle is broad and straight, as in scandinavicus, or slender and

hooked, as in alvews. The apex of the cuiller, usually very solid and spined, is level with

the dorsal edge of the harpe in some specimens, but in most it is raised high above that

edge. Usually the cuiller is dorsally better developed than ventrally, having a somewhat

scandinavicus-like appearance. The free proximal edge of the cuiller is more than one

third the total depth of the cuiller, as in scandinavicus, and in some specimens even

reaches half of the total depth of the cuiller. The ventral part of the lateral apophyses

is also variable, sometimes suggestive of the schansiensis type but usually shorter. Ex-

ternally speyer? is an unmistakable form, though there exists some variation (see Chap-

ter 3). However, west of the Lake Baikal matters become complicated. There it seems

impossible to make a sharp distinction between speyeri and alvens (and scandinavicus!).

Details will be given in Chapter 3. It makes me doubt the specific rank of speyeri and,

though it is not a very satisfactory solution, I am placing speyer7 as a subspecies of

alveus, and schansiensis as a separate species. However, only abundant material and

research on the spot can clarify the relations between the Asiatic forms of the alveus

complex.

R. DE JONG: Pyrgus in the Palaearctic region 23

Summarizing, in this study the following species have been recognized within the

alveus complex: alveus, armoricanus, bellieri, cinarae, jupei, schansiensis and warrenensis.

Other forms, by some authors considered specifically distinct, have been placed here as

subspecies of a/vews. In this review I have only given the general outline of the variation.

Many details may be found (at least of the European forms) in Warren (1953) and

Kauffmann (1954). I must, however, stress the fact that much of the confusion about

the alveus complex is due to a too detailed examination of a too limited material which

has led to attaching too much value to details. Of course, the present classification cannot

be regarded as definitive, and only is an attempt to summarize the relations within the

entire alveus complex.

Species of uncertain affinities

In some respects, the species classified here seem intermediate between the subgenus

Scelotrix and the alvezs species complex. The females possess a genital plate, but

it is entirely embedded in the membrane and not free, as in Sce/otrix. The male genitalia

(Fig. 46-49) resemble the type found in the a/veus species complex by the sharp,

hooked apex of the uncus and by the absence of a ventral connection between the

lateral apophyses. The valvae are as much Scelotrix-like as alveus-like. The lateral

apophyses are very characteristic. In P. serratulae the ventral part of the lateral apo-

physes is very large and strongly spined; although not connected ventrally,

the ventral parts are yet solidly connected by a strongly curved, sclerotized, smooth

/ )

1 mm

Fig. 46—49. Male genitalia. 46, Pyrgus serratulae (S France); 47, P. onopordi (S France); 48, P.

carlinae (Val d'Aosta); 49, P. cirsii (C Spain)

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

bridge, with a very concave ventral surface and embedded in the anal sheath. In P.

onopordi the ventral part of the lateral apophyses is bipartite, with a long ventral and a

short dorsal projection, both spined. In the superspecies carlinae the ventral parts of the

lateral apophyses are rather simply built, somewhat elongated and strongly indented.

They recall to a certain degree the type found in malvoides (subgenus Pyrgus), but

are better developed and lack the lamina transversa of that semispecies.

Other separating characters can be found in the aedeagus which is strikingly broad in

P. onopordi and spined in the superspecies carlinae, a unique feature in the Palaearctic

species of Pyrgus.

As these species appear to be closer to Scelotrix than to the alvezs species complex,

they will be dealt with in Chapter 3 before the a/vews species complex.

The American Pyrgus species

Apart from centaureae which undoubtedly is a rather recent invader from Asia (see

Chapters 2d and 3) the American Pyrgus species are quite different from their Palaearc-

tic congeners. From a phylogenetic point of view, however, the following notes are of

importance.

(1). All American species have a white spot at the base of space 2 upf, as in a number

of species of the subgenus Scelotrix.

(2). Though variable the males of the American species have a lamina transversa or

something resembling it. In P. oileus I even found a lamina transversa which is dif-

ficult to distinguish from the type found in subgenus Scelotrix and in a form of

P. communis (orcynoides) the (unspined) lamina transversa appeared to be ventrally

concave, thus suggesting the “bridge” of P. serratulae.

(3). In the American species the uncus is bipartite, but unlike P. malvae the two parts

are situated close together (more suggesting malvae kauffmanni, see Chapter 3).

In conclusion, the American species appear distantly related to the subgenus Scelotrix,

while their relationship to the alveus species complex is only evident through the

subgenera Pyrgus and Scelotrix.

2.c. Distribution

The distribution of the genus Pyrgus in the Palaearctic is very extensive, it comprises

the entire Palaearctic, with the exception of desert regions. The distribution in N and

NE Siberia is unknown.

Of the eleven species of the subgenus Sce/otrix (including one superspecies) six are

confined to Asia, one is Holarctic, two are cold-adapted European species and two have

a mainly European distribution.

The subgenus Pyrgus has a Palaearctic distribution.

The remaining ten species (including one superspecies) in the Palaearctic are mainly

European, only P. serratulae and the alvews complex extending far into Asia, thus giving

a distribution opposite to that of the subgenus Scelotrix.

In America the genus is distributed from Canada to Chile and Patagonia. Of the

twelve species recognized by Evans (1953) six species are confined to S America.

There is still little agreement about the classification of the American Pyrgus species,

but a different classification can only increase the number of S American species.

Apart from P. centaureae only two species (in the classification of Evans), P. ruralis

and P. communis, extend to north of the Sonoran region.

R. DE JONG: Pyrgus in the Palaearctic region 25

2.d. Geographic and evolutionary history

The geographic origin of the genus is very uncertain. It would fall too far outside

the scope of this work, to elaborate on the systematics of the S American species, but

they are undoubtedly descendants of N American invaders. A Neotropic origin of the

genus can therefore be left out of consideration. The following statements can be made.

(1). The genus is of Holarctic origin.

(2). The contact between the Palaearctic and Nearctic groups was severed very long

ago, at some time during the Tertiary. The recent distribution of the species suggests an

originally thermophilous group. Whether the contact was re-established later is difficult

to say, but during the Quarternary contact was only possible through cold-adapted

species. Indeed, centanreae migrated into America in rather recent times (see also

below).

(3). Apparently, in the Palaearctic, the subgenus Scelotrix has the most primitive

characters as it has some characters in common with the American species, which among

all others are the only ones to be considered primitive.

(4). There are no indications of an invasion from the Nearctic into the Palaearctic.

(5). The bipartite uncus of malvae appears to be a young character and is not an in-

dication of a close relationship to the American species. All other characters, of the

male as well as of the female genitalia of malvae, indicate a recent origin, compared to

malvoides with which malvae still hybridizes to some extent.

In the Palaearctic members of the genus, the uniformity in the male and female

genitalia of the subgenus Scelotrix is striking, at least, in comparison with the other

species of the genus. This uniformity concerns the genital plate of the female and the

dorsal parts (uncus and lateral apophyses) of the male genitalia. The meaning of this

(relative) uniformity is not yet known, but apparently it is not because Scelotrix has

remained unchanged from the time that the contact with the Nearctic disappeared (see

also the history of Scelotrix, below). Apparently, the breakdown of the Sce/otrix structure

of genitalia has accelerated the evolution, subsequently giving rise to many divergent

types. This occurred presumably more than once. Therefore I cannot unite e.g. P. ser-

ratulae and P. alveus into one subgenus and fail to see, why P. carlinae would be more

closely related to, e.g. P. alveus than to P. malvoides.

The principal development of the genital plate and the lamina transversa of the

lateral apophyses appears to have been a reduction and subsequent disappearance.

Special attention may be drawn to the supposed evolution of the lateral apophyses.

In Scelotrix they form a complete ring around the anal sheath, to which they are more

or less attached. The lamina transversa supports the anal sheath ventrally and e.g. in

P. maculatus it has a long sclerotized projection towards the distal end of the anal

sheath. Presumably, the spined lamina transversa serves as an extra grip during

copulation. Where it is reduced its supposed function is taken over by spines springing

from the ventral part of the lateral apophyses. I have tried to illustrate a possible

course of the evolution in Fig. 50. It is impossible to believe that the malvoides and

the serratulae types belong to the same evolutionary trend. The other types (carlinae,

onopordi, alveus), if not evolved directly from a Scelotrix type, may have originated

equally well from the serratzlae type as from the malvoides type.

In addition an attempt is made to reconstruct the outline of the tentative geographic

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

AES

malvoides -type malvae-type

en Ee serratulae-t

- type

carlınae -type

onopor di - type

alveus - type

Fig. 50. Possible evolution of lateral apophyses in Pyrgus

history of the genus in the Palaearctic. The distribution, variation and relationship in Asia

of P. malvae, P. serratulae and the alveus complex strongly suggest that they are new-

comers in that continent and invaders from Europe. On the other hand, the same can

be said of the centaureae group (centaureae, cacaliae, andromedae) of Scelotrix in Europe,

while the other Sce/otrix species in Europe (carthami, sidae) appear taxonomically more

distant from Asiatic species.

Thus, the geographic history of the genus may be outlined as follows. At some time

during the Tertiary period Sce/otrix extended throughout the Palaearctic. The geographic

origin of Scelotrix cannot be defined more exactly. Somehow, the European part of the

genus became isolated and has developed characters of its own. This separation of the

European group was followed by a new invasion of Scelotrix into Europe, a subsequent

isolation, etc. This occurred several, if not many times, giving rise to the serrafulae and

malvoides types of genitalia, respectively. P. carthami and P. sidae may have originated

in the same way. The last invasion of Scelotrix brought centaureae to Europe (see

below, the history of the subgenus Scelotrix).

It is impossible to date the successive invasions more or less exactly, apart from the

more recent invasions of the centaureae group. Therefore, the causes of isolation of the

European groups remain uncertain. The following considerations, however, may be of

interest.

(1). As stated above, the genus was thermophilous (at least adapted to a temperate

R. DE JONG: Pyrgus in the Palaearctic region 27

climate), when the contact with the Nearctic has been severed. Such a group had ample

possibilities of being distributed throughout the Palaearctic in Tertiary times (cf. Peter-

sen, 1954; Moreau, 1955).

(2). Though the subgenus Scelotrix may be supposed to have the most primitive

characters of the genus in the Palaearctic, only three species (P. maculatus, P. carthami

and P. sidae) appear to be adapted to a temperate climate, while the other species are

cold-adapted (see also the history of the subgenus Scelotrix, below).

(3). None of the cold-adapted Pyrgus species outside the subgenus Scelotrix can be

regarded as a direct descendant of a cold-adapted Sce/otrix species (see also below).

(4). From this follows that the adaptation to a cold habitat in Scelotrix and in the rest

of the genus is secondary. Consequently the adaptation to high altitudes of Asiatic

Scelotrix species may have contributed to the isolations of the European groups. The

gradual rise of the Asiatic mountains may have forced this adaptation (see below).

(5). Of course, also the cooling down of the Palaearctic in Pleistocene times may have

influenced the contact between the European and Asiatic members of the genus, but

apart from the centaureae group this influence is obscure.

Summarizing the geographic history of the genus, we may state that the subgenus

Scelotrix has a Palaearctic origin, while all other Palaearctic species have a European

origin (or originate from a European species).

Asiatic species invaded Europe many times, giving rise to several new species by

subsequent geographic isolation. Some European species re-invaded Asia and in the

alveus complex this may have given rise to species duplication in Asia (see below).

The geographic history of Scelotrix and of the alvews species complex will be outlined

below; that of the other species will be dealt with in Chapter 3 (including the geographic

history of the species of Scelotrix and the alveus species complex).

Geographic history of the subgenus Scelotrix

The general uplift of the Asiatic mountains lasted until rather recent times. Ac-

cording to Kostrowicki (1969) the Kunlun was even elevated during the Pleistocene by

1300-1500 m above its level of the first glaciation. This must have been of great

influence on the geographic history of the species of the subgenus as it may have

forced some species to adapt themselves to a much colder habitat. But apart from such a

general statement it is difficult to indicate the influence more exactly. This is partly due

to the lack of knowledge on the present distribution in Tibet. From the scanty data

available one may assume that e.g. P. oberthuri and P. dejeani are widely distributed

throughout Tibet, but as long as they are only known from border areas one cannot be

sure. Nevertheless, their known distribution (E Tibet, Turkestan/Transcaspia) sug-

gests that they belong to the fauna that has been forced to submit to an adaptation to

rough climatic conditions during the time of the uplift of the Tibetan highlands.

It is clear that the cold-adapted Asiatic species must be regarded as having originated

in Asia, while the warmth-loving P. carthami (European distribution) and P. sidae

(mainly Mediterranean distribution) may be regarded as remnants of the old, warmth-

loving Scelotrix group. Although its ancestors may have been Asiatic, P. carthami un-

doubtedly became separated from related forms in Europe. The geographic origin of

P. sidae may have been the western, warmer part of Central Asia and also the Mediter-

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

ranean. The E Asiatic P. maculatus (the only Pyrgzs species that has reached Japan) may

also be regarded as a remnant of (or a descendant from) the thermophilous Scelotrix

group. Like P. sidae it has developed a particular unh. Its origin, however, is obscured

by the presence of a subspecies in the mountains of Szechwan and by the presence of a

probably closely related species, P. bieti, in the mountains of Szechwan and Yunnan.

Turning again to the cold-adapted species, the history of P. badachschanus isolated

by its structural features and restricted to N Afghanistan is entirely obscure. The presence

of a white spot at the base of space 2 upf may indicate its high antiquity. The super-

species alpinus is known from the same region, but has a much wider distribution,

extending from Kashmir and Afghanistan through Pamir to N Turkestan and the Altai

as far as Kamschatka. As will be seen in Chapter 3, the distribution north and east of

Turkestan is supposed to be of recent age and the origin of the superspecies has to be

looked for in Pamir and adjacent regions.

According to Warren (1926), alpinus is closely related to the centanreae group (P.

centaureae, P. cacaliae and P. andromedae), but I am not able to understand, why their

relationship to the centaureae group would be closer than to other Scelotrix species.

The geographic history of the centaureae group may be outlined as a repeated ex-

pansion to Europe. Of the three species P. andromedae appears to be the least modern.

This is not only indicated by the presence of a white spot at the base of space 2 upf

(rarely present in P. centanreae, too), but also indirectly by the fact that some specimens

of centaureae sibiricus resemble P. cacaliae in the washed-out appearance of the wings.

Presumably three waves of migration from Siberia to Europe occurred, successively

bringing P. andromedae, P. cacaliae and P. centaureae to Europe.

In Chapter 3 the last invasion into Europe is supposed to be of postglacial date.

Thus the invasion from which the present P. cacaliae originated may be tentatively

dated at the Third Glacial, while the first invasion (P. andromedae) may have taken

place during, or shortly after, the Second Glacial. However, in Chapter 3 the expansion

of P. centaureae into America has been dated at the Third Glacial, i.e., the same

glacial during which P. cacaliae is supposed to have reached Europe. Externally, in the

male and in the female genitalia the Nearctic and the Palaearctic centaureae forms are

closer to each other than to P. cacaliae, suggesting that the contact between the Palaearctic

and Nearctic centaureae forms has been severed later than the contact between Siberian

P. centaureae (as far as we can speak of centaureae in that period) and European

P. cacaliae. This is quite possible, as P. cacaliae may have reached Europe at the

beginning of the Third Glacial and may soon have lost contact with Siberia.

According to the above reconstruction of the geographic history, P. cacaliae and

P. andromedae appear to be species that in Europe became specifically distinct from the

Siberian populations from which they originated, while centanreae received its specific

imprint in Siberia from where it spread to Europe and America.

At present the Scandinavian form of P. centaureae (ssp. centaureae) is distinct from

the Siberian form (ssp. s7biricus). In the male genitalia the ssp. centaureae appears to

differ less from the American forms than from ssp. szbiricus (the female genitalia have

been left out of consideration, because of their great variability). In some specimens of

ssp. sibiricus the cuiller is quite narrow and elongated which seems to be a recent

character. Thus it appears that if ssp. centaureae and ssp. sibiricus will eventually

become specifically distinct, it will be due to a differentiation of ssp. sbiricus and not

R. DE JONG: Pyrgus in the Palaearctic region 29

51 52

53 i 54

55 56

Imm

Fig. 5156. Male genitalia (right valve). 51, Pyrgus andromedae (Abisko, Sweden); 52, P. cacaliae

(Tirol); 53, P. centaureae centaureae (Lapland); 54, P. centaureae wyandot (New York); 55, P.

centaureae sibiricus (Altai); 56, P. centaureae sibiricus (E Sajan)

of ssp. centaureae. Thus both will still have to be called Siberian species, as both

originated there.

Virtually the male genitalia of the centaureae group are suggestive of such a develop-

ment. The general evolutionary trend of the valve appears to be a widening of the

gap between the distal edge of the harpe and the proximal edge of the cuiller, followed

by a narrowing of the cuiller. Thus, in P. andromedae the free proximal edge of the

cuiller practically touches the terminal curve of the harpe. In P. cacaliae the edge of the

cuiller is very close to the harpe but does nowhere touch it, while in centaureae cen-

taureae the harpe and the free proximal edge of the cuiller are clearly apart and in

some specimens of centaureae sibiricus they are far apart (Fig. 51—56). Apparently,

a repeated differentiation in the cenfaureae group took place in Siberia and not in

Europe. The denomination of P. cacaliae and P. andromedae as European species is

therefore, at least, debatable. Indeed, the farreaching intergradation of the Siberian and

European faunas is much more apparent in the denomination of P. cacaliae and P. an-

dromedae as Siberian species than as European species.

The fact that P. cacaliae and P. andromedae at present have retained some original

characters does not mean that they are identical to their parental Siberian populations.

Undoubtedly a part of the present characters evolved in Europe, but this does not alter

the fact that they were Siberian when they became specifically distinct from other forms.

If the Siberian populations had evolved at a slower speed and had retained more

of the original characters than the European ones, P. cacaliae and P. andromedae would

have been regarded as European species as they (apparently) would have received their

specific imprint in Europe.

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Phylogenetically the difference between a European and a Siberian origin in the

cold-adapted species appears gradual and not essential. The conception may be tenable

that a species ceases to exist when it is split into two or more intersterile forms, but of

course this is only a matter of definition (see also Chapter 4.a).

Unlike P. centaureae, P. cacaliae and P. andromedae do not belong to what is usually

meant by the denomination “Siberian fauna”, i.e. the fauna of the taiga (coniferous)

zone. They belong to the fauna of the alpine meadows. For their zoogeographic clas-

sification, see Chapter 5.

The geographic history of the a/vews species complex

As stated above the origin of the alveus species complex is obscure, but it may be

called a European group of species. In Asia (apart from the eastern Mediterranean)

it is only represented by alveus and schansiensis. This is a poor representation as

compared with the six species in Europe, the more so as the specific rank of schan-

siensis is debatable (it could be a subspecies of P. alveus). The slight differentiation

in Asia, the absence south of the Issyk-Kul (except P. cinarae?) and the fact that

the least modern species of the complex, P. cinarae and P. armoricanus, are European,

lead to the assumption that the alveus species complex has a European origin and that

it invaded Asia from Europe.

The supposition, that P. cinarae and P. armoricanus are the least modern species of the

complex, is based on the fact that they stand taxonomically somewhat apart, while all other

species are extremely close to a/vews; also because of their preference for a warm climate,

while the other species prefer a more temperate or even cold climate. The geographic

origin of P. cinarae and P. armoricanus cannot be defined more exactly than it is

European, but ecologically they do not belong to the same fauna, see Chapters 3 and 5.

The history of the other species can be reconstructed with a fair degree of probability.

The present distribution area of P. bellieri (SE France, NW Italy, C Italy) strongly

suggests an origin in an Italian refugium during some glaciation. In external characters

and in the male genitalia P. bellieri is an extreme variation of the SW European accretus

group (subspecies group of P. alveus) and undoubtedly they share a common origin. As

the occurrence of P. alveus in C Italy (ssp. centralitaliae) probably dates from the Last

Glacial (see Chapter 3), the origin of P. bellieri and the accretus group cannot have

been later than the Third Glacial and there is no reason to assume an earlier origin.

The origin of P. warrenensis (Alps), P. jupei (Caucasus) and alveus numidus (N

Africa) must have been earlier and may be dated at the Second Glacial. The fact that

they share some characters of the male genitalia (mainly the raised apex of the cuiller,

but also the more or less even width of the harpe) may be due to their common origin.

Though the resemblance appears rather slight and of little importance in view of the

variation in the a/veus complex, one cannot expect to find a total resemblance between

forms that have been isolated from each other since the Second Glacial. I think it even

possible that the E Asiatic alvens speyeri belongs to this Second Glacial group. The

variability in the male genitalia in this form could be due to later interbreeding with

other forms, at least partly. The origin of P. alveus is obscure; P. alveus itself may have

originated as well in Europe as in Asia, although the a/vews species complex as a whole

is of European origin. For P. alveus, see further Chapter 3.

R. DE JONG: Pyrgus in the Palaearctic region 31

The last species of the alveus species complex to be dealt with is P. schansiensis.

However, this form is very badly known and I give it only tentatively specific rank.

It seems advisable to leave the species out of consideration at this place.

3. THE SPECIES OF Pyrgus IN THE PALAEARCTIC

1. Pyrgus maculatus Bremer et Grey

Bremer et Grey, 1853 — Schmett. N. China's: 11, pl. 3 fig. 6.

Type-locality: Pekin.

Distribution (Map 1). — East Palaearctic. From Chita and Stanovoy Khrebet,

southward to S China (Yunnan, Kwang Si, Kwang Tung). The only species of the

Pyrgus group in Japan.

In the northern part of the range widespread in the Amur district, but in the north-

western part only known from Chita (Transbaicalia, 1 &, BM) and Mongolia (without

exact locality, 1 9, BM).

Habitat. — Although the species is common, the habitat has rarely been recorded.

According to Kurentsov (1949) in Primorsky Kraj (Maritime district) (N of Vladi-

vostok) along river-banks and roads. From lowlands up to 1550 m (Tai Shan, prov.

Shantung, China; BM), ssp. thibetanus to 2800 m (Ta Tsien Lou; BM), and perhaps,

much higher.

Biology. — Presumably bivoltine, as morphologically two broods can be distinguished

(see below), the first brood flying from March to May and the second from May to

July; possibly at high altitudes and in the north a single brood, at least summer

specimens from E Siberia are scarce. Kurentsov (1949) records the flight period north

of Vladivostok as May-June.

The larvae have been found by Graeser on Rubus idaeus and Spiraea (Rosaceae)

(Staudinger, 1892), on leaves which had been spun together.

Geographic variation. — The geographic variation of this species has long been

obscured by seasonal variation. Leech (1894) divided the forms maculatus, zona, albi-

strigus and thibetanus, here considered conspecific, into three species and placed

albistrigus as a variety of zona. All four forms were given specific rank by Warren

(1926). Evans (1949) was the first to recognize the seasonal variation, perhaps he

was also the first who could study an adequate number of specimens. I have seen all

the material Evans studied and can only follow his classification.

Pyrgus maculatus maculatus Bremer et Grey

Pl. 1 Fig. 1—3; Pl. 2 Fig. 25—27

Material examined. — 145 8 95 9: spring form; 1 & Chita (BM), 30 4 33 9

E Siberia (Amur, Sichote Alin, Stanovoy Khrebet, “E Siberia”) (BM, ML, RJ),

10 & 1 9 Askold (BM), 1 & 2 9 Manchuria (BM), 1 4 N Korea (Seishin) (ML),

Sgt © N’China, 1 gi 1 @ Pekin, 3.1 9 Shan Si, 1 g 1 9 S Shen Si, 4 ¢

4 9 Shantung, 9 g 4 2 Kuku-Nor (all BM), 20 9 15 2 Japan (BM, ML, ZMA,

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

RJ); summer form: 1 9 Mongolia, 1 & 3 9 E Siberia, 1 Q Manchuria, 8 g' 5 Q

Korea, 7 & 2 9 N China, 2 & Pekin, 1 g 1 2 Shan Si, 11 g' 5 2 Shantung (all

BM), 29 G 14 @ Japan (BM, RJ).

Distribution. — The northern part of the distribution area of the species; the southern

limit is not exactly known, somewhere between the Huang-ho and the Yang-tse-Kiang;

there may be a wide transitional zone with ssp. bockz.

Subspecific characters. — Unf apex beyond the sub-apical spots uniformly coloured.

Uph submarginal band of spots (only present in spring form) more or less angulated in

spaces 4—5. Male genitalia: style short and broad, not nearly reaching the end of the

cuiller; lamina transversa of lateral apophyses with short spines.

Two easily separable seasonal forms. Spring form: Above, with large spots, uph with

conspicuous discal and submarginal spots; unh ground colour usually chestnut, sub-

marginal spots forming a narrow, more or less continuous band, a very narrow white

spot mid space 7 (sometimes absent). $ 12.8—14 mm. The holotype of the species

belongs to this form. Summer form: Above, spots reduced, especially uph, where only

two (or rarely three) central spots are present; unh ground colour more greyish dark

brown, submarginal spots absent. This form was named zona by Mabille (1875) and

being, of course, sympatric with maculatus, has, for a long time, been considered a

separate species.

Variation. — There exists some variation in the extension of the spots above, and in

the ground colour and extension of the white submarginal band unh. Some specimens

from China can be separated only with difficulty from the following subspecies.

Pyrgus maculatus bocki Oberthiir

Pl. 1 Fig. 5—7; Pl. 2 Fig. 29—31

Oberthiir, 1912 — Et. Lép. Comp. 6: 71, figs. 1235, 1236.

Type-locality: Kiang Si.

Material examined. — 63 & 42 9: spring form: 3 g' 3 9 Leou Fang, 3 g' 3 2

Shanghai, 3 G 3 2 Chekiang, 6 g' 1 $ Szechwan (Chia Ting Fu), 8 g' 7 9 Hupeh,

1 & 2 2 Fukien, 3 4 Kiang Si, 1 g' Kwang Si; summer form: 5 g'° Leou Fang, 2 &

4 Q Shanghai, 4 $ 1 9 Chusan Island, 1 g 4 Q Ya Chiao Ling, 4 $ 1 Q Chekiang,

4 d 1 @ Szechwan (Chia Ting Fu), 7 g 4 9 Hupeh, 3 g' 4 Q Fukien, 2 ¢

Hunan, 2 g 4 9 Kiang Si, 1 G Kwang Tung (all BM).

Distribution. — The southern part of the distribution area of the species; for the

northern limit, see under subspecies maculatus.

Subspecific characters. — Very much like subspecies maculatus, but differing in the

spring form by a darker ground colour unh and by the absence of the submarginal band

and the small spot mid space 7 unh, and in the summer form by, usually four, discal

spots uph and the light greyish colour of the basal two-thirds of the hind wing below,

where the small spot mid space 7 may be present.

Variation. — Varies somewhat in the extension of the white spots above (particularly

uph) and in the colour unh. Some specimens of northern localities (Chekiang, Hupeh)

resemble ssp. maculatus.

R. DE JONG: Pyrgus in the Palaearctic region 33

Pyrgus maculatus thibetanus Oberthiir

PI. 1 Fig. 4, 8; Pl. 2 Fig. 28, 32

Oberthür, 1891 — Et. d’Entom. 15: 20; pl. 3 fig. 27.

Type-locality: Ta Tsien Lou.

Material examined. — 36 g 26 9: spring form: 25 g 9 ® Szechwan (Ta Tsien

Lou, Wa Ssu Kow, Maenia, Batang, Quase), 9 d 16 2 Yunnan (Li-Kiang, Tsekou,

Ta Pin Tze, Teng Yuch Ting): summer form: 2 & 1 9 Szechwan (Mosy mien, Ta

Tsien Lou) (all BM).

Distribution. — SW China.

Subspecific characters. — Unf apex beyond the sub-apical spots, inner half dark, outer

half white. Uph band of submarginal spots not elbowed. Male genitalia: style longer,

reaching end of cuiller; lamina transversa with long spine-like hairs on either side of

the aedeagus.

The difference between the spring form and the very few specimens known of the

summer form is slight. Above, both forms resemble the forms of ssp. Lock, below

they look like a pale summer form of ssp. maculatus, but the discal band unf is

usually wider and the termen unf is broadly pale.

Geographic history. — The apparent constancy of both ssp. maculatus and ssp. bocki

throughout their range clearly points to an origin of both subspecies in isolated areas,

situated in N China-Manchuria and S China, south of the Yang-tse-Kiang. The cause

of this separation is, however, not clear. A marine transgression could possibly inundate

a large part of eastern China and the lower Yang-tse-Kiang, and cause a separation in

this way.

Ssp. thibetanus may be older, as it differs externally as well as in the male genital

armature. Presumably it is a high altitude insect, but the data are very scanty. Its range

offers many possibilities for isolation by high peaks and deep valleys.

2. Pyrgus bieti Oberthür

Oberthür, 1886 — Et. d’Entom. 11: 26, pl. 6 fig. 50.

Type-locality: Ta Tsien Lou.

Distribution (Map 2). — Only known from the Chinese provinces Szechwan and

Yunnan. For exact localities, see below. The specimens recorded by Leech (1894) from

Amdo and Nan Shan probably belong to P. dejeani.

Habitat. — Not recorded. Undoubtedly an insect of high altitudes. The lowest altitude

probably is Tse-kou (Yunnan), at 1993 m (Oberthür, 1912).

Biology. — Unknown.

Geographic variation. — Although there seems to be some overlap in the characters,

there is a slight difference between the specimens from Yunnan and from Szechwan.

It is difficult to evaluate this difference, as most (76 %) of the specimens studied

originate from one locality, Ta Tsien Lou. However, the possibility of getting more

material for study appears rather remote at the moment. So we must feel satisfied for the

present with the division into two subspecies.

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pyrgus bieti bieti Oberthür

PI. 1 Fig. 9—10; Pl. 2 Fig. 33 —34

Material examined. — 83 g' 51 2:67 g 49 2 Ta Tsien Lou (BM, ML), 7 g Wa

Ssu Kow, 6 d 1 Q Yaragong, 1 g' Vrianotsang, 1 g' Batang, 1 9 Rama La (SE

Thibet), 1 8 Maenia (all BM).

Distribution. — W Szechwan.

Subspecific characters. — Unh ground colour greenish to ochreous, submarginal white

spots in spaces 1-c and 2 extend to termen, discal white band narrower than in ssp.

yunnanus.

Variation. — The ground colour unh may be as reddish as in ssp. yunnanus and

sometimes the submarginal spots in spaces Ic and 2 are separated from the termen by

traces of dark dots. There is also some variation in the width of the discal band unh

and in the extension of the spots above.

There is an obvious sexual dimorphism, males being much more distinctly marked

than females.

Pyrgus bieti yunnanus Oberthiir

PI. 1 Fig. 11—12; PI. 2 Fig. 3536

Oberthür, 1912 — Et. Lép. Comp. 6:66.

Type-locality: Yunnan.

Material examined. — 14 & 5 ® Yunnan (Li-kiang, Ta Pin Tze) (BM).

Distribution. — N Yunnan.

Subspecific characters. — Unh ground colour generally redder than in ssp. bieti,

submarginal spots in spaces 1-c and 2 separated from termen by dark spots, discal band

wider than in ssp. biet?

Variation. — The specimens examined varied in all characters, some specimens were

almost inseparable from ssp. biet.

Sexual dimorphism as in ssp. bieti.

Geographic history. — With the present data it is impossible to get a clear picture

of the history of the species. Evidently, the separation into subspecies is very young.

The extreme differences in altitude in the distributional area, caused by the steep and

deep river valleys, offer many possibilities for isolation. As P. bieti presumably is an

insect of high altitude, an amelioration rather than a worsening of the climate may

cause isolation. It is, therefore, not impossible that the differentiation observed is of

postglacial age.

3. Pyrgus oberthuri Leech

Leech, 1891 (June) — Entomologist 24, suppl.: 59.

Type-locality: Wa Ssu Kow.

Distribution (Map 3). — The Chinese provinces Szechwan and Yunnan. One male

is known from Fort Naryn (E Turkestan) (BM).

R. DE JoNG: Pyrgus in the Palaearctic region 35

Habitat. — Not recorded. Undoubtedly a species of high altitudes, as P. bieti. At

least to 4000 m (Wushi, Szechwan) (BM).

Biology. — Unknown.

Geographic variation. — Two subspecies can be distinguished which differ in the

male genital armature and the colour unh.

Pyrgus oberthuri oberthuri Leech

Pl. 1 Fig. 13—14; Pl. 2 Fig. 37—38

Material examined. — 27 & 9 9:16 g 3 Q Ta Tsien Lou, 9 4 6 9 Wa Ssu

Kow, 1 g' Batang, 1 & Fort Naryn (E Turkestan) (all BM).

Distribution. — SW China, E Turkestan.

Subspecific characters. — Unh ground colour dark greenish brown. Male genitalia:

apex of cuiller more rounded.

Note. — According to Evans (1949) smaller than ssp. delavayi, 8 13—14 mm, but

some males (Ta Tsien Lou) are as large as ssp. delavayi, 16 mm, and therefore, size

cannot be considered an important subspecific character.

Pyrgus oberthuri delavayi Oberthür

Oberthür, 1891 (July) — Et. d’Entom. 15: 20; pl. 3 fig. 31.

Type-locality: Yunnan.

Material examined. — 47 4 2 2:33 4 1 @ (Szechwan, Wushi 12000 ft), 14 3

1 2 Yunnan (Teng Yueh Ting, Ta Pin Tze, Li-kiang) (all BM).

Distribution. — SW China.

Subspecific characters. — Unh ground colour generally light greenish grey-brown

by white or creamish superscaling, except narrow dark brown lines along white spots.

Male genitalia: Apex of cuiller more pointed.

Note. — (1) The characters only refer to the males, as only two females are known.

Moreover, the female from Wushi has a reddish unh. (2) According to Picard

(1948b) there is a single geographic form, but he only refers to figures of Leech and

Oberthiir and evidently, he did not study much material.

Geographic history. — Presumably this species has a history parallel to that of P. bieti.

Although it seems to have a much wider distribution, it is nevertheless senseless to

speculate on the history on the base of the single male known from Turkestan.

4. Pyrgus dejeani Oberthür

Oberthür, 1912 — Et. Lép. Comp. 6: 66—67.

Type-locality: Ta Tsien Lou.

Distribution (Map 4). — Only known from Tibet, Sikkim and W Szechwan.

Furthermore, one male and one female are known from Tura (BM). According to

Evans (1949) this locality is in Transcaspian, but according to Warren (1936) it is in

the Namangan district in Ferghana. As far as I know, most specimens labeled “Tura”

came from Staudinger, who explained “Tura” (in Staudinger - Rebel, 1901) to be the

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Achal Tekke region in Transcaspian. Therefore I think that Evans is right. Apparently

it is a rare insect, but it may be widely distributed in Tibet.

Habitat. — Not recorded. In Sikkim at 4000 m (BM).

Biology. — Unknown.

Geographic variation. — There is some variation in size, g' from 12.4 to 14.7 mm.

The larger specimens are from Ta Tsien Lou. Other variations are unknown, probably

because of the scarcity of the material. (Pl. 1 Fig. 15—16; Pl. 2 Fig. 39—40).

Material examined. — 13 & 13 9:6 & 5 9 N Tibet (Amdo, Nan Shan), 2 ¢

2 2 SE Tibet (Charme, Gyantse, Pusho, Kyari Dzung), 1 9 E Tibet (Nenden),

3 d 3 2 W Szechwan (Ta Tsien Lou), 1 g 1 Q Sikkim (Chumbi Valley, 12.000 ft),

1g 1 2 Tura (all BM).

5. Pyrgus carthami Hiibner

Hübner, 1808—1813 — Samml. europ. Schmetterl. 1: pl. 143 fig. 720—723.

Type-locality: S Germany (selection by Alberti, 1938).

Nomenclature. — There is little uniformity in the use of the specific name of this species: both

the names carthami Hubner, 1809—1813 and fritillarius Poda, 1761, are in use. The difficulties

arise from the interpretation of the original description of fritillarius and from the interpretation of

the figure referred to by Poda. About carthami there has never been any difference of opinion, so I

prefer to use the name carthami, thus following Lempke (1953) and Alberti (1952-c, 1953-b).

Distribution (Map 5). — West Palaearctic: from Spain to E Russia and S Turkestan;

even known from Saisan. In Spain wide-spread, from Andalusia to the Pyrenees. Un-

doubtedly the species also occurs in Portugal, but I have not found any record of this;

it is not recorded by Zerkowitz (1946), nor by Da Silva Cruz (in litt.). Practically

throughout France and throughout C and S Europe. Not in Corsica and Sardinia (Verity,

1940), the northwestern part of Switzerland (Kauffmann, 1951) and the northwestern

part of peninsular Italy (Verity, 1940). No reliable records from the Peloponnesus.

The northern limit roughly runs from the mouth of the Loire through SE Belgium to

Stettin and Danzig (Friese, 1956). Once collected in the Netherlands (Lempke, 1953).

Farther east the most northern localities are Wolmar (Petersen, 1924), Podolsk (BM)

and Kasan (Krulikovsky, 1908). In Siberia only known from Saisan (1 &, BM).

In S Russia known from the Crimea (Korshunov, 1964) and Sarepta (Eversmann,

1844; ML, BM). Not in the Caucasus (Alberti, 1969). In Turkey evidently very rare,

recorded by Alberti (1938, 1969) from Konia and Taurus Mts. Farther east only

recorded from Askhabad (Alberti, 1938).

Habitat. — Flowery meadows, way-sides, mountain slopes, from lowlands to 2200 m

in the Alps (Verity, 1940; Kauffmann, 1951) and the Pyrenees (Caradja, 1893; Dufay,

1961); in the Sierra Nevada still common at 2500 m (Manley & Allcard, 1970).

Biology. — Monovoltine. In Switzerland flight period from the middle of May until

the end of July in the valleys, and from the beginning of June to the middle of August

in the mountains (Kauffmann, 1951). Records of two broods per year (e.g. Caradja,

1893; Manley & Allcard, 1970) are presumably biased by the long flight period.

Various plant species have been recorded as food plants; presumably some of them

are false, but this is difficult to prove. Most records undoubtedly do not rely on own

observations. I think the following summary is complete as far as the recorded food

plants are concerned.

R. DE JONG: Pyrgus in the Palaearctic region 37

Malvaceae: Althaea, A. officinalis, A. hirsuta (Verity, 1940; Schmidlin, 1948; Forster

& Wohlfart, 1955; Manley & Allcard, 1970; Higgins & Riley, 1970); Malva, M.

sylvestris (Verity, l.c.; Schmidlin, l.c.; Forster & Wohlfahrt, l.c.; Manley & Allcard,

l.c.); Carthamus lanatus and coeruleus (Guillaumin, 1964).

Rosaceae: Potentilla, P. opaca, P. verna (Bergmann, 1952; Friese, 1956; Higgins &

Riley, l.c.).

Compositae: Centaurea (Verity, l.c.; Schmidlin, l.c.; Manley & Allcard, L.c.).

Gramineae: (Schmidlin, l.c.; Guillaumin, 1.c.).

Geographic variation. — A highly variable species of which many forms have been

described as subspecies. As is mostly the case when subspecies are distinguished for the

sake of the distinction, the splitting-up has obscured the pattern of geographic variation,

rather than cleared the picture. Alberti (1938) was the only one who studied the

geographic variation throughout the range of the species. Although he evidently had too

little material available from Spain and France, his conclusions are very valuable, as he

looked for the relations between the subspecies. The treatment of the geographic

variation by Evans (1949) is inadequate; he only recognized the subspecies carthami

and moeschleri, which as will be shown below, do not represent the geographic variation

very well.

The variation comprises the following characters: size, extension of white spots above,

extension of white spots unh, extension of white superscaling above and unf, colour

and extension of the dark areas unh. Presumably, these characters are greatly influenced

by climatic conditions. It is therefore not necessary to assume in advance a historic

relation between forms that are of the same size, that have the same white superscaling,

etc. The relations supposed below are, at any rate, historically understandable.

nevadensis subspecies group

The position of ssp. nevadensis from the Sierra Nevada has long remained obscure,

due to lack of material. Neither Warren (1926), nor Alberti (1938) had seen this

subspecies. While he had only two specimens from the Sierra Nevada available for

comparison, Zerny (1927a) classified also specimens from Albarracin (C Spain) as

this subspecies. As a result the variation of the species in Spain remained unclear and

herewith also the position of the forms outside Spain.

The group is characterized by the pale yellow ground colour unh. This colour can be

almost white (nemausensis) but is usually overlaid by dark scales, especially around the

white spots. In this way lighter and darker patches are formed. In specimens from the

Pyrenees the darkening is usually more uniform. Size usually medium to small, in some

cases very large. Extension of white spots and light superscaling above very variable.

Unf usually light grey.

The group has a vast distribution, but the northern and eastern limits are very unsharp,

owing to intergradation with other subspecies groups. The distribution comprises Spain,

France and peninsular Italy. The transition to the carthami subspecies group is very

gradual. The greater part of E France should probably be considered a transitional zone,

as the influence of the carthami subspecies group is clearly perceptible in SE France

(Basses Alpes) and even in peninsular Italy, while the influence of the nevadensis sub-

species group can be traced in many German specimens. Such transitional populations

are historically easily understandable, but they cannot be classified. I regard all French

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

populations as belonging to the mevadensis group only for practical reasons. In N Italy

the influence of the valesiacus subspecies group is perceptible in some specimens from

the Ligurian Alps (by the size and the colour unh). Farther east, the populations of the

southern Alps show only valesiacus characters. Only rare specimens (e.g. Naturns,

Feltre) recall the nevadensis subspecies group. According to Verity (1940) specimens

transitional between ssp. valesiacus and ssp. “carthami’” occur in the Turin district. Four

subspecies have been described within the nevadensis subspecies group. They only

comprise a part of the total variation of the group. The remainder of the variation

will be discussed after the four subspecies. Further research will possibly indicate, that

the nevadensis subspecies group constitutes one single subspecies.

Pyrgus carthami nevadensis Oberthür

Pl. 3 Fig. 37, 46

Oberthür, 1910 — Et. Lép. Comp. 4: 384; pl. 55 fig. 474.

Type-locality: Sierra Nevada.

Material examined. — 30 & 11 2 Sierra Nevada (BM, ZMA, HC).

Distribution. — Sierra Nevada.

Subspecific characters. — Size moderate, 9 14—16 mm. Upf spots large, little

white superscaling. Uph spots usually well developed, especially the submarginal spots;

central band often vague. Unf dark to light grey. Unh spots well developed, especially

the central spot; spots bordered by dark brown lines; the darkening of the very light

yellow ground colour variable, but not extending over more than half of the ground

colour.

Note. — Oberthiir (1910), naming this subspecies, quoted the description of Rambur

(1858) who stated that the specimens from the Sierra Nevada differ from French

specimens, e.g. “par le dessous des ailes inférieures, dont la bord externe présente une

nuance d'un roussätre obscur, mais ne s’unissant pas complétement avec le dessin des

ailes comme chez l’Alvezs” (p. 67). I do not exactly follow the meaning of this, at

any rate I did not observe a reddish termen.

Pyrgus carthami pyrenaicus Warren

Pl. 3 Fig. 38, 47

Warren, 1926 — Trans. ent. Soc. Lond. 74: 69.

Type-locality: Pyrenees.

Nomenclature. — Verity (1928) thought it necessary to replace the name pyrenaicus by micro-

carthami as Tutt (1906) had given the name pyrenaica to a form of P. malvoides. This was not a

subspecific name and therefore it is not necessary to replace Warren's name (see also under P. mal-

voides).

Material examined. — 51 & 24 9:12 g 9 9 Spanish Pyrenees (prov. Huesca)

(RJ), 27 & 5 2 Hautes Pyrénées (BM, ML, ZMA, RJ), 12 g 10 9 Pyrénées Orien-

tales (BM, ML, ZMA).

Distribution. — Confined to the Pyrenees.

Subspecific characters. — The most important character is the small size, 4 13—14.5

mm. Above, spots medium-sized, on hindwing often obscure, light suffusion slight or

R. DE JONG: Pyrgus in the Palaearctic region 39

wanting; unh ground colour sometimes rather uniformly darkened, spots variable but

usually not as wide as in ssp. nevadensis.

Variation. — This subspecies seems to be most constant in the Central Pyrenees (at

both the Spanish and the French sides), and even there some specimens cannot be

separated from those from C France or C Spain. In the Eastern Pyrenees larger specimens

increase in number, giving this region the character of a transitional zone. Three males

examined, from the Spanish side (prov. Gerona), were indistinguishable in size (14.8—

15.2 mm) and in light superscaling, from specimens in C Spain. Therefore, I did not

include these specimens within ssp. pyrenaicus.

Pyrgus carthami lucasi Reverdin

Reverdin, 1928 — Bull. Soc. Lép. Genève 6: 91—92; pl. 2 fig. 5 and 6.

Type-locality: Forét de Benon.

Material examined. — 4 g° 4 9 Forêt de Benon (ML).

Distribution. — According to Picard (1948c), it is the race of the “plaines atlan-

tiques”.

Subspecific characters. — According to the original description it only differs

constantly from ssp. pyrenaicus by the presence of small white streaks in the dark

ground colour unf between the veins, beyond the median spots. However, Alberti

(1938) showed that the choice of this character is not very fortunate, as the white

streaks are present in many other forms.

Although I studied a limited number of specimens, I cannot consider the combination

of small size and very light ground colour unh the most important character, as stated

by Alberti (l.c.). The specimens examined show very well the characters mentioned by

Picard (1948c): reduction of the dark lines surrounding the ground colour unh, the

ground colour varying from vivid yellow orange to pale yellow. Sizes, measured by me,

were: d 11.5, 13.6, 13.8, 14.0 mm, 9 13.2, 14.8, 15.0, 15.1 mm.

Note. — The vivid yellow-orange ground colour unh may indicate influence of the

carthami subspecies group.

Pyrgus carthami nemausensis Picard

Picard, 1948 — Lambillionea 48: 28—29.

Type-locality: Nimes.

Material examined. — None.

Distribution. — According to Picard, confined to the rifle-range of Nimes.

Subspecific characters. — A very pale form; above not only a rather strong suffusion

with white scales, but the normal brown-black tinge is also very bleached; ground colour

unh very light: in the female only yellowish patches have been left, in the male the

ground colour is white with only traces of the dark lines around the spots. Even these

traces may disappear, making the unh unicoloured white (Picard named this form

deficiens). Somewhat smaller than the form of C France, with narrower wings.

Note. — This form is an extreme of the variation which may be found throughout

the range of the nevadensis subspecies group, especially in Spain and S France. I have

studied specimens from Albarracin (C Spain) and Digne (SE France) which show the

same characters, although not in an equally extreme way.

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

I doubt the usefulness of distinguishing subspecies when they are only based on an

extreme variation and, moreover, have a very limited distributional area. The rifle-

range of Nimes is a dry and hot locality. Perhaps the extreme environmental conditions

have caused the extreme variation. In this connection it is useful to quote the case of

the hesperiid Ochlodes venatum concoulense Picard. Picard (1948-c) distinguished

this subspecies on account of a series of specimens captured in three successive years at

a rather wet locality in S France (Mont Lozére), while everywhere in the surroundings the

subspecies septentrionale was flying. According to Gaillard (1952) the subspecies (sic!)

concoulense had lost its characters after three years of drought and had become septen-

trionale! (see also Chapter 1.b).

Remainder of the nevadensis subspecies group

The remainder of the local variation within the nevadensis subspecies group has not

yet been named. The name carthami has been used by most authors to cover this remain-

der variation. Alberti (1938) selected S Germany as the type locality of the species.

However, in my opinion another subspecies group occurs in S Germany, viz. car-

thami subspecies group; so, it is impossible to use the name carthami as a subspecific

name in the nevadensis subspecies group. It should be noted that some specimens from

S Germany are inseparable from French, or even Spanish or Italian specimens. This is

only to be expected, as in E France and S Germany the nevadensis and carthami sub-

species groups are mixed. In Spain the variation of the revadensis group is large. In

size the specimens vary from pyrenaicns to nevadensis, but as already noted by Alberti

(1938), very large specimens occur in the Sierra de Gredos; I saw a male from Navarre-

donde in the Sierra de Gredos of 16.5 mm and a female of 17.4 mm! (ZMA). Many

specimens from Albarracin are very strongly suffused with white scales above, some are

darker and a few are as dark as ssp. nevadensis, without reaching the same extension

of the white spots. At nearby Cuenca some specimens are almost inseparable from

specimens taken in the Sierra Nevada.

In France the specimens are less variable, 13—15.5 mm, with usually an obvious

white suffusion above. The white spots above are well-developed but not large and they

are often faintly visible on the hind wing, in particular the median spots. Some

specimens from the Basses Alpes and the Alpes Maritimes look like specimens from

Albarracin, but in the same region the dark suffusion of the hind wing below may cover

almost all the pale yellow ground colour. This ground colour is rather vivid yellow in

some specimens, indicating the influence of the carthami subspecies group.

In Italy the variation is about the same as in Spain, but I have not seen such large

specimens as from the Sierra de Gredos. In the Abruzzi most specimens are inseparable

from specimens taken in C Spain (including Albarracin). Specimens from Calabria are

rather large (up to 16.2 mm) and dark above, due to reduction of white spots and

scantity of white superscaling.

The only known specimen from the Netherlands, a male of 13.9 mm, was listed by

Lempke (1953) as ssp. septentrionalis, but in my opinion it belongs to the nevadensis

subspecies group.

Material examined. — 242 G 132 9:59 og 32 2 Spain (Andalusia to Gerona)

(BM, ML, ZMA, HC, RJ), 151 4 80 9 France (Pyrenees to Alps and to Fontaine-

blau) (BM, ML, ZMA, HC, RJ), 1 4 Belgium (Virton) (ML), 1 & Netherlands

(De Lutte) (ML), 29 4 20 9 Cand S Italy (BM, ML, ZMA, RJ), 1 & Sicily (BM).

R. DE JONG: Pyrgus in the Palaearctic region Al

valesiacus subspecies group

A well-defined group of medium-sized to large forms, occurring in the southern

Alps and southward to Bosnia and, largely separated, in N Germany. Above, light

suffusion generally scarce or absent, giving the specimens a dark appearance. Spots uph

usually faintly developed, in some specimens absent. Ground colour unh dark.

Three subspecies can be distinguished.

Pyrgus carthami valesiacus Mabille

Pl. 3 Fig. 40—41, 49—50

Mabille, 1876 — Bull. Soc. Ent. France (5) 5: 214.

Type-locality: Valais.

Material examined. — 90 8 44 9: 26 G 13 ® Switzerland (Valais) (BM, ML,

ZMA), 62 g 31 9 N Italy (Val d'Aosta to Trentino and prov. Udine) (BM, ML,

ZMA, RJ, HO), 2 $ Austria (Landeck) (RJ).

Distribution. — Southern Alps, not at the French side.

Subspecific characters. — Large, d to 16.8 mm. Above, white suffusion absent

(typical form) or obvious (f. major Rebel). Upf spots well developed, uph spots faint

or wanting. Unh ground colour a pale shade of yellow, but darker than in the zevaden-

sis subspecies group, with strong dark superscaling, giving the colour a dirty yellow-

green tinge with or without light patches. Unf black with sharply contrasting white spots.

There is a strong intermingling with ssp. carthami in Valais as stated by Verity (1940)

and Kauffmann (1951) (perhaps also with influence of the nevadensis subspecies

group). To the north and to the east the transition to ssp. carthami seems rather rapid.

The most northern locality from which I have seen ssp. valesiacus is Landeck in W

Austria. Eastward from the Gross Glockner all Austrian populations seem to belong to

ssp. carthami.

According to Kauffmann (1951) the occurrence and relative abundance of f. major

depend on micro-climate and weather conditions during the year.

Pyrgus carthami analogus Alberti

Pl. 3 Fig. 39, 48

Alberti, 1938 — Stett. ent. Zeit. 99: 244.

Type-locality: Trebevic, Sarajevo.

Material examined. — 63 & 6 9: 21 & Croatia (BM), 42 g 6 9 Bosnia

(Trebevic) (BM, ZMA, RJ).

Distribution. — Yugoslavian mountains; southern limit unknown.

Subspecific characters. — Medium-sized, $ 14—16 mm. On the upperside, this

form looks like ssp. valesiacus; uph spots often rather well-developed. Unh spots reduced,

especially the central spot, which almost always lacks the projection towards the base

of the wing; ground colour rather dark yellow, in most cases evenly overlaid by dark

scales, usually a warmer tinge than in ssp. valesiacus. The colour and reduction of spots

unh give the form an appearance that recalls P. serratulae balcanicus.

Note. — The populations from Croatia can best be listed as this subspecies, although

they show the influence of the neighbourhood of ssp. valesiacus and ssp. carthami.

42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pyrgus carthami septentrionalis Alberti

Pl. 3 Fig. 42, 51

Alberti, 1938 — Stett. ent. Zeit. 99: 236—237.

Type-locality: Neumark (Bärwalde).

Material examined. — 4 & 3 2 Berlin and environments (BM, ML, RJ), 1 &

Livonia (BM).

Distribution. — From Berlin north and north-east to the Baltic.

Subspecific characters. — As large as ssp. valesiacus or slightly larger. Upperside

white suffusion very scanty; spots somewhat reduced as compared to ssp. valestacus;

spots uph small but obvious. Unh dark grey-greenish, sometimes brownish, without light

patches; dark colour almost to the termen, white spots strongly contrasting, isolated but

well developed.

The specimens enumerated above agree well with Alberti’s description.

carthami subspecies group

Size medium to large, usually with well-developed white spots on both fore and

hind wings, light superscaling variable, usually well marked, rarely absent; ground

colour unh usually yellow to vivid orange-yellow, rarely as pale as in the mevadensis

subspecies group; dark suffusion brown, variable, but rarely covering all the ground

colour of the hind wing and not sharply outlining the white spots as in the nevadensis

subspecies group.

This group is widely distributed in E Europe, but occurs as far west as W Germany;

see also under the mevadensis subspecies group.

Pyrgus carthami carthami Hübner

Pl. 3 Fig. 43—44, 52—53

Material examined. — 101 g 60 9:29 g 12 @ Germany (Bavaria to Mainz)

(BM, ML, RJ), 14 g 13 9 Switzerland (excl. Valais) (BM, ML), 27 & 18 2

Austria (eastward from Gross Glockner) (BM, ML, RJ), 15 & 11 9 Hungary (BM,

ML, ZMA, RJ), 1 g 1 9 Czecho-Slovakia (BM), 14 & 5 9 Rumania (BM, RJ),

1 & Bulgaria (BM).

Distribution. — From N Switzerland and W Germany through C and E Europe.

The eastern limit is unknown. Some Hungarian specimens cannot be separated from

ssp. moeschleri.

Subspecific characters. — g° 14—16.1 mm. Differs from the only other subspecies of

the group by its smaller size and less developed spots above, especially on hind wing.

Variation. — In S Germany and Switzerland and, very rarely, in Austria, some

specimens have a pale yellow unh as in the nevadensis subspecies group. To the east the

specimens vary towards ssp. moeschleri.

Pyrgus carthami moeschleri Herrich-Schäffer

Pl. 3 Fig. 45, 54

Herrich-Schäffer, 1854 — Syst. Schmett. Eur. 6: 175.

Type-locality: designated hereby Sarepta.

Material examined. — 33 g' 21 2:29 g' 18 Q Sarepta (BM, ML), 1 & 1 9

R. DE JoNG: Pyrgus in the Palaearctic region 43

Guberli (BM), 1 & 1 9 Kisilskaja (RJ), 1 & Saisan (BM), 1 G Urumi (BM),

1 2 “Broussa’” (BM).

Distribution. — SE Russia, SW Siberia; according to Alberti (1938) the same form

occurs at Askhabad.

Subspecific characters. — Large, & 15.4—17 mm. Upperside, spots large, light

suffusion obvious; uph median band conspicuous and even one or two basal spots

obvious; upf obvious submarginal light streaks between the veins.

Geographic history. — The outline of the history of the species from the Last Glacial

to the present appears little complex. We may assume three or four refugial areas for

this species during the Last Glacial: (1) Spain, (2) N Italy - S Alps, (3) SE Europe

and, possibly separated from (3), (4) S Russia.

From Spain the nevadensis subspecies group spread postglacially far to the north,

but in E France it met carthami which spread westward from a SE European refugium,

possibly through the valley of the Danube. By this expansion ssp. carthami cat off the

northward expansion of the valesiacus subspecies group from its refugium in the southern

Alps. Alberti (1938) may be correct in supposing (indirectly) that the valesiacus sub-

species group was adapted to less favourable climates than the carthami subspecies group,

thus making the discontinuous distribution of the valesiacus subspecies group under-

standable. In its south-eastward extension the valesiacus subspecies group came into

contact with ssp. carthami. The result of this contact can be seen in ssp. analogus in

the smaller size and the warmer tinge of the ground colour unh, as compared to ssp.

valestacus.

Ssp. carthami and ssp. moeschleri, though closely related, may have had separate

refugial areas. Material from SW Russia probably could elucidate this point.

The history of the species before the Last Glacial cannot be reconstructed on the

ground of the present data.

6. Pyrgus sidae Esper

Esper, 1782 — Eur. Schmett. 1, 2: 178, pl. 90 fig. 3.

Type-locality: Volga region.

Distribution (Map 6). — Southern Palaearctic, from S France to Turkestan.

In France only known in the south, in the departements Hérault, Gard, Bouches-du-

Rhône, Var, Alpes Maritimes and Basses Alpes (Verity, 1947; Picard, 1948c; Gaillard,

1952; see also below, sub “Material examined”). In Italy widely distributed from

Liguria to the Monti Aurunci and from N Italy recorded from several localities in the

northeast (Verity, 1940).

From Hungary through the Balkans and Turkey to Turkestan, perhaps not con-

tinuously (a.o. Caradja, 1895; Bachmetjev, 1902; Rebel, 1903, 1904, 1913; Thurner,

1964; Groum-Grshimailo, 1890). In Iran only known from two specimens in the BM

labelled “Persia”. See also below, Material examined. Widely distributed in Trans-

caucasia, but from the Caucasus only known from Teberda and Itkol (Romanoff, 1884;

Alberti 1969). In Russia from Taganrog (Alpheraky, 1908) to the southern Urals

(Eversmann, 1844). Recorded by Lederer (1853) from the Irtisch River between

Ustkamenogorsk and Ustbuchtarminsk. This is the most eastern record of the species, but

it possibly concerns Syrichtus antonia and for that reason I consider the record unreliable.

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Habitat. — Dry and open, flowery places. Most common at low altitudes, particularly

in France and Italy (Warren, 1926; Verity, 1940), but in SE Europe to 2000 m (own

observation), likewise in the Caucasus at Itkol (Elbrus) (Alberti, 1969), and in Turke-

stan to 2700 m (Ugam Valley) (BM).

Biology. — Monovoltine; flight period from the middle of May to the end of July,

depending on altitude and local conditions. According to Guillaumin (1964) the larvae

have been found on Abutilon avicennae (Malvaceae).

Geographic variation. — There is a marked geographic variation in size, colour and

design and it is not difficult to distinguish three subspecies. However, the subspecific

classification of specimens from Central Asia is unsatisfactory.

Pyrgus sidae occiduus Verity

Pl. 4 Fig. 55—56, 64—65

Verity, 1925 — Ent. Rec. 37: 76.

Type-locality: Tuscany.

Material examined. — 88 4 54 9:62 G 30 9 France (Hérault, Bouches-du-Rhône,

Var, Alpes Marit, Basses Alpes) (BM, ML, MP, RJ), 26 8 24 © Italy (Firenze,

Monti Aurunci, Monti Meta, ‘Italy ') (BM, ML).

Distribution. — S France and Italy. For details of the distribution, see under the

distribution of the species.

Subspecific characters. — Generally smaller, & (and 9) 14—15.5 mm, but much

variation exists and there is a great overlap with the other subspecies. The smallest

specimen I have seen is a male from S France with a fore wing length of only 12.8 mm,

but also specimens (both & and © ) can be found of 16.6 mm or even more. Fig. 57 is

based on individual measurements of 30 specimens from S France and Italy (males and

females do not differ in size), which gives an impression of the difference in size with

more eastern populations and of the great overlap.

There are other differences which show less overlap. The usually bright orange

coloured bands unh in the nominate subspecies vary from pale yellow to pale orange

in ssp. occiduus, rarely reaching the bright orange of ssp. sidae; these bands are less

heavily outlined in black, more regularly formed and narrower in ssp. occiduus. The

last-mentioned character becomes clear when we measure the width of the central spot

along vein 5 and the breadth of the outer orange band along vein 5. Measured in this

manner, the central spot is wider than the orange band in ssp. occiduus, often twice

or even more times as wide; in ssp. sidae the central spot is, at the most, as wide as the

orange band and generally it is obviously narrower.

Finally, the spots on upperside are somewhat smaller and less square than in ssp.

sidae.

Note. — According to Evans (1949) two more or less intergrading forms exist, a

small form with yellow bands and a larger form, resembling ssp. sidae. The variation,

however, appears rather continuous and in my opinion it is undesirable to recognize two

forms.

R. DE JONG: Pyrgus in the Palaearctic region 45

fore wing length

mm 19,

184 ö a sn

17 à he

16 . :

15] :

14 . î

T T T T

France - Dalmatia - Bulgaria Turkey

Italy Mt. Olympus (Sliven) (Aksehir)

Fig. 57. Pyrgus sidae. Length of fore wing in four localities (regions). @ = one specimen. The line

connects the mean values (arrows)

Pyrgus sidae sidae Esper

PI. 4 Fig. 58—60, 67—69

Material examined. — 173 4 70 9:1 & 1 Q Hungary (Budapest) (BM), 1 &

2 9 Dalmatia (BM), 1 g' 1 9 Herzegovina (BM), 1 & Montenegro (RJ), 37 &

8 9 Macedonia (Yugoslavian and Greek) (BM, ML, RJ), 22 g° 23 Q Bulgaria

(Rila, N Rhodope, Sliven) (BM, RJ), 15 & 7 9 Bosphorus (BM), 60 g 15 2

Turkey (BM, ML, RJ), 5 g 1 @ Syria (BM), 1 G 1 @ Iran (“Persia”) (BM), 21 G

9 9 Sand E Russia (BM, ML, RJ), 2 $ Pamir (BM), 6 g° 2 9 Turkestan (Alex-

ander Mts, Naryn, Ugam Valley, Semirechensk, N Tianshan) (BM, ZMA).

Distribution. The whole distributional area of the species, except France, Italy and

NE Iran.

Subspecific characters. — Larger than ssp. occiduus, generally 16—18 mm, with

larger spots above, and bright orange bands unh that are strongly indented, broad and

heavily outlined in black. See also ssp. occiduus.

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Variation. — There is a general increase in size from west to east, Fig. 57. Apart

from this, the Bulgarian specimens differ obviously from the Turkish (when observed

in a series), by the smaller spots and less white superscaling above. Macedonian

specimens resemble Turkish. (Pl. 4 Fig. 58—60, 67—69).

The Central Asian specimens are small, & 14.4—16.4, 9 14.6—16.4 mm, upper-

side strongly marked in both fore and hind wing, bands unh pale to deep orange,

irregular, as wide as in Turkish specimens or narrower.

Pyrgus sidae evansi Alberti

Pl. 4 Fig. 57, 66

Alberti, 1957 — Mitt. Deutsch. Entom. Ges. 16: 27.

Type-locality: Arwas.

Material examined. — 3 &: 2 g Askhabad (paratypes; BA), 1 g' Arwas (ML).

Distribution. — Only known from the Koppe Dagh region: Askhabad, Arwas,

Nochur Tekke (Alberti, 1957).

Subspecific characters. — On the upperside the spots are well delevoped, but on the

hind wings usually only the submarginal spots are distinct. Approximately as large as

ssp. sidae, 15—18 mm. Underside with a dun tinge, orange bands faded yellowish,

with straight sides, somewhat like ssp. occiduus; median band wide.

Geographic history. — The recent history of the species appears little complex. Ob-

viously, ssp. occiduus originated during the Last Glacial in an Italian refugium. It is

possible that in Postglacial times it came into contact with ssp. sidae which spread

northwestward from a refugium in the Balkans and Turkey, having resulted in the

great variability of ssp. occiduus. The variation found in ssp. sidae may be of postglacial

age and is perhaps due to local climatic differences. From its East Mediterranean

refugium ssp. sidae extended its range not only westward, but also northward into

Russia, either along the west-side of the Black Sea, or along the Caucasus.

Presumably, the species was also isolated in a Last Glacial refugium in NE Iran (ssp.

evansi), but the present subspecific position of the Central Asian populations is not

clear. The material is insufficient to speculate on the history of the Turkestanian

populations.

7. Pyrgus badachschanus Alberti

Alberti, 1939 — Entom. Rundschau 56: 107—109.

Type-locality: Badakhshan (Sebaktal).

Distribution. — Except from the type-locality (Sebaktal = Sebak Valley) in NE

Afghanistan only known from Panjao (SW Koh-i-Baba Mts, W of Kabul) and from

the upper Panjshir Valley (N of Kabul) (Wyatt & Omoto, 1966).

Habitat. — The two type specimens came from the “Alpenwiesenzone, 2800 bis

3000 m” (Alberti, Lc.). Wyatt & Omoto (l.c.) collected the species in the upper

Panjshir Valley at 3800 m.

Biology. — Unknown.

Geographic variation. — Unknown.

Material examined. — 2 d 1 9, Hindu-Kush (upper Panjshir Valley) (CW).

(PIS rig BIS 33):

R. DE JONG: Pyrgus in the Palaearctic region 47

8. Superspecies Pyrgus alpinus Erschoff

Taxonomy. — Under the name of Pyrgus alpinus, Evans (1949) united three forms

which, according to him, have similar genitalia. Alberti (1952a), however, pointed

out that there are clear cut differences in the male genitalia, and the possibility of

distributional overlap without interbreeding led him to consider the three forms separate

species. Apart from some additional material I examined the same material as Evans did

and can only confirm Alberti’s conclusions. A distributional overlap is not yet known,

but the three forms apparently approach each other very closely. The use of the term

“superspecies” may indicate the supposed close relationship between these three forms

or semispecies which among Palaearctic Pyrgus species have a unique feature in com-

mon, viz. the absence of a costal fold in the male.

The differences in the male genitalia are mainly the shape of the cuiller, the length

of the style and the shape and spines of the antistyle (Fig. 58—60):

alpinus — cuiller relatively long, dorsal ridge often slightly concave; style almost

reaching distal end of cuiller; antistyle elongated, blunt, slightly spined.

cashmirensis — cuiller slightly shorter, apex usually strongly spined; dorsal ridge

often concave; style as long as in alpinus; antistyle short and broad, angular, strongly

spined.

darwazicus — cuiller rounded; style short, not nearly reaching distal end of cuiller;

antistyle short, but pointed, more or less spined.

As far as presently known these three types of genitalia are constant: no transitional

genitalia have been found. Even in the remarkable race cashmirensis pseudoalpinus —

externally indistinguishable from alpinus — the genitalia are entirely of the cashmirensis

type:

Alberti based pseudoalpinus on two males from Gilgit and the Baroghil Pass (N

58 1 mm a

Fig. 58—60. Male genitalia of superspecies Pyrgus alpinus. 58, P. alpinus (C Asia); 59, P. cashmi-

rensis (Pahalgam, NE Kashmir); 60, P. darwazicus lilliput (Anjuman Pass, NE Afghanistan)

Chitral) and supposed that the specimens in the BM from NW Kashmir (Hunza,

Yasin, Astor, Gilgit, Chitral), identified by Evans (1949) as alpinus, belonged to

cashmirensis psendoalpinus. Although I have seen all these specimens, I had the

opportunity to examine only a limited number of male genitalia. Based on the structure

of these genitalia, eleven males from Misgah (Hunza) appeared to belong to alpinus

mustagatae, seven males from Gilgit and Chitral to cashmirensis pseudoalpinus and one

male from Darkot in the extreme north of Chitral to “normal” P. alpinus. This is largely

in agreement with Alberti’s opinion (see further below under P. cashmirensis).

The easternmost locality of P. cashmirensis known is the Safed Koh in E Afghanistan

(I have seen the genitalia of the male in the BM). Wyatt & Omoto (1966) described

the new subspecies cashmirensis lilliput from the Anjuman Valley and the Anjuman

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pass in N Afghanistan. A distributional overlap of P. darwazicus and P. cashmirensis

seemed possible as darwazicus celsimontius was known to occur in N Afghanistan. I

examined three male paratypes of /i/liput. On dissection of two of them the genitalia

appeared to belong to the darwazicus type and not to the cashmirensis type.

Geographic history of the superspecies. — Undoubtedly, the history of the super-

species is not very old, but it is (yet) impossible to date the differentiation into three

species. All three appear to be confined to high altitudes. Therefore, warm periods,

rather than cold, may have led to their initial isolation. But, evidently glaciations could

also have brought about changes in these highly elevated mountainous areas.

8.a. Pyrgus alpinus Erschoff

Erschoff, 1874 — Lep. Turkestan: 24; pl. 2 fig. 18.

Type-locality: Kokand.

Distribution (Map 7). — C Asia from Serafshan throughout the mountains of W

Turkestan, at least to Issyk-Kul; further, Altai and Kamschatka (Alberti, 1952a) and

Vilui (Herz, 1898). Towards the south reaching Darkot (N Chitral) and Hunza (N

Gilgit) (BM).

Habitat. — According to Groum-Grshimailo (1890) it is confined to the alpine

meadow zone (in Pamir), from 3200—4300 m, in the Kounjout Mts. even higher; on

the Col Beik at 4800—5000 m.

Biology. — Presumably monovoltine. Groum-Grshimailo (l.c.) recorded as flight

period June-July, but specimens are also known from August and September.

Geographic variation. — As pointed out by Alberti (19522) specimens from

Serafshan to Kamschatka are all alike. However, east of Pamir a small form occurs in a

restricted area.

Pyrgus alpinus alpinus Erschoff

Material examined. — 894 40 9:3 4 Serafshan (BM, ML), 2 g 1 9 Samarkand

(BM), 2 g 1 @ Issyk-Kul, 32 & 13 9 Tian Shan (BM, ML, RJ), 23 & 12 2

Naryn River (BM, ML), 17 4 8 @ Ferghana (BM, RJ), 1 g 1 9 Alai (ML), 3 gd

2 9 Kashgar (BM), 1 ¢ Darkot (Chitral) (BM). Apart from these specimens I found

a number of specimens in the BM collections of which Evans, evidently, did )not

trust the labels: 3 g 1 Q “Amoor’ (Crowley Bequest), 1 4 “Amur”, 1 g “Russia”,

1 ©. ‘Siberia’.

Distribution. — The whole range of the species except that of ssp. mustagatae.

Subspecific characters. — For differences with ssp. mustagatae, see under that sub-

species.

From the other species of the superspecies (except cashmirensis pseudoalpinus) the

present species is easily distinguished by the strongly spotted upper side and par-

ticularly, by the two obvious white spots at the tornus uph.

Pyrgus alpinus mustagatae Alberti

Alberti, 1952 — Zeitschr. Lepid. 2: 80.

Type-locality: Mustagh Ata, Yarkand.

Material examined. — 11 & Hunza (Misgah) (BM).

R. DE JONG: Pyrgus in the Palaearctic region 49

Distribution. — Presently only known from the type locality and from N Gilgit

(material examined).

Subspecific characters. — Small, 11.2—12.2 mm. Above as ssp. a/pinus. Unh spots

reduced, central spot not projecting towards the base of the wing.

Note. — Alberti (l.c.) considered it a high altitude race but did not record exact

altitudes of material examined. As ssp. alpinus also seems to be confined to high altitudes

(see above, the habitat of the species), it is not very significant to call ssp. mzstagatae

a high altitude form.

Geographic history. — The absence of geographic variation over such a wide range

may indicate a fast and recent expansion of the range, not earlier than Last Glacial and

even now, the range may be more continuous than the present data suggest. Presumably

P. alpinus spread from C Asia, as all related forms and species occur there. Besides, if

alpinus originally was an East Siberian species one would have expected its occurrence

in N America, for this highly cold-adapted species must have been able to cross Beringia.

8.b. Pyrgus cashmirensis Moore

Moore, 1874 — Proc. Zool. Soc. Lond. 1874: 274; pl. 42 fig. 7.

Type-locality: Kashmir.

Distribution (Map 7). — From Safed Koh and Chitral through Kashmir to Kumaon;

Bhutan (BM). The known localities suggest that the species may also occur in Nepal.

Habitat. — In Gilgit at least to 4300 m (BM).

Biology. — Unknown.

Geographic variation. — Although the male genitalia do not seem to vary geogra-

phically, there is a remarkable variation in external characters leading to the recognition

of two subspecies: in Chitral and Gilgit a form occurs which is externally indistinguish-

able from P. alpinus. This was discovered by Alberti (1952) who had two males

available. As mentioned earlier, I examined male genitalia of only few specimens.

As I am not entirely certain about other specimens from Chitral and Gilgit, only the

specimens thus examined by me will be enumerated below. It is, however, interesting

to note that 55 g and 25 ® from Chitral, Gilgit, etc. in the BM vary in external

characters from nearly P. cashmirensis to completely P. alpinus.

Pyrgus cashmirensis cashmirensis Moore

Material examined. — 76 & 28 9:1 g' Safed Koh (BM), 4 & 4 ® Baltistan

(BM), 141 G 17 @ Kashmir (BM, RJ), 12 & 2 2 Ladakh, 5 g 1 2 Lahoul, 10 &

3 9 Kulu, 1 & Kumaon, 2 & Bhutan (all BM).

Distribution. — The range of the species except Chitral and Gilgit.

Subspecific characters. — Easily distinguished from ssp. psendoalpinus (and from

P. alpinus) by the reduced spots, above; spots upf small, uph faint or absent.

Pyrgus cashmirensis pseudoalpinus Alberti

Alberti, 1952 — Zeitschr. Lepid. 2: 85.

Type-locality: Gilgit.

Material examined. — 6 o Chitral, 1 & Gilgit (BM).

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Distribution. — Confined to Chitral and Gilgit. Shirözu & Saigusa (1963) recorded

this form from W Pakistan under the name Pyrgus alpinus alpinus. They gave nice

figures of male genitalia (fig. 2) and the upper- and underside of two males (not one

male and female, as stated). The localities in W Pakistan are: Mochodan-Tikatoki and

Tikatoki-Diwan Gal.

Subspecific characters. — Externally like P. alpinus.

Variation. — The male examined from Gilgit had a transitional facies and resembled

more ssp. cashmirensis than P. alpinus. There may be a continuous variation from the

cashmirensis facies to the alpinus facies.

Geographic history. — Presently, nothing can be said on this subject.

8.c. Pyrgus darwazicus Groum-Grshimailo

Groum-Grshimailo, 1890 — in Romanoff, Mém. Lép. 4: 501; pl. 21 fig. 5.

Type-locality: Darwaz (Touptschek).

Distribution (Map 7). — In collections, a very rare species of which only few localities

are known. In the BM there are specimens from Darwaz, Samarkand and Trans-Alai.

According to Alberti (1952a) further only known from the Alai, the Sebak Valley and

the Nuksan Pass (NE Hindu-Kush). From these scanty data Alberti thinks that P.

alpinus and P. darwazicus are possibly partly sympatric (P. alpinus is also known from

Samarkand and Alai). However, by the locality “Samarkand” usually the province

is meant and not the city (the same applies to Bokhara). The province has a

length of more than 500 km and a mountainous part stretching over more than 400 km.

Likewise, the Alai is an extensive region. Therefore, on the ground of the few specimens

known there is no reason to consider the two species sympatric in these regions.

As stated above, the race described by Wyatt & Omoto (1966) as Pyrgus cashmirensis

lilliput appeared to be a race of P. darwazicus. The known localities of P. darwazicus

can thus be supplemented by Bala Quran and the Anjuman Pass, both in the Anjuman

Valley in the Hindu-Kush, southwest of the Sebak Valley.

Besides, Wyatt & Omoto recorded possible P. darwazicus specimens from the Anjuman

Pass (1 g') and from the Shiva Mts. (NE Badakhshan) (1 g', 2 9). I examined

one of the females of the Shiva Mts. This rather worn female is geographically as well

as externally intermediate between ssp. darwazicus and ssp. celsimontius, perhaps

somewhat closer to ssp. celsimontins. I also examined the genitalia, but assurance on the

identification cannot be given, as the female genitalia of a true P. darwazicus are

unknown to me. For the present time it seems best to classify the Shiva female as

P. darwazicus (it will be enumerated below under ssp. celsimontins).

Habitat. — All known specimens are from higher altitudes (as far as recorded):

2800—4100 m, the alpine meadow zone (Groum-Grshimailo, l.c.; Alberti, Lc. Wyatt

& Omoto, l.c.).

Biology. — Unknown.

Geographic variation. — There is a marked variation in size, in the extension of the

spots and sometimes some variation in the shape of the antistyle, but the material is

too scarce to be certain.

R. DE JONG: Pyrgus in the Palaearctic region 51

Pyrgus darwazicus darwazicus Groum-Grshimailo

Material examined. — 3 & 2 9:2 8 Darwaz, 1 g' 1 9 Samarkand, 1 9 Trans-

Alai (all BM).

Distribution . — Southern C Asia, north of Afghanistan.

Subspecific characters. — Dark, with reduced spots; upf spots very small, uph spots

very faint or absent, unh spots reduced and with vague outline, recalling Pyrgus cacaliae.

Pyrgus darwazicus celsimontius Kauffmann

Kauffmann, 1952 — in Alberti, Zeitschr. Lepid. 2: 85—86; figs. 3, 4.

Type-locality: Sebak Valley (Badakhshan).

Material examined. — 1 9 Shiva Mts. (CW).

Distribution. — Probably a limited distribution in Badakhshan.

Subspecific characters. — Strongly spotted, somewhat like P. alpinus. Uph with very

obvious central spots, but remainder of median band almost absent, submarginal spots

indicated by greyish scales. Smaller than ssp. darwazicus, according to the original

description g' 24—26 mm, ® 29 mm, undoubtedly “from tip to tip”.

Variation. — The female examined has less distinct spots than the figure of ssp.

celsimontius in the original description. It may be transitional to ssp. darwazicus; see

also above. Wing-span from tip to tip: 27.5 mm, length of fore wing 14.1 mm.

Pyrgus darwazicus lilliput Wyatt & Omoto

Pl. 1 Fig. 18, 36

Wyatt & Omoto, 1966 — Entomops 1966 (6): 195—196.

Type-locality: Bala Quran (Anjuman Valley, Hindu-Kush).

Material examined. — 3 4 Anjuman Valley (paratypes) (CW).

Distribution. — Only known from the Anjuman Valley.

Subspecific characters. — Very small, 10.5 mm; according to the original

description the males are 10—12 mm, the female 13 mm. Above, better marked than

in ssp. darwazicus, more or less like P. cashmirensis; uph submarginal median spots

clearly visible, according to the original description in the female very distinct. Underside,

dark, recalling ssp. darwazicus.

In the male genitalia the antistyle seems to be less pointed. However, no data on

individual variation can be presented, as genitalia of only two males were examined.

Likewise, the individual variation in the nominate subspecies is unknown (cf. Alberti,

1952a).

Geographic history. — At present no information can be given on this subject.

9. Pyrgus andromedae Wallengren

Wallengren, 1853 — Of. K. Vet. Akad. Förh. 10: 25.

Type-locality: Dalarne (selection by Opheim, 1953).

Distribution (Map 8). — Arcto-alpine and confined to Europe. In Scandinavia

confined to the mountains from the North Cape south to Dovre and Dalarne, most

localities being situated within the Arctic Circle (for detailed distribution map, see

Nordstrém, Opheim & Valle, 1955).

Widely distributed in the Pyrenees, from Col du Pourtalet in the Basses Pyrénées

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

(RJ) to Mont Canigou in the Pyrénées Orientales (Dufay, 1961); from the Spanish

side only recorded from Ordesa, Sanctuario de Nuestra Sefiora de Las Bordas (Valle de

Aráú) (Rondou, 1932; Manley & Allcard, 1970). In the Alps it is one of the most

universally distributed species of the genus and almost anywhere above 1700 m one

may expect to find it (Warren, 1926). Known from the Basses Alpes and Alpes

Maritimes to S Bayern, Styria and the Triglav (NW Yugoslavia) (a.o. Alberti, 1927;

Verity, 1940, 1947; Kauffmann, 1951; see also under “Material examined”).

Further only known from the high mountains of Yugoslavia: Treskavica, Prenj, Velez

and Baba planina near Gacko, all in Bosnia (Rebel, 1904); Bretherton (1966) records

the species from the Shar planina, between Servia and Macedonia.

Habitat. — Mountain slopes, moorland, usually near water. According to Warren

(1926), banks of a mountain torrent are a favorite haunt, though swampy ground of

any kind is equally suited. In the Alps usually from 1600 to 2500 m (Kauffmann,

1951), but single specimens may be found up to 3300 m (Warren, 1926, Verity,

1940). Unlike P. cacaliae it is rarely abundant (Warren, l.c.). In Scandinavia in the

upper birch zone and beyond the tree limit (Nordstrom, e.a., 1955; Gullander, 1959).

Biology. — Monovoltine; flight period between the end of May and half August,

depending on altitude (Kauffmann, l.c.). Larval stages and food plants unknown.

Geographic variation. — In spite of the great disjunction in the distributional area,

geographic variation is unknown for this species. Even a fervent splitter like Kauff-

mann (1951) failed to find geographic variation.

Material examined. — 121 8 56 2:19 8 21 9 N Scandinavia (Norway, Sweden,

Finland) (BM, ML, RJ), 3 g 1 @ Pyrenees (Basses Pyr., Col du Pourtalet; Hautes

Pyr.) (BM, RJ), 7 & 2 2 French Alps (Basses Alpes, Hautes Alpes, Savoie) (BM),

47 d 16 9 Switzerland (BM, ML, ZMA), 5 d 2 9 Bavaria (Oberstdorf, Allgau,

Garmisch) (BM, ML, RJ), 2 & Italia (Aosta) (BM), 36 4 13 @ Tirol and Austria

(BM, ML, RJ), 2 & Bosnia (Trenj) (BM), 1 £ Hungary (“Hungrie’) (BM).

Geographic history. — The postglacial history of this species must have run almost

parallel to that of P. cacaliae. Apart from coincidences, the difference in the distribu-

tional areas may be due to a slight difference in habitat preference. There are no traces

of a disjunction during the Last Glaciation. See also Chapter 2.d.

10. Pyrgus cacaliae Rambur

Rambur, 1839 — Faune entom. de l’Andalousie 2: pl. 8 figs. 6, 7, k.

Type-locality: Grande-Chartreuse de l'Isère (Verity, 1925).

Distribution (Map 9).— Pyrenees, Alps, southern Carpathians, Balkan Mts. and Rila

Dagh. In the Pyrenees only few localities are known, situated in Andorra, and in the

departements Pyrénées Orientales, Ariége and Hautes Pyrénées (Caradja, 1893; Rondou,

1932; Picard, 1946, 1947). In the Alps widespread, from the Alpes Maritimes to the

Schneeberg in Lower Austria (see Material examined). In the southern Carpathians

only known from the Bucegi Mts and from Kronstadt (Niculesci, 1968 in litt.; Higgins

& Riley, 1970; BM). In Bulgaria known from the Central Balkan Mts (Drenowsky,

1909, 1910) and the Rila Dagh (Züllich, 1936—37; BM).

Habitat. — Moist and dry, flowery localities at high altitudes. In the Pyrenees not

below 2000 m (Picard, 1947), in the Alps usually from 2000 m to 2600 m, rarely from

R. DE JONG: Pyrgus in the Palaearctic region 53

1500 m upward (Verity, 1940; RJ), in the Balkan Mts. not below 1800 m (Drenowsky,

1910). In the Alps much more local than P. andromedae, but often very abundant

(Warren, 1926; Verity, 1940).

Biology. — Monovoltine, from the last part of June to half August; the flight period

in the Alps lasts about five weeks (Kauffmann, 1951).

Thanks to a partly succeeded breeding experiment of Kauffmann (1954c) we

know that the larvae hibernate at a very young stage: a few days after hatching they

enter diapause, in the soil or between leaves, spun together; for food plants Potentilla

aurea and Sibbaldia procumbens (both Rosaceae) are considered; the often mentioned

Tussilago farfara (Compositae) (e.g. Verity, 1940; Higgins & Riley, 1970) is probably

not accepted as food plant by the larvae.

Geographic variation. — A species restricted to high altitude is apt to produce mor-

phologically distinguishable, local forms. The present species, however, shows little

variation, individual or geographic. The variation mainly concerns the extension of the

markings. In the same locality specimens without spots above (forma caecus Reverdin,

1912) occur together with rather well-marked specimens.

According to Picard (1947), eleven out of twelve specimens studied from the

Pyrenees differed from specimens from the Alps in a greatly enlarged basal spot in

space 7 unh. On this ground he described a new subspecies pyrenaeus. I have seen no

material from the Pyrenees, but it is, of course possible that the enlarged basal spot is

a constant characteristic of the Pyrenean populations and it may be useful to distinguish

the subspecies pyrenaeus. But as the difference is small and the material scanty and as,

for the present study, the distinction of subspecies is only a means, not an aim,

mentioning of Picard’s description may suffice.

Kauffmann (1951) described the subspecies prosensis from St. Gotthard-Hospiz and

neighbouring localities. This form differs from the typical form in being larger,

14.2—15.1 mm against 13.7—14.9 mm, with larger spots above, unf whitish and unh

yellowish with still more indistinct markings. Kauffmann (p. 342) says that “this sub-

species forms about two-thirds of the population”. This is, of course, a careless way of

saying that about two-thirds of the specimens of the subspecies have the above-

mentioned characteristics. Apparently, it is an extreme variation. However, the com-

bination of a large size, large spots and yellowish unh seems to be more characteristic

than the occurrence of all these characters apart, as all can be found elsewhere, par-

ticularly the yellowish unh. The large spots may be the most characteristic, but as there

is so much overlap in all three characters mentioned above, I doubt whether it is useful

to recognize prosensis as a separate subspecies.

According to Higgins & Riley (1970) the white spots upf are larger in the specimens

from the Carpathians than in specimens from the Alps. I saw only 1 4 and 1 9 from

Kronstadt. Like the Bulgarian specimens these did not differ remarkably from specimens

from the Alps.

Material examined. — 173 J 121 9:5 G 3 Q Alpes Maritimes (BM), 34 &

11 Q Basses Alpes (BM, RJ), 62 8 53 ® Swiss Alps (BM, ML, ZMA, RJ, HC),

4 © Bavaria (Allgäu; ‘“Germania”) (ML, BM), 56 4 39 ® Tyrol (from Ortler to

Gross Glockner) (BM, ML, ZMA, RJ, HO), 9 £ 8 © Carinthia (BM, RJ), 1 «4

Radstädter Tauern (RJ), 2 & Steiermark (Hochschwab) (ML, RJ), 2 g 1 © Lower

Austria (Schneeberg) (ML, RJ), 1 & 1 9 Rumania (Kronstadt) (BM), 3 ¢ 3 2

Bulgaria (Rila Dagh, Musalla Mts) (BM).

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Geographic history. — The present localities of P. cacaliae must be considered post-

glacial refugia for this cold-adapted species. During the Last Glacial the species ap-

parently, had a continuous distribution area, somewhere between the land ice in the

north and the glaciers of the Alps. At any rate, there are no traces of an earlier than

a postglacial disjunction. For a reconstruction of the more ancient history of the species,

see Chapter 2.d.

11. Pyrgus centaureae Rambur

Rambur, 1839 — Faune entom. de l’Andalousie 2: pl. 8 fig. 10.

Type-locality: Dalécarlie (= Dalarne).

Distribution (Map 9). — Discontinuously Holarctic: Scandinavia, Siberia, North

America. In Scandinavia widely distributed, from S Norway and C Sweden to the

Barents Sea and throughout Finland (for details, see Nordström, Opheim & Valle,

1955). East of Finland the species is only known from the formerly Finnish districts

(Kaisila, 1947). Undoubtedly the species occurs further east, but records are totally

absent.

The distribution in Siberia is very imperfectly known: lower course of the Yenisey,

between 68° and 69° N Lat. (Trybom, 1877), Altai and Sajan Mts (a.o. Elwes, 1899;

see also below, Material examined), and Vitim (Herz, 1898; BM). It is rather impro-

bable that the species is absent from the intermediate regions.

In America widely, but possibly discontinuously distributed, from NW Yukon (NW

Canada) through British Columbia southward to the high mountains of Colorado and

through the NW Territories and Manitoba eastward to Newfoundland and Labrador and

from there southward to N Carolina. For details, see Klots (1951), Evans (1953) and

Freeman (1958).

Habitat. — A typical bog species, in the mountains up to just above the timber line

(Petersen, 1954; Nordström e.a., 1955; Gullander, 1959). In Canada and Colorado as

in Europe, but from New York to N Carolina (ssp. wyandot) on open, grassy hillsides

(Klots, 1951).

Biology. — Monovoltine; flight period June-July. Larval food plant Rubus chamae-

morus (Rosaceae) (Nordstrém, Wahlgren & Tullgren, 1941; Gullander, 1959).

Geographic variation. — Although America falls outside the limits of the region set

for this study, the geographic variation in this part of the range may throw light on the

history of the species and for this reason we should also deal with the American forms.

Moreover, I will stress the subjectivity of considering the Eurasian and American forms

conspecific. There is no proof of their conspecificity and we cannot state that they are

or are not conspecific, only that we consider them as such.

The variation affects design, male and female genitalia. Three subspecies groups can

be distinguished:

(1) European — subspecies centaureae and schoyeni

(2) Siberian — subspecies sibiricus

(3) American — subspecies freija, loki and wyandot.

(1) European subspecies group. — Above with well-developed spots; unh, sub-

marginal spots usually detached from the termen, except the spot in spaces 4—5,

sometimes also the spot in space 1-c attached to the termen. Male genitalia: cuiller bluntly

rounded at apex, without any specialized terminal structure; antistyle sharply turned

R. DE JONG: Pyrgus in the Palaearctic region 55

upwards at its ventral end. Female genitalia: sternum of eighth abdominal segment

usually less sclerotized and lines of corrugation less developed than in other subspecies

groups, but there exists much variation (Fig. 25, 26; see also Opheim, 1959).

Pyrgus centaureae centaureae Rambur

Material examined. — 116 4 76 9:12 g' 5 £ Norway (Langfjordvand, Finmark,

“Norway’) (BM, ML), 1 9 Dalarne (RJ), 93 4 58 ® Swedish Lapland to Torne

Lappmark) (BM, ML, ZMA, RJ), 2 4 4 9 Finland (Enotekio, Lamm, ‘‘Finland’’)

(BM, RJ), 9 & 9 2 “Scandinavia” (BM).

Distribution. — The European part of the distributional area of the species except the

localities of ssp. schoyeni (see below).

Subspecific characters. — See above. Differs from ssp. schoyeni by the heavier white

markings unh; particularly the submarginal spots better developed.

Variation. — Rarely specimens occur with a basal spot in space 2 upf. Opheim

(1959), who named this form #ripuncta, knew only of eight such specimens; I have

seen ten others. This variation is interesting as the basal spot in space 2 upf occurs in

all American and in all Chinese Pyrgus species and also in P. badachschanus and P.

andromedae.

Pyrgus centaureae schoyeni Opheim

Opheim, 1959 — Astarte 18: 4, fig. 1.

Type-locality: SE Norway.

Material examined. — None.

Distribution. — Isolated lowland moors in the extreme southeast of Norway (Skarnes-

Saeterstoa in Sor-Odal and Bjorkelangen in Nordre Holand) (see Opheim, 1959).

Subspecific characters. — Unh ground colour very dark, spots somewhat reduced,

particularly the submarginal spots and the outer spot in space 7.

(2) Siberian subspecies group. — Above spots reduced, though in some females

strongly developed. The upperside varies from a sharply marked to a rather washed-out

appearance, somewhat recalling P. cacaliae. Unh varies from sharply marked as in ssp.

centaureae to rather vaguely marked (but with markings much sharper than in P. caca-

liae). The submarginal spots unh in spaces 1-c and 4—5 usually extending to the termen.

Male genitalia: cuiller variable, from narrow throughout and pointed at apex, to

broadening and bluntly rounded at apex, with very fine spines at or near apex; anti-

style as in ssp. centaureae. Female genitalia: sclerotization and corrugation of the sternum

of the eighth abdominal segment heavier than in ssp. centaureae, in addition strong

corrugations in the intersegmental membrane between the seventh and eighth segments

(Warren, 1957).

Pyrgus centaureae sibiricus Reverdin

Reverdin, 1911 — Bull. Soc. Lép. Genéve 2: 78—80, pl. 11 fig. 13, 14, pl. 13 fig. 6.

Type-locality: Altai.

Material examined. — 14 & 14 9:10 & 11 9 Altai (Ongodai, Bashkaus, Tchuja

Mts, Arazan), 4 & 2 9 Sajan Mts. (Sajan, E. Sajan), 1 © Vitim (all BM).

Distribution. — The Siberian part of the range of the species, but it is unknown

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115. AFL. 1, 1972

whether the specimens of Trybom (1877) from the lower Yenisey belong to this

subspecies or rather to the Scandinavian form.

Subspecific characters. — See above.

Note. — I cannot refrain from recording a case, in which measuring by two standards

makes systematics inacceptable. Warren (1926: 41—42) described the new species

chapmani, based on four males and one female from Munko Sardyk (Sajan Mts),

Vitim and “S Siberia”, and added (p. 41): “This species is very close to H. sibirica, and

though abundantly distinct anatomically, I am not able to separate the two by their

superficial facies” and (p. 42): “We must rest content for the present only to record

the fact that these two species exist, and hope that the future will bring us fresh light on

the subject.” At that time, systematics were based on the typological species concept and

any morphological difference was considered proof of specific distinction. The future

brought fresh light on the subject: Warren (1957) had examined one female of sibiri-

cus and one female of chapmani and as the genitalia of these specimens were similar,

he concluded (p. 376): “The slight differences in the male genitalia must be individual

variations, the few specimens dissected being insufficient to demonstrate this.” I fail

to see why four males are insufficient to demonstrate individual variation and why two

females are sufficient to demonstrate constancy. Besides, in 1926 Warren was not able to

separate the females of sibiricus and chapmani, but in 1957 forgot to mention how he

could distinguish the females then. Perhaps he only examined two specimens of sibiri-

cus (or of chapmani).

I was not able to study female genitalia of sibiricus, but fortunately, could examine

those of 12 males, in which I found a large range of variation, from the narrow and

pointed chapmani type of cuiller, through the sibirica type, to almost the centaureae type

(Fig. 55, 56). Therefore I think Warren was correct in considering chapmani and

sibiricus conspecific, not because their females are alike, but rather, because there is no

reason to consider them distinct, as in every species one can expect some individual

variation in any character.

Moreover, if one accepts the biological species concept, one can only consider szbirieus

a subspecies of centaureae. This was done by Evans (1949), who listed chapmani as a

synonym of sibiricus, and sibiricus as a subspecies of centaureae. Unfortunately, Warren

(1957) neglected both Evans’s action and the biological species concept.

(3) American subspecies group. — Upf bar at end of cell usually better defined

than in the Eurasian forms; unh submarginal spots separated from the termen throughout

(except along the veins). According to Evans (1949) the basal cell spot unh is more

or less widely separated from the spot near base of space 7, whereas in ssp. centaureae

they usually adjoin. There is, however, too much variation in both ssp. centaureae and

the American forms, to use this as a distinguishing character.

Male genitalia: cuiller somewhat variable, apex usually pointed, dorsal edge of cuiller

finely spined, sometimes only a little serrated; antistyle spined or indented, but not

with a sharply upturned point as in ssp. centaureae. Female genitalia: corrugation of

the sternum of the eighth abdominal segment rather stronger and the proximal lines

running straighter than usually in ssp. centaureae, more recalling the structure of ssp.

sibiricus, but without the corrugation of the intersegmental membrane. (I did not

examine female genitalia of American specimens, but refer to the figures and description

by Warren, 1957).

R. DE JoNG: Pyrgus in the Palaearctic region 57

Pyrgus centaureae freija Warren

Warren, 1924 — Trans. ent. Soc. Lond. 72: 56—57, pl. B, fig. 1—7.

Type-locality: Labrador.

Material examined. — 19 ¢ 5 £ Labrador (BM, ML).

Distribution. — Northeastern part of the American range of the species; Labrador,

south to Gaspé. Western limit of this subspecies unknown (cf. Klots, 1951; Evans,

1953).

Subspecific characters. — Unh white areas well-developed, giving the hind wing a

much whiter appearance than in the other American forms. Small, 4 13.0—13.8 mm.

According to Klots (1951), the style in the genitalia of males from Manitoba to La-

brador is shorter than in ssp. wyandot. I only examined the genitalia of nine American

males (wyandot 2, loki 3, freija 4), but found the longest style in Labrador specimens.

Note. — The history of freija in literature is full of misconceptions. It is not neces-

sary to go deeply into the subject, but it may be useful to summarize the difficulties

briefly.

The form was described as a distinct species occurring in Lapland and America. Bell

had called attention to freija as an American species in 1926; already two years later

Lindsey (1928) questioned the specific rank of freija, comparing specimens from New

Jersey, Labrador and Lapland. Therefore, Warren (1957) is incorrect, saying that:

“those workers who stated that freija and centanreae were the same species were only

comparing two races of freija; the latter and wyandot” (p. 375).

Another problem, viz. the occurrence of freija in Lapland, was dealt with extensively

by Opheim (1951, 1953, 1959). Only two females from Lapland were known, attributed

by Warren (1951, 1957) to freija, but Opheim showed that they fall within the varia-

tion of Scandinavian centaureae.

Pyrgus centaureae loki Evans

Evans, 1953 — Cat. Amer. Hesp.: 214.

Type-locality: Long Peak Trail (Colorado).

Material examined. — 27 & 10 9:5 8 2 9 Canadian Rockies (BM, RJ), 22

8 ® Colorado (South Park, Glen Cove, Bullino, Hall Valley) (BM).

Distribution. — The western part of the American range of the species; eastern limit

unknown.

Subspecific characters. — Above more strongly marked than ssp. freija; unh white

spots reduced, ground colour somewhat yellower. Larger than ssp. freija, 4 13.8—14.8

mm. The unh appears somewhat intermediate between ssp. freijd and ssp. wyandot, but

still closer to ssp. freija.

Pyrgus centaureae wyandot Edwards

Edwards, 1863 — Proc. Ent. Soc. Philadelphia 2: 21, pl. 5.

Type-locality: Long Island.

Material examined. — 11 8, 7 9:1 8 New York, 8 g 5 9 New Jersey, 1 gd

1 9 N Carolina, 1 & 1 2 Virginia.

Distribution. — New York south to N Carolina (Klots, 1951; Evans 1953).

Subspecific characters. — Size as in ssp. frezja. Unh spots reduced, particularly the

submarginal spots, somewhat recalling the unh of P. malvae but spots larger; ground

colour yellowish brown.

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Geographic history. — For the reconstruction of the geographic history of the species

the occurrence and differentiation in America is elucidating. P. centaureae is undoubted-

ly a Pleistocene invader in America, as all close relatives are cold-adapted, Palaearctic

species (see also Chapter 2.4). The change of habitat (undoubtedly connected with a

change of food plant) in ssp. wyandot appears to be of a recent age.

The Bering Strait area could only be crossed by land during glacial and perhaps in

early interglacial times (Petersen, 1954). If P. centaureae would have reached America

during the Last Glacial, the ice shield which isolated ice-free Alaska from the rest of

ice-free America (cf. Petersen, l.c., fig. 2), would have prevented a further expansion

of the range in N America until postglacial times. The occurrence of P. centaureae in

the high mountains of Colorado is only understandable, if we suppose that this species

was present in N America before the Last Glacial. It seems that an invasion into

America earlier than during the Third Glacial (i.e. during the Second Glacial or earlier)

would have left behind much more differentiated forms, as the American populations

would have been isolated from the Palaearctic ones during the very long Second Inter-

glacial. Consequently we may assume, that P. centaureae invaded America during the

Third Glacial. Thus the species came into being in the Palaearctic not later than during

the Third Glacial.

The differentiation of the species in the Palaearctic does not give a reason for sup-

posing an earlier origin. The differentiation into a European and a Siberian group may

even be of a postglacial age, but it is impossible to date the differentiation more exactly

as long as the occurrence in N Russia and Siberia is so imperfectly known. Isolation

during the Last Glacial could have been caused by the inundated W Siberian lowlands

(cf. Moreau, 1955).

In Europe P. centaureae is a postglacial invader which spread to the west with the

taiga to which its habitat (mire) is so closely connected. The absence of the food plant

(Rubus chamaemorus) from the C European mountains may have caused the absence of

P. centaureae there.

According to Petersen (l.c.) the mire species are adapted to a cold, continental climate.

This can be demonstrated by the distribution of P. centaureae in Norway, where it is

absent from the coastal regions. It is hardly probable that the species has been isolated

on the western coast of Norway during the Last Glacial, as supposed by Opheim (1951),

to explain the variability of P. centaureae in Scandinavia. At least, the optimal habitat of

this species was not present during the Last Glacial along the Norwegian coast.

The origin of ssp. schoyeni is of a very recent, no doubt postglacial age. The fact that

it is less variable than ssp. centaureae (Opheim, 1959) can be due to the restricted range

of the former subspecies and to a restricted number of ancestor specimens.

12. Superspecies Pyrgus malvae Linnaeus

Taxonomy. — The taxonomy of the superspecies has been discussed in Chapter 2.b

and is summarized here. The superspecies consists of three semispecies: P. malvae, P.

malvoides and P. melotis. P. malvae and P. malvoides interbreed; interbreeding of P.

malvae and P. melotis is not improbable, but has not yet been established.

Distribution. — The superspecies is distributed throughout the Palaearctic (with the

exception of very cold or dry regions). The distribution of the semispecies is as follows:

P. malvoides — SW Europe (Portugal, Spain, S France, S and E Switzerland, W

Austria, Italy).

R. DE JONG: Pyrgus in the Palaearctic region 59

P. melotis — south of the Caucasus and the Black Sea.

P. malvae — the remainder of the distribution area of the superspecies.

Geographic and evolutionary history. — Various authors (Picard, 1948a, 1950a; War-

ren, 1957; Sichel, 1962a) have attempted to outline the geographic history of superspe-

cies malvae, but only Sichel has taken the far-reaching influence of the Pleistocene

glaciations into account. As shown below, each of the three semispecies can be divided

into two subspecies. These subspecies clearly indicate the refugial areas of the super-

species during the Last Glacial:

malvoides malvoides — Spain

malvoides modestior — Italy

melotis melotis — Syria

melotis ponticus — E Turkey

malvae malvae — SW Siberia / SE Russia / SE Europe

malvae kauffmanni | — S Manchuria and Korea.

The following notes are added:

(a). The differentiation into ssp. melotis and ssp. ponticus is not well understood. The

Taurus Mountain System may have had influence on the geographic isolation which led

to the differentiation of both forms (cf. Kosswig, 1955, fig. 12). However, the

occurrence of ssp. melotis on the Island of Milo is rather puzzling. This led Sichel (1.c.)

to the assumption of a second refugium in W Anatolia. If the specimen in the British

Museum labelled “Smyrna” really originates from that locality, a W Anatolian refugium,

indeed, would not be impossible. This refugium, however, seems rather improbable,

because (1) it does not explain that ssp. ponticus instead of ssp. melotis occurs at the

southern coast of Turkey, and (2) it would mean that ssp. melotis is the older form.

The latter suggestion is improbable, as the differentiating character of ssp. melotis

(mainly the white unh) does not seem primitive; or else, melotis has to be polytopic, for

which no evidence, neither in favour nor to the contrary, exists.

There is a much more trivial explanation for the occurrence of ssp. melotis on the

Island of Milo (? and W Turkey), viz. it may have been brought there by man as there

has been lively shipping between Syria and the Aegean region since the Phoenicians.

(b). With the available data it is difficult to explain the large transitional zone between

ssp. malvae and ssp. kauffmanni (Transbaicalia, Amur region), but there is no reason

to regard it differently than as a zone of postglacial secondary intergradation.

Apparently the differentiation which eventually led to the origin of the three semi-

species, took place in the Third Glacial. From the present distributional areas of the

semispecies we may conclude that P. malvoides originated in a West Mediterranean

refugium, P. melotis in an East Mediterranean refugium and P. malvae in an Asiatic

refugium.

Sichel (l.c.) observed that P. malvoides does not occur in NW Africa, on the Balearic

Islands and on Sardinia (and according to him, neither on Corsica). He concluded that

the semispecies arose rather late, but he did not give an exact period. Evidently, he

meant that the presence of P. malvoides in SW Europe of a rather recent time, but as

P. melotis is absent on Crete and Cyprus and P. malvae in Sakhalien and Japan, this

reasoning leads to the idea that the presence of superspecies malvae in the Palaearctic is

young (i.e. the superspecies originated in a recent time or it originated elsewhere and

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

colonized the Palaearctic in a recent time) or that superspecies malvae up to a rather

recent time only occurred in the interior of the Palaearctic. The only morphologic

character pleading for an origin outside the Palaearctic is the bipartite uncus, a unique

character among the Palaearctic Pyrgus species, but a normal feature in the American

ones. Without other points of evidence I will leave a possible Nearctic origin of super-

species malvae out of consideration (see also below). Furthermore, if there is any truth

in the assumption that P. malvoides and P. melotis originated in Mediterranean refugia,

during the Third Glacial, their precursor must have lived in the Mediterranean during

the Second Interglacial and the origin of superspecies malvae must date back at least as

far as the Second Glacial. As the difference between P. malvae and P. melotis is less

than between P. malvoides and the other semispecies (particularly in the lateral apophy-

ses), it is not improbable that the differentiation into a malvoides ancestor and a malvae-

melotis ancestor took place in the Second Glacial and thus the origin of superspecies

malvae must date from an earlier period.

So the absence of superspecies malvae from NW Africa and many islands cannot be

attributed to a recent origin of the semispecies or of the superspecies and the cause of

the absence must be looked for elsewhere, probably in some unknown ecological factor.

At present nothing can be added to the geographic history of superspecies malvae,

but a few words may be said on the evolutionary history. Where there exists a distinct

genital plate in the Pyrgus species, it is entire, except in malvae malvae in which it is

split into two parts. Therefore, the bipartite condition in malvae malvae undoubtedly has

been derived from an undivided plate as found in the semispecies malvoides and melotis

and in malvae kauffmanni.

A comparable situation exists in the male genitalia. The long processes of the lateral

apophyses found in P. me/otis and P. malvae probably evolved from a simpler type such

as found in P. malvoides. There is a gradual evolution in these processes, from the

slender and smooth type of P. melotis through the rather slender and spined type of

malvae kauffmanni to the coarse and heavily spined type of malvae malvae.

The same tendency is found in the style-stylifer-antistyle: a shortening and widening

from P. melotis (and P. malvoides) through malvae kauffmanni to malvae malvae.

Thus very probably, semispecies malvae is the most recent among the semispecies of

superspecies malvae. Of course, this does not mean that every character of semispecies

malvae is of recent origin, but it is very unlikely that a primitive character persisted only

in this semispecies. In this light the bipartite uncus of semispecies malvae seems a recent

character, independently originated from the bipartite uncus found in American Pyrgus

species; this disposes of the only argument in favour of an American origin of the super-

species malvae.

Warren (1957) considered the bipartition of the uncus a very old character, as he found

it also in Zerynthia, a genus of the Papilionidae which is supposed to be primitive.

However, comparison of the unaus of an arbitrary species of the Hesperiidae with that

of any species of the Papilionidae does not seem appropriate, as these families since long

are considered very distantly related (cf. Kiriakoff, 1960). Warren is undoutedly

correct in stating that the type of the genitalia found in ssp. kauffmanni (named corea-

nus by Warren) can be regarded as the primeval type of the semispecies malvae (given

specific rank by Warren). Just as the lateral apophyses and the style-stylifer-antistyle, the

uncus appears to change gradually from P. malvoides through P. melotis and malvae

R. DE JONG: Pyrgus in the Palaearctic region 61

kauffmanni to malvae malvae: in P. malvoides the uncus is usually apically incised, in

P. melotis it is sometimes partly bifid, in malvae kauffmanni the two parts of the uncus

lie close to each other, and in malvae malvae the uncus consists of two diverging parts.

Strangely enough, the linking of P. malvoides to P. malvae through ssp. kauffmanni is

regarded as improbable by Warren and he even states (1957: 373): “The formation of

the apophyses, so markedly different in malvoides, suggests that it has originated from

some entirely independent source.” Thus he ignores the known hybridization between

P. malvae and P. malvoides.

12.a. Pyrgus malvae Linnaeus

Linnaeus, 1758 — Systema Naturae, Ed. X: 485.

Type-locality: Aland Is.

Distribution. (Map 10). — Palaearctic. From Great Britain to Korea and from the

Mediterranean to approximately 65° N Lat. (In Finland). The most widely distributed

Pyrgus species (i.c. semispecies).

In Great Britain, throughout England and Wales, north to S Yorkshire (Warren,

1926; Ford, 1946); records from Ireland refer to misidentifications (Baynes, in litt.).

Throughout W, C, E and SE Europe. In Norway only in the south, in Sweden north to

approximately 64° N Lat., in Finland a little more to the north; for a detailed distribu-

tion map, see Nordström, Opheim & Valle (1955). In SW Europe replaced by P. mal-

voides, south of a line roughly running from Rochefort (W France, Charente Maritime)

through C France, via Lyon, south of the Lake of Geneva, via St. Maurice (Valais) to

Landquart (Grisons) and from there via Innsbruck curving to Istria (for references, see

Chapter 2.b and under P. malvoides). In Italy it is only known from Sappada in the

northeast (Verity, 1940).

Throughout the Balkans and Greece; see also under semispecies melotis. East of the

Bosphorus the distribution is poorly known; I saw specimens from Brussa (24 4 9),

Zonguldak (1 4 ) and Amasia (1 G ) (BM). From the last locality also melotis ponticus

is known, which form occurs much further west, near Ankara (Cubuk Baraj). This

makes the correctness of the label of this single P. malvae specimen from Amasia at least

questionable. De Lattin (1950) recorded a male and a female of P. malvae taken at

Sivrice in E Turkey and he stated: „Wie die Genitaluntersuchung ergibt, handelt es sich

um echte malvae, nicht um malvoides Elw. u. Edw.” (p. 326). Possibly, these specimens

belong to melotis ponticus, which sometimes has a partly bipartite uncus and also in

other respects resembles P. malvae much more than P. malvoides. However, De Lattin

also recorded a male of P. melotis from Sivrice of which he examined the genitalia. This

may indicate a variable population or a distributional overlap of ssp. melotis and ssp.

ponticus (in which case they cannot be considered conspecific) or of P. melotis and

P. malvae.

According to Alberti (1969), not yet found in the Caucasus (replaced there by melotis

ponticus), but north of the Black Sea only P. malvae occurs and not melotis ponticus. In

Russia widely distributed, at least as far north as Kotlas (61° N Lat.) (Krulikovsky,

1909).

Throughout Siberia; some northern localities are Antsiferovo on the Yenisey (59° N

Lat.) (Trybom, 1877) and the Vilui in NE Siberia (63°—64° N Lat.) (Herz, 1898).

Southward to Pamir (Groum-Grshimailo, 1890) and Korea (Alberti, 1955; Warren,

1957).

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Habitat. — Flowery places, on dry and on marshy ground; from sea-level to 2000 m

in the Alps and in Macedonia (own observation) and to 3000 m in Pamir (Groum-

Grshimailo, 1890). At least in Switzerland more common at low than at high altitudes

(Wiltshire & De Bros, 1966).

Biology. — Monovoltine; usually flying in May and June. In southern localities it

may be bivoltine, e.g. in Macedonia (Thurner, 1964); in the British Museum there is

a nice series of Hungarian specimens from every month from April to October. Accord-

ing to Kurentsov (1949), north of Vladivostok two broods, in May and August.

P. malvae hibernates as chrysalis (Forster & Wohlfahrt, 1955; Gullander, 1959; etc.).

Various species of Rosaceae have been recorded as food plants, belonging to the

genera Comarum, Fragaria, Potentilla, Rubus and Agrimonia (e.g. Verity, 1940; Berg-

mann, 1952; Forster & Wohlfahrt, 1955; Guillaumin, 1964). Some authors also mention

Coronilla (Papilionaceae) (Verity, Bergmann) and Dipsacus (Dipsacaceae) (Verity) as

food plants.

Geographic variation. — It is surprising that P. malvae, though individually highly

variable, doet not vary geographically from England to Lake Baikal. Only in the extreme

east of the distribution range, in the Sikhote-Alin range and Korea, a form occurs which

is obviously different by the genitalia and slightly by the facies. Transbaicalia and the

Amur region are populated by transitional forms.

Pyrgus malvae malvae Linnaeus

Material examined. — 729 § 372 9: 44 G 29 © England (BM, ZMA, RJ), 141

d 55 ® Netherlands (ML, RJ), 36 & 12 9 Belgium (BM, ML, ZMA, RJ), 2 &

Luxembourg (ZMA), 128 4 68 2 France (south to Charente Maritime, Cantal and

Savoie) (BM, ML, ZMA), 5 & 2 2 Norway (BM), 1 4 Sweden (BM), 6 g 4 2

Finland (BM, ML), 1 4 Denmark (BM), 65 & 50 © Germany (BM, ML, ZMA,

RJ), 19 & 6 © Switzerland (Geneva, Lausanne, Vaud, St. Gallen, Luzern, Zürich)

(BM, ML, RJ), 62 & 38 2 (N Tyrol, Carinthia, Austria Inf.) (BM, ML, RJ), 2 &

3 Q Istria (BM), 8 g 12 Q Czechoslovakia (BM, ZMA), 46 G 21 9 Hungary (BM,

ML, ZMA), 59 g 21 ® Yugoslavia (BM, ML, ZMA, RJ), 6 G 3 2 Bulgaria (BM,

ML), 4 & Rumania (ML), 25 4 8 ® Greece (Macedonia, Olympus, Parnassus) (BM,

ML), 29 & 10 © Bosphorus (BM), 4 g 4 2 W Turkey (Brussa, Zonguldak,

Amasia) (BM), 17 g 10 ® Russia (Leningrad to Novorossiisk and Uralsk) (BM),

1 f 1 9 W Siberia (Turgai, Kansk) (BM) 7 & 2 9 Altai (BM), 1 d E Sajan

(BM), 2 & Irkutsk (BM), 10 4 3 9 Turkestan (Alexander Mts to Alai) (BM, ML).

Distribution. — The range of semispecies malvae, eastward to Lake Baikal.

Subspecific characters. — Differs from ssp. kauffmanni in the male genitalia by the

two parts of the uncus which are set obliquely to the tegumen, by the coarser and more

strongly curved processes of the lateral apophyses and by the broader style-stylifer; in the

female genitalia by the genital plate which is divided into two parts. The differentiation

in the facies is slight, see ssp. kauffmanni.

Variation. — Apart from many forms of individual variation two forms have been

described as subspecies:

elegantior Verity, 1934. — Large, 11—12 mm; above a less black, warmer tinge,

spots smaller, particularly the submarginal spots; unh ground colour more brownish or

reddish than greenish, veins outlined in yellow instead of white. According to Verity

R. DE JONG: Pyrgus in the Palaearctic region 63

(1947) and Kauffmann (1951) this form occurs in the west of Switzerland and in

France south of the Lake Geneva. However, specimens agreeing with the above descrip-

tion also occur in S Germany and, particularly, in Yugoslavia, where they fly with other

forms. Perhaps an ecophenotypic variation.

luciarius Kauffmann, 1951. — Large, 11.7—12.8 mm; above ground colour very

black with large spots and many whitish hairs. Unf black with strongly contrasting white

spots; uph ground colour ochreous-brown to chocolate-brown or dark grey, rarely

reddish, veins little contrasting.

According to Kauffmann this form occurs in C Switzerland. It is, however, not at all

confined to that country but it occurs as an individual variation throughout C, E and SE

Europe. I have seen specimens of this form taken as far north as the Netherlands, but

almost all these examples are smaller and still further north the specimens are invariably

smaller than /ciarius.

A striking individual form is faras Bergsträsser, 1780. In this form the spots upf are

strongly enlarged and partly coherent, while the spots uph are reduced. Extremes of this

variation from W. Germany have more than half of the upperside of the fore wings white,

only intersected by brown veins (PI. 4 fig. 61, 70).

Transitional zone: Transbaicalia and Amur region. According to Warren (1957) this

region is populated by forms transitional between ssp. malvae and ssp. kauffmanni. I

have examined the genitalia of only five males from this region. Indeed, they turned out

to be somewhat intermediate, though more resembling ssp. malvae than ssp. kauffmannt.

With regard to the facies I could not distinguish these and other specimens from this

region from ssp. malvae.

Material examined. — 11 d 11 2:6 g 2 9 Chita, 3 g 7 Q Jablonowyj Khrebet

(= Apfelgebirge), 2 3 2 9 Amur (all BM).

Pyrgus malvae kauffmanni Alberti

Alberti, 1955 — Zeitschr. Wiener Entom. Ges. 40: 43—46.

Type-locality: ““Amur Radde”.

Material examined. — 9 G 1 9:9 8 Tjutju-ho (400 km N of Vladivostok), 1 9

Amur (all BM).

Distribution. — Presumably confined to the Ussuri region and Korea. Data on the

distribution in Korea are very scarce; Alberti (l.c.) mentions Seishin (N Korea); Warren

(1957) described the same form from Hakugan under the name coreanus.

Subspecific characters. — Male genitalia: the two parts of the uncus not diverging as

those parts in ssp. malvae; processes of lateral apophyses more slender, less strongly

curved and less strongly spined; style-stylifer more slender. Female genitalia: genital

plate entire, very much like the plate found in P. malvoides. Unh ground colour more

evenly brownish, spots often reduced. I have seen the same unh in some specimens of

melotis ponticus.

Geographic history. — See above, under the history of superspecies malvae.

12.b. Pyrgus malvoides Elwes & Edwards

Elwes & Edwards, 1897 — Trans. Zool. Soc. Lond. 14: 160; pl. 23, figs. 27, 27-a.

Type-locality: Biarritz.

Distribution (Map 10). — SW Europe. Throughout Spain and Portugal (Manley &

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Allcard, 1970). In France north to the following departements: Charente Maritime, Gi-

ronde (northern part), Dordogne, Lot, Corrèze, Cantal, Puy-de-Dôme, Loire, Rhône,

Isère, Savoie, Haute Savoie (Picard, 1948a; Guillaumin, 1964). In Switzerland confined

to Valais, Ticino and Grisons (Kauffmann, 1951). In Austria confined to the western

part (Otztaler Alpen; Innsbruck) (Kauffmann, 1955).

Habitat. — As that of P. malvae. At least in Switzerland (Valais) it is most common

at 1500 m and higher, while P. malvae is more common at lower altitudes (Wiltshire

& De Bros, 1966). In the Alps it has been collected up to 2500 m (Ticino; Verity,

1940), in the Pyrenees up to 2000 m (Gavarnie; own observation).

Biology. — Bivoltine; in Spain flying from March to September at low altitudes

(Manley & Allcard, 1970), in France from the middle of May to the middle of June

and from the middle of July to the middle of October, depending on altitude (Guil-

laumin, 1964). In the Alps (and Pyrenees?) at higher altitudes probably single-brooded

(Kauffmann, 1950-a).

It hibernates in the pupal stage (Kauffmann, 1954c; etc.).

Kauffmann (1950a, 1954c) reared the larvae and recorded as food plants: Fragaria

vesca, Potentilla verna and P. erecta (Rosaceae).

Geographic variation. — As in P. malvae various forms, apparently ecophenotypic

variations and differing in size, spotting and colour, have been described as subspecies,

while two geographically defined forms can be distinguished on the basis of the male

genitalia. Sichel (1961, 1962b) was the first to recognize the variation in the genital

armature. He found a nearly constant difference between specimens from Peninsular Italy

and Sicily (34 males) and specimens from the remaining part of the range (9 males

from Switzerland, 41 males from N Italy, 23 males from France, 12 males from Spain

and 8 males from Portugal). I have examined the genitalia of 15 males from peninsular

Italy, 5 males from N Italy, 20 males from Switzerland, 20 males from SE France and

the Pyrenees, and 5 males from Spain. I can only support the findings of Sichel and I

therefore distinguish the subspecies malvoides and modestior. These subspecies also

differ to a certain extent in the ground colour unh, but there is much overlap (see

below).

Pyrgus malvoides malvoides Elwes & Edwards

Material examined. — 385 & 232 9:16 4 14 9 Portugal (BM), 63 & 28 2

Spain (BM, ML, ZMA, HC, RJ), 94 & 54 Q French Pyrenees (BM, ML, ZMA, HC,

RJ), 123 4 86 Q France (Landes, Gers, Aveyron, Puy-de-Dôme, Bouches-du-Rhône,

Var, Alpes Maritimes, Basses Alpes, Hautes Alpes, Dröme, Isere, Savoie, Haute Savoie)

(BM, ML, ZMA, RJ), 49 g° 31 9 Switzerland (Valais, Ticino, Grisons) (BM, ML,

ZMA, RJ), 39 & 16 9 N Italy (BM, ML, HC, HO, RJ), 1 g 3 2 Istria (BM, HS).

Distribution. — That of the semispecies except peninsular Italy and Sicily.

Subspecific characters. — Male genitalia: distal edge of cuiller more strongly curved

than in ssp. modestior; the continuation of this curve intersects the dorsal ridge of the

harpe or passes well above it. External characters: Ground colour unh in both broods

usually of a warmer and more reddish tinge than in ssp. modestior but in mountainous

regions often ochreous or brown, sometimes greenish, particularly in the first brood.

Variation. — In Spain and France the male genitalia show little variation, but in

Switzerland and N Italy traits of ssp. modestior are apparent in the genitalia of some

males. Sichel (1962b) found the curvature of the cuiller in 14 % and the continuation

R. DE JONG: Pyrgus in the Palaearctic region 65

of the profile of harpe and cuiller in 7 % of the males examined from Switzerland and

N Italy to be of the modestior type, but he does not mention whether he found both

characters in the same specimens.

Of 25 males from Switzerland and N Italy examined I found two specimens which

showed both characters of ssp. modestior and two specimens which showed one. Thus it

seems advisable to list the Swiss and N Italian populations as ssp. malvoides. The

material examined is insufficient to establish how ssp. malvoides and ssp. modestior

pass into each other, but from Emilia southward malvoides traits in the male genitalia

appear to be very rare (see ssp. modestior).

For forms described on the basis of differences in size, colour and extension of the

white spots is referred to Verity (1940, 1947) and Kauffmann (1951). Only two forms

are mentioned here:

(1) pyrenaicus, Tutt, 1906, has given some nomenclatural difficulties. It was described

as an individual variation taken in the Pyrenees. Warren (1926) gave the Pyrenean form

of Pyrgus carthami the same name. Verity (1928) therefore thought it necessary to

replace this name by microcarthami, but Warren (1929: 149) pointed out clearly that

“the fact that there happens to be an ab. pyrenaica, Tutt, of another species in the genus

in no way invalidates my race pyrenaica”. It is surprising to find the name microcarthami

as a subspecific name for N Spanish malvoides in Manley & Allcard (1970).

(2) semiconfluens Reverdin, 1911. An individual variation comparable with the form

taras in P. malvae, viz. spots upf very enlarged and more or less confluent. I have not

seen such extreme specimens as mentioned under P. malvae.

Pyrgus malvoides modestior Verity

Verity, 1929 — Trab. Mus. Ci. Nat. Barcel. 11: 7.

Type-locality: Tuscany.

Material examined. — 53 d 25 9:41 8 17 9 Peninsular Italy (Bologna to Cala-

bria) (BM, ZMA, HC, RJ), 12 8 8 @ Sicily (BM).

Distribution. — Peninsular Italy and Sicily.

Subspecific characters. — Male genitalia: distal edge of cuiller less narrowly curved;

the continuation of this curve coincides with the dorsal ridge of the harpe.

External characters: the specimens of the first generation as in ssp. malvoides, but

rarely with the warm reddish tinge unh often found in ssp. malvoides. The specimens of

the second generation usually with a paler unh than specimens of ssp. malvoides, but

otherwise as variable as in that subspecies.

Variation. — As stated above, males with malvoides-like genitalia are rare in Penin-

sular Italy and Sicily. I saw one male taken at Pistoia (Tuscany) with the harpe-cuiller

profile of the malvoides type and one male from Calabria with both the curvature of the

cuiller and the harpe-cuiller profile of the malvoides type.

An interesting individual variation is the form which has been named melotiformis

by Verity (1929): ground colour unh extremely pale, recalling P. melotis.

Geographic history. — Possibly the plains of the Po have retarded the postglacial

extension of ssp. modestior towards the north. Evidently ssp. modestior reached N Italy

and Switzerland not earlier than ssp. malvoides. The data available are not yet conclusive,

but it can be supposed that ssp. modestior and ssp. malvoides came into a postglacial

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

contact in the southern Alps, forming a hybrid population which extended its range

into the central Alps.

See further above, the history of superspecies malvae.

12.c. Pyrgus melotis Duponchel

Duponchel, 1832 — Lép. France; Suppl. 1: 251; pl. 42 figs. 1, 2.

Type-locality: Island of Milo.

Distribution (Map 10). — East Mediterranean. W Anatolia to the Caucasus (Alberti,

1969) and the Jordan Valley (Hemming, 1932; BM); Milo Is. The distributional limits

are badly known. The westernmost localities from where I have seen specimens of this

species are Gubuk Baraj (north of Ankara), Aksehir, Egerdir and Antalya. According

to Alberti (1969), it is widely distributed in Transcaucasia, but apparently there are no

records of its occurrence in Iraq and Iran.

Habitat. — Not recorded but presumably as that of P. malvae. From 250—275 m

below sea level (Jordan Valley; Hemming, 1932) up to 2600 m (Caucasus; Alberti,

1969). In the Lebanon very common up to + 1700 m (Nicholl, 1901). In Israel (and

possibly elsewhere) not in the Eremian regions (Amsel, 1933).

Biology. — Bivoltine. Ssp. melotis may have a third brood (Graves, 1925; Hemming,

1932). According to Milyanovsky (1964) also in Abkhaziya a third brood.

Food plants unknown.

Geographic variation. — There is much confusion about the taxonomic status of the

forms ponticus and melotis (see also Chapter 2.b). They are usually considered specific-

ally distinct, evidently without any other reason than the fact that they differ with regard

to the facies and the genitalia. The difficulties are partly or entirely due to the scarcity

of ssp. ponticus in collections, e.g. in the British Museum there are only 14 gf and 9 9.

I could examine rather numerous specimens with many transitional forms between ssp.

ponticus and ssp. melotis. As stated above (Chapter 2.b), no constant differences were

observed in the male genitalia. In both ssp. ponticus and melotis these are rather variable;

in ponticus this variation may be partly geographic, see below. The facies is very

variable in both forms; the most important, if not the only, difference, is the colour unh;

in ssp. melotis the unh is partly to entirely white, with very faint spots, in ssp. ponticus

the unh is usually brownish-ochreous or greenish with well-defined spots. However,

intermediate forms occur, especially in N Syria and S Turkey.

Pyrgus melotis melotis Duponchel

Material examined. — 79 4 26 9:4 gd 1 9 “Greece” (BM), 1 g “Smyrna”

(BM), 1 9 Antakya (BM), 70 8 23 Q Syria (BM, ML), 4 Lebanon (BM, ML,

RJ), 1 g 1 ® Jordan Valley (BM). According to Evans (1949) the specimens labelled

“Greece” in the British Museum probably originate from Milo Is.

Distribution. — N Syria to the Jordan Valley. Presumably also further north: De

Lattin (1950) recorded this subspecies from Gaziantep and from Sivrice, which is much

further north.

Subspecific characters. — Unh white or with white streaks, spots faintly visible.

Variation. — Already Graves (1925) noted the great variation in this form with

regard to the distinctness of the spots unh, the size of the spots above and the size of

R. DE JONG: Pyrgus in the Palaearctic region 67

the butterfly. I have seen males from N Syria measuring 10.9 to 15.0 mm! Hemming

(1932) described the “subspecies” jordana from the Jordan Valley, characterized by its

large size, large spots above and a very white unh. Evidently, Hemming did not know

that these characters do occur in other areas of distribution of ssp. melotis as well (he

only compared his jordana with specimens of ssp. melotis from Lebanon).

The unh varies from white to nearly ponticus-like; some specimens from N Syria are

difficult to distinguish from ponticus.

Pyrgus melotis ponticus Reverdin

Reverdin, 1914 — Bull. Soc. Lép. Genéve 3: 66—72; pl. 3 figs. 6, 12; pl. 4 figs. 2, 3.

Type-locality: Amasia (Turkey).

Material examined. — 106 4 26 9 Turkey (Antalya, Egerdir, Aksehir, Cubuk Baraj,

Amasia, Kedos, Yozgat, Gümüshane, Seki Yaila, Boz Dagh, Mersin) (BM, RJ).

Distribution. — The range of the semispecies except that of ssp. melotis, i.e. north

of Syria.

Subspecific characters. — Unh not white or with white streaks; brownish, ochreous or

greenish with well-defined spots.

Variation. — g' 10—14 mm. The specimens from the Boz Dagh (18 g' 2 9) are

rather large (12—14 mm), dark above (with very little light superscaling), spots uph

reduced. Most specimens from Aksehir (75 g 14 ®) are smaller and much more

extensively spotted, but some are indistinguishable from the examined specimens from

the Boz Dagh.

I studied the genitalia of 25 males from Aksehir and 15 males from the Boz Dagh.

All specimens from Aksehir appeared to possess an entire uncus while all specimens

from the Boz Dagh showed a partly bifid uncus. With more material available from

various parts of the range of ssp. ponticus, a marked geographic variation may be proved

to exist.

The single specimen (4) from Mersin has a pale ground colour unh, indicating the

influence of ssp. melotis.

Geographic history. — See above, the history of superspecies malvae.

13. Pyrgus serratulae Rambur

Rambur, 1839 — Faune entom. de |’ Andalousie: pl. 8 figs. 9, m.

Type-locality: Spain.

Distribution (Map 11). — From Spain to Transbaicalia; presumably continuously

distributed from France to Transbaicalia, but isolated populations, at least in Europe,

occur north and south of the main range.

In Spain presumably confined to the larger mountain regions: from the Pyrenees and

the Cantabrian Mountains south to the Montes Universales and Sierra de Gredos, and

probably discontinuous in Sierra Nevada (Warren, 1926; Manley & Allcard, 1970;

material examined).

In Italy also confined to the larger mountain regions: the Alps, Sibillini Mts., Abruzzi

and Monte Meta (Verity, 1940).

Throughout France, except the northwestern part. In W Germany northward to a

line approximately from Koblenz to the Harz, in E Germany through the valley of the

Saale southward (Alberti, 1927; Bergmann, 1952). Further north found in Mecklenburg

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

and Pommern (Urbahn, 1939; Friese, 1956); two specimens have been taken in Den-

mark (Wolff, 1939), but were probably of foreign origin (Opheim, 1956). Throughout

C, E and SE Europe. From S Greece (Taygetos Mts.) only recorded by Rebel (1902).

Throughout Turkey; south towards the Lebanon (BM), southeast through the moun-

tains of Iraq and the Zagros Mts. to Fars (Wiltshire, 1957; Brandt, 1939). According

to Romanoff (1884) south of the Caucasus widely distributed, but the distribution in

N Iran is almost unknown, as the species is recorded by Alberti (1957) only from NE

Iran.

In the Caucasus widely distributed (Wojtusiak & Niesiolowski, 1947; Alberti, 1969).

Few records of the occurrence in S Russia: Gniliza (east of Charkow) (Alberti & Soff-

ner, 1962), Crimea (Korshunov, 1964) and Sarepta (BM). In E Russia known from

Uralsk, Guberli (BM), and Kasan (Krulikowsky, 1908). The records from Vologda

and Kotlas (Krulikowsky, 1909) probably concern P. alveus.

East of the Urals possibly continuously distributed, but only recorded from the

Tjumen district (Slastshevsky, 1911), the Altai (Elwes, 1899), N Mongolia (Forster,

1965) and Jablonowyj Khrebet (= Apfelgebirge) (BM). Herz (1898) recorded the

species from Vilui in NE Siberia, but it may be a misidentification for P. alveus.

Habitat. — More or less flowery places, clearings, meadows, etc., from sea level

(W France, N Germany) up to 2400 m in the Spanish Pyrenees (Gerona; HC) and still

higher in the Alps (Kauffmann, 1951); in Iraq from 1300 to 2000 m, in S Iran (Fars)

above 2600 m (Brandt, 1939), in the Altai up to 2300 m (Elwes, 1899). Often in large

numbers at damp places (mud and dung).

Biology. — Monovoltine; flight period, four to five weeks, date of emergence

strongly influenced by altitude, in the lowlands from the middle of May, in Switzerland

at 100 m from June, at high altitudes often from the middle of July (Warren, 1928;

Kauffmann, 1951). The altitude appears to be more important for the flight period than

the latitude: in N Germany adults from the middle of May to the middle of June

(Friese, 1956). In Iraq bivoltine, flying in May and from July to October (Wiltshire,

1957).

The species hibernates in the larval stage (Koch, 1963).

Various species of Potentilla and Alchemilla (Rosaceae) have been recorded as food

plants (Verity, 1940; Bergmann, 1952; Guillaumin, 1964). In addition, Verity mentioned

Aira montana (Gramineae).

Geographic variation. — The sensitiveness to local conditions appears not only in the

strong delay in the date of emergence in the mountains, but also in the variation in size,

colour and design. This has led to the distinction of many ‘‘subspecies’’ even in cases

where it is clear that the supposed subspecies is a climatic, non-hereditary form. Thus

Kauffmann (1951: 349) says: “Die Zentraleuropäische Verhältnisse zwingen mich,

neben den von Evans aufgestellten Unterarten noch die f. planorum Verity als Subspezies

hinzuziehen” and he recognizes the subspecies serratulae, caecus and planorum in Swit-

zerland. However, one page further he admits that planorum may be a climatic form:

“Als Form ist planorum überall in der Schweiz zu finden mit anderen Formen vermischt,

reichlicher in der Ebene und spärlicher im Gebirge. Sie ist wahrscheinlich die somatische

Form von trockenem Klima” and he speaks of caecus in a similar manner. Even the great

lumper Evans (1949) thought it justified to recognize the subspecies caecus. As stated

in Chapter 1.b it is not desirable to call climatic forms subspecies and therefore I

recognize only a single subspecies in SW and C Europe.

R. DE JONG: Pyrgus in the Palaearctic region 69

In the eastern part of the range problems have arisen, due to the misleading lumping

by Evans (1949) of different forms into the subspecies wralensis, and due to the abuse

of the name major by various authors for forms which have little to do with the true

major from Turkey (e.g. Higgins & Riley, 1970: 316—317: “Size varies, large speci-

mens, f. major Staudinger Pl. 58 with fw 15/16 mm occur in lowlands in SW France

and elsewhere (Vendée, Gironde, Charente, etc.) and in Balkans (esp. Greece)’’).

The following classification may be justified from the phylogenetic point of view.

Pyrgus serratulae serratulae Rambur

Pl. 5 Fig. 73—76, 82—85

Material examined. — 702 4 276 9: 149 & 49 9 (Sierra Nevada and from Montes

Universales and Sierra de Gredos to the Pyrenees) (BM, ML, ZMA, HC, RJ), 4 &

3 © Andorra (ZMA), 146 G 65 ® French Pyrenees (Basses Pyrénées to Pyrénées

Orientales) (BM, ML, ZMA, RJ), 179 8 60 Q France (Alpes Maritimes, Basses

Alpes, Hautes Alpes, Savoie, Cantal, Puy-de-Dôme, Ille-et-Vilaine, Auvergne, Vendée,

Charente, Eure, Aube, Lozére, Seine-et-Marne, Bas-Rhin, Marne) (BM, ML, ZMA, RJ),

105 8 39 9 Switzerland (Valais, Berne, Uri, Ticino, Grisons) (BM, ML, ZMA, RJ),

45 g 23 @ Italian Alps (BM, ML, RJ), 9 & 3 2 C Italy (BM, ZMA), 35 g 20 9

Austria (Tyrol to Mödling) (BM, ML, ZMA, RJ), 14 8 7 9 W Germany (northward

to Niedersachsen) (BM, ML, RJ), 5 g 1 2 Czechoslovakia (BM), 8 & 4 9 Hun-

garia (BM), 1 g 1 9 Banat (ML), 2 g' 1 @ Bosnia (BM, ML).

Distribution. — W and C Europe; the zone of contact with the major subspecies group

is not exactly known.

Subspecific characters. — Highly variable and, therefore, difficult to define. Small to

large, d' 10.8—14.6 mm. Above, usually less spotted than in other subspecies. Below,

spotting variable, usually less spotted than in other subspecies, unh the median spots 2

and 3 and the submarginal spots 3 and 7 usually faint or absent, rarely prominent.

Rarely with clear white borders unf and unh. Colour unh dark olive-green to warm

yellow.

Variation. — As stated above, highly variable according to local conditions. Some

obvious variations are:

Forma caecus Freyer: small, 10.8—12.4 mm; above, spots very reduced, may be

absent in female; below, colour faded, unh greyish-greenish with unsharp markings. This

form is found almost everywhere above 2000 m in the Alps, together with specimens of

the nominate form from which it cannot be well distinguished, as there seems to exist a

continuous variation in the extension of the spotting. The ratio caecus: serratulae changes

from year to year and depends on the climate (Kauffmann, 1951). It is, however, not

confined to the Alps. Bergmann (1952) mentions the form from several localities in

Thiiringen (he attributes it to a cold and damp climate); other records are e.g. Caradja

(1895): Poland. In addition, I saw some (rare) specimens from the Pyrenees.

Forma serratulae Rambur: somewhat larger, 9 11.6—13.2 mm; upf all spots of

median row present, uph plain dark brown or more or less clearly spotted; unh cold

greenish with clearly defined, somewhat reduced spots. Found in all mountain regions,

varying towards f. planorum which is usually more common at lower altitudes.

Forma planorum Verity: slightly larger than serratzlae, in Switzerland 12.8—13.7 mm

(Kauffmann, 1951), but in France variable, may be as large as f. magnagallicus. Above,

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

spotted as in f. serratulae; unh various shades of a warm, yellowish colour; spots unh

often more or less isolated and slightly rounded. Most numerous in the lowlands, but

occurs everywhere with f. serratulae.

Forma magnagallicus Verity (= occidentalis Lucas). The specimens may be large, 3

to 14.5 mm, in the northwestern part of the distributional area in France, from Vendée

to Seine-et-Marne and Aube, otherwise like f. planorum. The variation from f. planorum

to f. magnagallicus appears continuous; the males captured at Fontainebleau (Seine-et-

Marne) vary from 13.2 to 14.5 mm.

The forms diniensis Heinrich and arvernensis Picard are not clearly defined local

forms which fall within the variation of subspecies serratulae.

A large, planorum-like form, 4 12.8—13.9 mm, occurs in the Sierra de Gredos, upf

well-spotted, unh spots reduced and colour of a strange, deep yellow shade.

major subspecies group

Evans (1949) united all forms east of Greece into a single subspecies, wralensis.

He did not use the older name major Staudinger, 1879, as this name is preoccupied by

major Fabricius, 1787 (= Pyrgus carthami). Warren (1926) described zralensis from

two males and one female from Uralsk. I have seen nine males and four females from

SE Russia. Although this is not a large number, I believe that zralensis is possibly a

transitional form between the Turkish form (named major by Staudinger) and the

Siberian form. But if the Ural and the Turkish forms are not identical, the Turkish form

needs a new name. As it is not my intention to propose new names in the present paper,

I only call attention to this case, further using the name major Staudinger.

Much less material is available concerning this subspecies group than concerning ssp.

serratulae. Although there are clear-cut differences within the major subspecies group, it

is possible that with more material becoming available, a part of the variation may prove

to depend on climatic conditions only. Therefore, the following classification is provi-

sional.

The group differs from ssp. serratulae by a usually large size, more extensive spotting

upf, clearly defined white borders below, unh median row spots 2 and 3 and submarginal

spots 3 and 7 usually well-developed. The spots of the median row unh isolated by

ground colour along the veins and ground colour unh often greyish-powdered.

Usually, the specimens can easily be distinguished from specimens of ssp. serratulae,

but the major group shows the same degree of variation as ssp. serratulae.

Pyrgus serratulae balcanicus Warren

Pl. 5 Fig. 77, 86

Warren, 1926 — Trans. ent. Soc. Lond. 74: 97—98; pl. 29 figs. 7—12.

Type-locality: Cetinje (Montenegro).

Material examined. — 24 g 4 9:1 9 Montenegro (Cetinje) (BM), 13 & 1 2

Yugoslavian Macedonia (BM, HC, RJ), 11 4 2 9 Olympus (BM, ML).

Distribution. — Macedonia and Montenegro.

Subspecific characters. — Size variable, Yugoslavian Macedonia g' 13.7—14.8 mm,

Olympus 3 14—15.5 mm. Upperside dark brown with little light superscaling (may

be absent), upf spots well-developed, uph spots faintly visible. Unf dark grey or black,

R. DE JONG: Pyrgus in the Palaearctic region 71

white border narrow or obscure; unh dark brownish yellow with usually well-developed

spots (on the average less extensive than in subspecies major) which are clearly isolated

by the ground colour along the veins; white border unh narrow or obscure.

Note. — The name infraobscuratus Verity (1938) (type-locality: Mt. Olympus) is clearly a

junior synonym of balcanicus. For some obscure reason Evans (1949) considered balcanicus a

synonym of serratulae and infraobscuratus, a synonym of uralensis.

Pyrgus serratulae major Staudinger

Pl. 5 Fig. 78, 87

Staudinger, 1879 — Horae Soc. Entom. Ross. 14: 292.

Type-locality: Asia Minor.

Material examined. — 51 & 13 9: 2 g 1 2 Hungary (BM), 5 q Bulgaria

(Sliven) (BM), 2 & Parnassus (BM), 22 & 5 $ Turkey (BM, RJ), 1 9 Kasikoparan

(BM), 20 8 6 @ Syria (Akbès, Aintab) (BM).

Distribution. — From Greece and Bulgaria to Syria and Armenia.

Subspecific characters. — Large, 4 to 16.1 mm. Upf with well-developed spots, often

more extensive than in ssp. balcanicus, and with rather slight superscaling; uph spots

rather faint; unf light to dark grey with obvious white border; unh spots large, isolated by

ground colour along the veins, ground colour pale olive-greenish or yellowish (in worn

specimens darker, owing to absence of light superscaling), in Syria sometimes dark

brown, white border usually conspicuous, sometimes very narrow.

Variation. — The Hungarian specimens are about intermediate between ssp. major

and ssp. balcanicus. As also ssp. serratulae is known from Hungary, more recent and

more exact data are wanted.

The two males from Mt. Parnassus have less extensive spots unh and white border

unh very narrow, thus resembling ssp. balcanicus.

Pyrgus serratulae alveoides Staudinger

Staudinger, 1901 — Cat. Lep. Pal.: 96.

Type-locality: Syria.

Material examined. — 7 & 2 ® Lebanon (BM).

Distribution. — Lebanon and through the mountains of Iraq and the Zagros Mts.,

southward to N Fars (Wiltshire, 1957).

Subspecific characters. — Small, $ 12.4—13.3 mm; upf and uph strongly spotted

and with obvious white superscaling; unf and unh with broad white border; spots unh

variable, reduced to strongly developed; ground colour unh pale yellow or pale ochreous

green. Ssp. alveoides differs in the male genitalia from other forms of the species in

having a much more elongated cuiller, but transitional specimens also occur.

Pyrgus serratulae kotzschi Alberti

Pl. 5 Fig. 79, 88

Alberti, 1957 — Mitt. Deutsch. Entom. Ges. 16: 27—28.

Type-locality: Kuh i Mirabi (NE Iran).

Material examined. — 2 8 Kuh i Mirabi (paratypes) (BA).

Distribution. — Only known from the type-locality. According to the original descrip-

tion, a similar form occurs in the Shahkuh Mts.

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Subspecific characters. — Large, S to 16 mm; above, spots small, uph faintly visible;

below greyish; unh spots very large and more connected than in ssp. major, ground

colour faded grey-olive; white borders below, obvious.

Pyrgus serratulae grisescens Alberti

Pl. 5 Fig. 80, 89

Alberti, 1969 — Faun. Abh. Staatl. Mus. Tierk. Dresden 2: 141—142; figs. 3a—b.

Type-locality: Teberda (Caucasus).

Material examined. — 6 g': 3 4 Teberda (NW Caucasus, 1700 m; paratypes), 3 d'

Passanauri (Georgia, 1700 m) (BA).

Distribution. — Confined to the Caucasus.

Subspecific characters. — Size as in the Central European ssp. serratulae, & 13.5 mm,

above also recalling ssp. serratulae, but unh the large spots are major-like, although they

are not as sharply isolated as in that subspecies; ground colour unh a peculiar shade of

faded greenish grey (in worn specimens the colour may be darker owing to the absence

of superscaling with light scales); white borders below narrow or absent.

Pyrgus serratulae uralensis Warren

Pl. 5 Fig. 81, 90

Warren, 1926 — Trans. ent. Soc. Lond. 74: 98; pl. 29 figs. 1—6.

Type-locality: Uralsk.

Material examined. — 9 G 4 9:1 9 Sarepta, 2 £ Guberli, 5 g' Ural (all BM),

2 3 @ Uralsk (BM, ML).

Distribution. — SE Russia; but see also below, under Siberian form.

Subspecific characters. — Smaller than the Turkish ssp. major, 9 13—14.8 mm.

Upperside and underside very much like ssp. major, the females indistinguishable from

that subspecies, but the males variable: unh spots large and isolated to reduced and more

coherent, a clearly defined white border along termen or the ground colour extending

nearly or entirely to the termen. Upperside often more extensively spotted than ssp.

major, particularly the spots uph may be conspicuous.

Note. — Although hampered by the limited number of specimens available, I still am

of the opinion that wralensis is a hybrid population between the SE European and SW

Asiatic form (ssp. major) and the Siberian form, some specimens being very much like

ssp. major, other specimens being almost indistinguishable from specimens from the

Altai. Therefore, I believe that the use of the name zralensis for the subspecies group is

undesirable. I even doubt the usefulness of the subspecific distinction of wralensis, but

the name can be provisionally used to indicate the hybrid population.

Pyrgus serratulae — Siberian form

The Siberian populations of P. serratulae are poorly known. These are listed by Evans

(1949) and Forster (1965) as ssp. wralensis. This attribution is not strange in view of

the variation of ssp. wralensis described above, but on the whole, Siberian specimens

show so little resemblance with subspecies of the major group, that they should be

R. DE JoNG: Pyrgus in the Palaearctic region 73

excluded from the major group and be considered to belong to a separate subspecies.

This Siberian form will remain unnamed for the present in view of the limited number

of specimens available.

Material examined. — 9 ¢ 5 9:8 g 12 Altai (Ongodai, Tchuja Valley), 1 ¢

4 9 Jablonowyj Khrebet (= Apfelgebirge) (BM).

Distribution. — Presumably from the Urals to E Siberia.

Subspecific characters. — Altai $ 13—14.3 mm, Jablonowyj Khrebet & 12.7 mm.

Upperside more extensively spotted than ssp. major, particularly uph, but less than ssp.

alveoides; unh recalling ssp. wralensis, spots not as reduced as in some specimens of ssp.

uralensis, hardly or not at all intersected by the ground colour.

Geographic history. — Although the material available from Siberia is very limited,

there seems to exist little variation, particularly in comparison with the situation in SE

Europe and W Asia. This suggests that the occurrence of the species in Siberia is of a

more recent age than the occurrence in SE Europe and W Asia, also because of the

absence of closely related species in Asia. The slight variation in Siberia also points to

the assumption that the Siberian form arose from a single, geographically isolated

population in S Siberia, possibly during the Last Glacial. The W Siberian lowlands may

have become inundated during the Last Glacial (cf. Moreau, 1955), in this way isolating

the Siberian populations from the more western ones. Thus the geographically as well

as morphologically intermediate position of ssp. wralensis becomes understandable, as

these populations apparently met postglacially, giving rise to a zone of secondary inter-

gradation.

Ssp. major evidently originated in a refugium in SE Europe or SW Asia during the

Last Glacial. It is at present impossible to be exact on the extension of this refugium,

whether it was restricted to Asiatic Turkey or also included Greece (and S Russia?), but

apparently it was situated north of the region where ssp. alveoides originated. There is

no reason to date the origin of ssp. alveoides further back than in the Last Glacial or at

best in the Third Interglacial. In a Glacial (Pluvial) rather than in an Interglacial Period

it may have had ample opportunity to extend its range, but I rather do not infer the

history of ssp. alveoides without having seen specimens from the Zagros Mts.

Ssp. kotzschi is possibly an extreme, postglacially developed form of ssp. major, but

the entirely unknown situation in NW and N Iran makes a reliable conclusion im-

possible. Likewise, ssp. grisescens may be a postglacial descendant from ssp. major, but

it may equally well have originated in an isolated area during the Last Glacial.

The characters of ssp. balcanicus seem to be too constant to conclude that this sub-

species is only a product of postglacial hybridization of the subspecies major and serratu-

Jae. Evidently it had a refugium of its own during the Last Glacial.

Subspecies serratulae presumably arose in a refugium in Spain and (or) Italy, during

the Last Glacial. From the present geographic variation it is not possible to infer that

the subspecies occurred in W Europe during and before the Third Glacial, but of course

this is not impossible.

So we cannot trace the history of the species far back. It is interesting to note that the

species is evidently a western immigrant in Siberia.

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

14. Pyrgus onopordi Rambur

Rambur, 1839 — Faune entom. de l'Andalousie: pl. 8 fig. 13, p.

Type-locality: Granada.

Distribution (Map 12). — NW Africa and SW Europe. In NW Africa widely

distributed, from S Morocco to E Algeria in High Atlas, Middle Atlas and Tell Atlas

(BM, RJ). Throughout Portugal and Spain, but absent in NW Spain (Agenjo, 1952).

The species is widely distributed throughout S France: I have seen specimens or reliable

records from the following departments: Basses Pyrénées (HC), Hautes Pyrénées

(Verity, 1947; BM), Pyrénées Orientales (Dufay, 1961), Tarn (RJ), Lot (BM),

Charente (BM), Vienne (BM), Gard (Picard, 1948c; Gaillard, 1952), Bouches-du-

Rhône (Verity, 1947; BM, ML), Vaucluse (ZMA), Var (BM), Alpes Maritimes

(Verity, 1947; BM, RJ), Basses Alpes (Verity, 1947; various collections), Hautes Alpes

(BM, RJ), Savoie (BM). Throughout Continental and Peninsular Italy (Verity, 1940);

recorded by Schwingenschuss (1942) from Sicily, (possibly a misidentification); ap-

parently absent from Corsica and Sardinia. In Switzerland only in Valais and the Grisons

(Kauffmann, 1951). Apparently absent east of the Adriatic Sea, apart from the records

by Warren (1926) from Bosnia and Kronstadt (Rumania).

Habitat. — Flowery and usually dry localities, meadows, waysides, etc. in the low-

lands, in Switzerland rarely above 1000 m (Kauffmann, 1951), in Morocco, however,

up to 2700 m (Djebel Anngour; RJ); caught at the same height in the Sierra Nevada

by Manley & Allcard (1970).

Biology. — Bi- or trivoltine; flight period in Switzerland from April to October

(Kauffmann, l.c.).

The larvae have been reared by Powell in N Africa on Malope malachoides (Malva-

ceae) (cf. Guillaumin, 1964).

Geographic variation. — The variation of this rather variable species has been dis-

cussed by Verity (1925) in his well-known erratic way, giving many new names rather

than making the variation surveyable. He recognized four “races” and, in 1947, he even

mentioned five “races” in France alone. I find it impossible to define subspecies of this

species. The variations may be described as follows.

The first generation is largest in N Africa and many Spanish localities, & 12.2—14

mm, smallest in the Alps, $ 12—12.5 mm. Spots above variable, smallest in the Alps,

where the ground colour is rather pale. Ground colour unh in various shades of greenish

or ochreous grey or fulvous to warm reddish fulvous, more or less overlaid by dark

scales. The colour seems to depend on microclimatic conditions. In a series of eight males

from the Djebel Anngour district (High Atlas), 2700 m, June, the colour varies from

a very warm reddish ochreous with slight dark superscaling to an evenly darkened cold

greyish-green. In the Alps the colour is pale greyish-green with less contrasting white

spots. Particularly in N Africa, but also elsewhere, the white spots unh are often outlined

by capillary black streaks. These streaks are generally absent in specimens from dry

localities in Italy and S France.

The ground colour unh in the second generation is in general yellowish or fulvous,

more or less overlaid by dark scales; usually of a warmer tinge than in the first genera-

tion, sometimes very vivid reddish fulvous. In some specimens from the Alps the ground

colour unh is cold pale greenish grey, the same as in the first generation. About as large

as the first generation, in the Alps usually a little smaller, $ 10.6—12.2 mm.

R. DE JONG: Pyrgus in the Palaearctic region 75

A possible third generation is differing not much from the second generation.

We may summarize the variation as follows: the higher the temperature (north to

south, high to low, spring to summer), the warmer the colour. Local differences can

obviously complicate this general picture.

The difference between pale greenish grey and warm fulvous is not as great as one

would suppose at the first sight. It is only due to the intensity of the pigmentation of the

yellow scales, combined with the intensity of the dark superscaling. Pale yellow scales,

suffused by few dark scales give the impression of pale greenish grey, dark yellow

scales with many dark scales give a brownish tinge, etc.

For the particulars of microclimatic, rather than geographic, forms may be referred to

Verity (1919, 1925, 1928, 1940), who is the author of twelve of the fifteen names,

given to varieties of this species. Here, only the name of the extreme form of the Alps

is mentioned, as it is often used in literature: conyzae Guenée. This form is, however,

not confined to the Alps; I have seen specimens from Septemes near Marseille which

are indistinguishable from first brood specimens from the Alps, while the more

frequently occurring form of Septémes is larger and unh warm ochreous.

Material examined. — 322 & 227 9:39 g 29 £ Morocco (BM, RJ), 54 8 54 9

Algeria (BM), 39 g' 22 2 Spain (BM, ML, ZMA, RJ), 7 3 7 @ Portugal (BM),

142 & 86 9 France (Basses Pyrénées, Hautes Pyrénées, Vienne, Charente, Lot, Tarn,

Bouches-du-Rhöne, Vaucluse, Var, Alpes Maritimes, Basses Alpes, Hautes Alpes, Savoie)

(BM, ML, ZMA, HC, RJ, HO), 2 g 5 2 Switzerland (Valais) (BM, ML), 4 4 2 9

N Italy (ML, ZMA, RJ, HO), 35 $ 22 9 C Italy (BM, ML, ZMA).

Geographic history. — The origin as well as the geographic history of the species is

entirely obscure.

15. Superspecies Pyrgus carlinae Rambur

Taxonomy. — Until Picard’s publication in 1950, carlinae and cirsii were considered

separate species, as they differed constantly in facies and genitalia. Picard (1950b)

found in the French Alps intermediate specimens almost as numerous as typical

specimens, particularly in La Bessée (Hautes Alpes), but he did not give exact data.

Kauffmann (1951, 1954b) could not decide on conspecificity, although he knew Picard’s

paper. Virtually the only reason for Kauffmann’s considering carlinae and cirsii separate

species was that they differ elsewhere. In this, as well as in other cases it is apparent that

the biological species concept was beyond this author. Apparently, Picard estimated the

number of intermediates too high. Guillaumin (1964) mentions a score of 14 % inter-

mediates in the localities where carlinae and cirsii come into contact (Hautes Alpes: La

Bessée, Le Lautaret). Additional studies are still in progress (Guillaumin, in litt.). For

the present study the most important fact is the interbreeding, as it indicates an in-

complete stage of speciation. Contact between the lowland form c/rs// and the mountain

form carlinae is only known from the French Alps. Apparently, they interbreed wherever

they have contact. It is nonsensical to consider them separate species. I prefer to

call carlinae and cirsii semispecies, as members of the same superspecies, thus indicating

the supposed, rather progressed stage of speciation. But of course, it is also tenable to

call them subspecies of a single species, see also Chapter 1.b. The decision is not im-

portant for the present study.

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Biologically the hybridization between P. carlinae and P. cirsi is interesting as P. car-

linae hibernates in the egg stage and P. cirsii in the larval stage (see Guillaumin, 1964).

Externally P. carlinae and P. cirsii can easily be distinguished by the more square-

shaped and much larger spots upf and the better marked uph in P. crsi7 (see also Kauff-

mann, 1954d). In the male genitalia differences can be found in the lateral apophyses

(four to six unequal teeth in P. carlinae, more than six, more or less equal, teeth in

P. cirsii) and in the subharpal plate (united to the harpe in P. carlinae, not united in

P. cirsiî). In the female genitalia the antevaginal region is sclerotized in P. carlinae,

membranous in P. cirsii. Good figures can be found in Warren (1926), Opheim (1956)

and Guillaumin (1964, 1966).

Distribution (Map 13) and biology. — Mainly confined to Europe: P. carlinae only

in the western and southwestern Alps, usually from 1500 m upward, P. cirsii mainly in

the lowlands, rarely at 1500 m or higher, in SW Europe, but also found in E Turkey.

Both semispecies are monovoltine, flying from July to September. As far as known,

both live on Potentilla verna.

Geographic history of the superspecies. — The hybridization between P. carlinae and

P. cirsii indicates a recent history of the superspecies and Picard (1950b) may be correct

in supposing that P. cérsi originated during the Last Glacial in a Spanish refugium and

P. carlinae in S France. However, P. cirsii may have a much wider distribution in E

Europe than we can conclude from the present data and we do better not to speculate on

refugial areas as long as the distribution of P. cirsii is insufficiently known. Although

it is still difficult to understand in what way the differentiation into the cold-

adapted P. carlinae and the warmth-loving P. cirsii took place, it is interesting to note

the short time, probably necessary for such an extreme ecological differentiation.

P. carlinae seems to have more primitive characters: the fusion of harpe and subharpal

region and the genital plate. This is important for assuming phylogenetic relations, but

it does not mean that the ancestor of P. carlinae and P. cirsii was adapted to high

altitudes. It may have been more eurytherm than the present semispecies. In this connec-

tion it is important to note that P. cirsii may fly at higher altitudes in regions where

P. carlinae is absent (Pyrenees, E Turkey).

Virtually the history of the superspecies is very obscure. It is hard to say how far the

contact between P. carlinae and P. cirsii may have been stimulated by human agency

(deforestation).

15.a. Pyrgus carlinae Rambur

Rambur, 1839 — Faune entom. de l’Andalousie: pl. 8 figs. 11, 12.

Type-locality: Alps.

Distribution (Map 13). — Confined to the western and central Alps, from the Alpes

Maritimes to Engelberg (C Switzerland) and Tomsee (Ticino). According to Kauff-

mann (1954b), records of more eastern localities are based on misidentifications (or on

false data). This is quite possible but the fact that Kauffmann had not seen specimens

from more eastern localities cannot be a definitive proof for the absence of the species

in the eastern part of the Alps.

Habitat. — Alpine meadows, often abundant in damp places; usually from 1500 m

upward (Guillaumin, 1964), in Switzerland from 1000 m upward, occasionally as low

as 700 m (Kauffmann, 1951). According to Verity (1940) in Savoie up to 3200 m.

R. DE JONG: Pyrgus in the Palaearctic region 77

Biclogy. — Monovoltine; flight period from the middle of July to the beginning of

September (Kauffmann, 1951).

Food plant, Potentilla verna (Rosaceae) (Verity, 1940; Guillaumin, 1964); accord-

ing to Kauffmann (1954c) also other species of Potentilla are probably accepted as food

plant by the larvae.

The species hibernates as egg (Kauffmann, l.c.; Guillaumin, l.c.).

Geographic variation. — The species is rather variable, particularly in the colour unh

which can be red, brown, ochreous yellow or greenish. Kauffmann (1951, 1954b)

recognized four subspecies based on size, extension of the spots above and colour unh:

carlinae (type-locality: Alps), atratus Verity (type-locality: Vaud, Switzerland), ochro-

ides Kauffmann (type-locality: C Alps) and cottianus Kauffmann (type-locality: Cottian

Alps). The species appears to be rather sensitive to differences in microclimate, resulting

in a high local variability. Therefore it is not surprising that the four subspecies recognized

by Kauffmann are not clearly defined and overlap to a great extent in their characters.

The variation appears to be clinal, at least as far as the colour unh is concerned; it can

be described as follows:

In the S Alps (Alpes Maritimes, Basses Alpes) the colour is reddish, red-ochreous or,

more frequently, bright ochreous to yellow-ochreous. More to the north the red colour

disappears. In the Cottian Alps the colour varies from light brownish ochreous to olive-

brown, the veins being clearly outlined by lighter scales especially in the darker

specimens. Still further north, in the Val d’Aosta and throughout the Swiss localities the

colour is still darker and a reddish tinge is absent, the veins are clearly contrasting with

the ground colour; many specimens, however, are not darker than specimens from the

Cottian Alps.

Small specimens with reduced spotting above and greenish unh may occur everywhere

with other forms; they are perhaps most common in the northwestern part of the range.

It does not appear useful to divide this variational pattern into subspecies.

Material examined. — 354 § 107 9:37 & 2 @ Alpes Maritimes (St. Martin de

Vesubie) (BM), 33 8 3 9 Basses Alpes (BM), 58 & 17 Q Hautes Alpes (BM,

ZMA, RJ), 26 4 26 ® Savoie (BM), 97 & 40 9 Valais (BM, ML, ZMA, RJ), 19 g'

5 Q rest of Switzerland (BM, ML, ZMA, RJ), 75 & 9 @ Val d'Aosta (BM, ZMA,

RJ), 7 g 1 9 Piedmont (BM), 2 & 4 2 Tirol-Carinthia (BM).

Geographic history. — See the history of the superspecies.

15.b. Pyrgus cirsii Rambur

Rambur, 1839 — Faune entom. de l’Andalousie: pl. 8 figs. 12, o.

Type-locality: Fontainebleau.

Distribution. (Map 13). — SW and C Europe. According to Manley & Allcard (1970)

distributed throughout the Iberian Peninsula, but reliable records of the occurrence in

Portugal have not been found by the author. Throughout the greater part of France,

north to approximately 49° N Lat. (Verity, 1947). In Switzerland found only in the

Jura. In W Germany north to approximately the northern boundary of Bavaria, espe-

cially between the Danube and the Main (Alberti, 1927). Further east very incompletely

known, partly due to the fact that the species has long been classified as a variety of

Pyrgus alveus, and to the fact that the name fritillum Schiff. has frequently been used

for this and for other species. According to Verity (1947), near Vienna, particularly

west of the city. Warren (1926) even mentioned Sofia.

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Very surprisingly, De Lattin (1950) recorded the species from E Turkey (west side

of Van Lake) and Alberti (1969) added an other locality in E Turkey (Göle, in the

Vilayet Kars). These discoveries suggest a much wider distribution of the species in SE

Europe than at present known.

Habitat. — Dry, flowery places, usually at low altitudes, in the Swiss Jura not above

1200 m (Kauffmann, 1951), in the French Alps up to 1500 m and in the Pyrenees up

to 2000 m (Guillaumin, 1964). Recorded in Spain between 900 and 1750 m (Manley

& Allcard, 1970) and in E Turkey between 2300 and 2600 m (Alberti, 1969).

Biology. — Monovoltine; flight period in the Swiss Jura from the end of July until

the beginning of September (Kauffmann, 1951). Manley & Allcard (1970: 14), how-

ever, stated: “There are two generations each year, but we have found the species only in

July and August”. Their record of two generations is undoubtedly copied from Verity

(1925), but I have not been able to find any indications of more than a single brood

per year.

Food plant: Potentilla verna (Rosaceae) (Guillaumin, 1964). According to this

author the species hibernates in the larval stage, but it is not known whether this is also

the case at higher altitudes.

Geographic variation. — As in P. carlinae the variation of P. cirsii seems to be clinal

in a north-south direction. The specimens in Spain (ibericus Gr.-Gr. = fabressei Obth.)

are usually large, & to 14.7 mm, with extensive spotting and colour unh ochreous to a

warm fulvous or reddish, slightly overshaded by darker scales (usually in patches). In

addition, smaller specimens with darker unh (more saturated with dark scales) occur

throughout Spain; I have seen such specimens from as far south as Sierra de Alfacar.

In S France the specimens are usually smaller; in a series of 18 males from Bagnols-

les-Bains (Lozére) the largest male was 13.1 mm; but in other places the specimens may

be as large as in Spain, e.g. Luberon (Var) & to 14.5 mm. The colour unh is variable,

often darker, with a deeper red, than the colour in Spanish specimens, but sometimes

also ochreous in various tinges. The smaller and deeper tinged French form was named

parafabressei by Verity who (1925, 1947) stated that the zbericzs-like specimens in S

France are confined to the hottest and driest localities. The specimens from N France to

C Europe are still smaller, 4 to 13 mm, but often smaller, with dark underside, unf

deep black, unh strongly overlaid by dark scales to a brown or greenish colour, but

parafabressei-like specimens also occur. This is the nominate form. A form occurs in the

northern part of the Swiss Jura which was named /ramelensis by Kauffmann (1951),

and which is characterized a.o. by the absence of a reddish tinge unh. I have not seen

specimens from that region, but the description agrees very well with many French

specimens. There is therefore no reason for the separation. Apart from this the name

tramelensis may be synonymous with nigrocarens, given by Verity (1925) to a specimen

from Tramelan (figured by Reverdin, 1910), but Verity left it an open question whether

this is a race or an individual variation. Specimens with a cold greyish-greenish unh,

named herrichii by Oberthür (1910) occur throughout the range of the species.

The distribution of the various forms suggests that the variation is clinal and largely

or entirely due to microclimatic differences.

The five males and the two females from NE Turkey do not differ constantly from a

series from Pottenstein (north of Nürnberg) (Alberti, 1969). However, De Lattin

(1950) described the subspecies twrcivola on a single male, adding to his description:

“Eine grosse Rasse (Vfllange 10 mm), die infolge ihrer intensiven Weisszeichnung auf

R. DE JONG: Pyrgus in the Palaearctic region 79

den ersten Blick os stark von der gewöhnlichen fritillum der westlichen Fluggebiete

abweicht, jedoch im Genitale (Abb. 8) nicht von ihr zu unterscheiden ist, und die daher

zweifellos als östliche Vikariante zu dieser Art gezogen werden muss” (p. 326). Accord-

ing to my measurements a fore wing length of 10 mm cannot be considered large, but

very small instead. For the rest, bold statements based on a single specimen should be

avoided.

Material examined. — 275 4 109 2:75 g 39 9 Spain (Andalusia, Old Castille,

New Castille, Aragon, Catalonia) (BM, ML, ZMA, RJ), 2 4 2 9 Andorra (BM),

186 4 64 9 France (Ariège, Haute Garonne, Lozère, Bouches-du-Rhône, Vaucluse,

Var, Alpes Maritimes, Basses Alpes, Hautes Alpes, Charente, Lot, Dordogne, Puy-de-

Dôme, Saône-et-Loire, Haute Savoie, Loire Inférieure, Eure-et-Loire, Yonne, Seine-et-

Marne, Doubs, Bas-Rhin) (BM, ML, ZMA, RJ), 5 & Switzerland (Vaud) (BM), 7 &

3 9 W Germany (Nürnberg, Fürth, Dillingen) (BM, RJ), 1 9 Austria (ML).

Geographic history. — See the history of the superspecies.

16. Pyrgus cinarae Rambur

Rambur, 1839 — Faune entom. de l' Andalousie: pl. 8 figs. 4, 5, j.

Type-locality: Sarepta.

Distribution (Map 14). — Discontinuously from Spain to Turkestan. In Spain only

recorded from the Montes Universales (C Spain) (cf. Manley & Allcard, 1970), but in

the BM there is a female from Sierra de Alfacar in S Spain, collected by Ribbe.

The species is widespread in Macedonia (Thurner, 1964), which is the nearest place

to Spain, where it has been collected. Further in the Balkans only known from Sliven

(Bulgaria) (Rebel, 1903; see also Material examined). In Turkey widely, but evidently

locally spread. According to Staudinger (1881) near Brussa, in Pontus and in Lydia; in

addition Warren (1926) mentioned Kurdistan and Wagner (1929), Aksehir; see also

Material examined. From S Russia only few records: apart from specimens in various

collections labelled “S Russia”, there are specimens from Sarepta and Guberli in the

BM, whereas Alberti & Soffner (1962) recorded the species from the eastern Donez

region. In the BM there is also a male labelled ‘Turkestan’, but I doubt the label as no

other specimens from that region seem to be known.

Habitat. — Dry, hot localities with scarce vegetation, steppes, mountain slopes, in

Macedonia up to the timber line (Thurner, l.c.).

Biology. — Unknown.

Geographic variation. — There is a slight geographic variation in the colour unh.

Warren (1927) described the Spanish form under the name c/orinda, based on the

following characters: ground colour unh bright yellow with veins outlined in light

yellow; spots uph slightly buff or cream in the male and of a deep, almost ochre, shade

in the female. According to Warren “the effect is very much more striking than the

description would lead one to suppose” (p. 81). Virtually the difference between

specimens from Spain and from the eastern Mediterranean is very slight: some eastern

specimens are indistinguishable from Spanish ones.

It is only a matter of taste whether one regards the Spanish form as a subspecies or

not. That an isolated population is different, is not surprising, but it is interesting that

in this instance the difference is so slight. This does not need to be an objection against

a subspecific separation.

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pyrgus cinarae clorinda Warren

Warren, 1927 — Ent. Rec. 39: 81—82.

Type-locality: Tragacete, Villacabras (Cuenca).

Material examined. — 5 & 1 2:2 oO Tragacete, 2 4 Huelamo, 1 ¢ Villacabras,

1 ® Sierra de Alfacar (all BM).

Distribution. — C (and S?) Spain.

Subspecific characters. — See above.

Pyrgus cinarae cinarae Rambur

Material examined. — 87 d 17 9:7 & 1 ® Macedonia (Skopje, Naussa, Seli,

Ohrid, Petrina planina) (BM, ML, RJ), 24 g° Bulgaria (Sliven) (RJ), 33 & 9 2

Turkey (Kedos, Aksehir) (BM, RJ), 22 8 7 9 S Russia (Sarepta, Guberli, “S Russia’)

(BM, ML), 1 g° Turkestan (BM).

Distribution. — From Macedonia eastward.

Subspecific characters. — Ground colour unh varies from yellow to dull olive-green;

spots uph in male often cream-coloured, in female always somewhat more darkened.

Geographic history. — The slight difference between Spanish and E Mediterranean

specimens suggests that the discontinuity in the distribution is of a recent age, presum-

ably Postglacial, but it is also possible, that it is only a reflection of a stable gene

configuration or of stable habitat circumstances. In the first mentioned case the species

may have spread to Spain during the Boreal or Atlantic climate phase, in the other cases

it is possible that the Spanish population is a relic of a continuous Mediterranean

distribution in the Third Interglacial. Thus, the reconstruction of the history of cinarae

is highly unsatisfactory, but it seems closely connected with the history of the fauna of

the dry and warm mountain slopes of the Mediterranean.

17. Pyrgus armoricanus Oberthür

Oberthür, 1910 — Et. Lép. Comp. 4: 411; pl. 57 figs. 509—517.

Type-locality: Rennes (W France).

Distribution (Map 15). — West Palaearctic. From NW Africa through S and C

Europe to SE Russia and NE Iran (see below, Material examined). In NW Africa only

recorded from the Middle Atlas in Morocco and Algeria (Picard, 1950d; Higgins &

Riley, 1970). The northern distribution limit runs through Belgium (cf. Lempke, 1953)

and N Germany (Urbahn, 1939; Alberti, 1940; Friese, 1956). A few localities still

further north are known: in Denmark in N and W Seeland, Jutland (a single specimen)

and Bornholm (Wolff, 1939) and in Sweden in the extreme south (Scania: Benestad,

Sandhammaren, Löderup: Nordström, Opheim & Valle, 1955). The northern limit

further eastward is very badly known, partly due to a possible confusion with Pyrgus al-

veus. The northernmost locality in Russia from where I have seen a reliable record, is

Tula (Warren, 1926). The species is wide-spread in S Russia (Alberti & Soffner, 1962).

In Iran southward to Shiraz (Brandt, 1939) and eastward to the Achal Tekke region

(BM), apparently restricted to the mountains, as in Iraq (Wiltshire, 1957). Through

W Syria southward to the Lebanon.

Habitat. — Flowery and often dry places, in S Russia in the steppes and even in the

R. DE JONG: Pyrgus in the Palaearctic region 81

semi-desert (Alberti & Soffner, 1962); most common in the lowlands. In Switzerland

and Italy infrequently above 600 m and very rarely at 1200 m (Verity, 1940; Kauff-

mann, 1951), in Iraq restricted to the mountains, from 1300 to 2000 m (Wiltshire,

1957).

Biology. — Bivoltine, but a partial third brood occurs at least in Italy; flight periods

mainly May—June and August—September, but specimens have been found in the

Mediterranean region during all months from February to October (Warren, 1926). In

Iraq adults of the two generations occur in May and July—October respectively (Wilt-

shire, 1957).

Females depositing eggs on Fragaria vesca and Potentilla reptans (Rosaceae) have

been observed by Rehfous (iz Reverdin, 1912) and on Potentilla verna by Picard

(1950c). Rehfous reared the larvae until the beginning of November and as they were

still far from full-grown at that time, it is highly probable that P. armoricanus hibernates

as larva.

Geographic variation. — The male genitalia vary geographically with regard to the

shape of the apex of the cuiller: this apex is sharply pointed and the free proximal edge

of the cuiller is concave in the western populations (west of the Balkans) (armoricanus

type); in the eastern populations the apex is blunt and the free proximal edge of the

cuiller is more or less straight (persiczs type). The blunt apex has not been found west

of the Balkans, but the pointed apex has been found in many parts of the eastern range

of the species, in the Balkans, in Turkey and in Iran. This has been known since long

(Reverdin, 1916; Warren, 1926; Alberti, 1940), but records concerning the proportion

of both forms are lacking. The blunt apex apparently predominates in the southeastern

Balkans and further eastward. I examined the genitalia of 103 males from the eastern

part of the range of the species and found the following proportions:

locality number armoricanus number persicus

examined type transitional type

Hungary 10 8 1 1

Croatia 6 6

Bosnia 2 1 1

Dalmatia 7 2 3 2

Montenegro 1 1

Macedonia 3 1 2

N Greece (Olympus-Saloniki) 17 7 10

Bulgaria 7 2 1 4

Rumania 1 1

Bosphorus 5 1 1 3

Turkey 30 1 3 26

Armenia 1 1

Georgia 1 1

Syria il 1

Lebanon 4 4

Iran (Elburz Mts) 1 1

Achal Tekke 2 1 1

S Russia (Zymljanskaja) 4 4

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

It is, of course, often difficult to call a structure ‘‘transitional’’, but even if we list all

specimens considered transitional as armoricanus, then still is the predominance of the

persicus type of cuiller in specimens collected east of the Balkans evident. Within these

two groups (armoricanus and persicus) various forms have been described as subspecies

but at least some of these belong to the ecophenotypic variation (e.g. cacaoticus). The

following classification has to be considered provisional, but this is not very important

for the reconstruction of the geographic history of armoricanus.

armoricanus subspecies group

Differs from the persicus subspecies group only in the sharply pointed apex of the

cuiller.

Distribution. — The western part of the range of the species, eastward to Hungary

and the northern Balkans. Perhaps much further east, as Warren (1926) recorded a

male from Tula in C Russia with genitalia intermediate between armoricanus and per-

SECHS.

Pyrgus armoricanus maroccanus Picard

Picard, 1950 — Bull. Soc. Sci. nat. Maroc 28: 110.

Type-locality: Morocco.

Material examined. — 29 g 10 2:16 & 7 £ Morocco, 13 8 3 ® Algeria (BM).

Distribution. — Morocco and Algeria.

Subspecific characters. — Rather large, $ 13—14.2 mm, with large and obvious spots

above, also uph.

Note. — A slightly differentiated form, indistinguishable from some specimens from

Spain, S France and C Italy.

Pyrgus armoricanus armoricanus Oberthür

Material examined. — 386 & 167 2:50 & 14 9 Spain (BM, RJ), 194 & 109 2

France (BM, ML, ZMA, RJ), 2 9 Belgium (ML), 2 4 1 9 Germany (Kreuznach,

Germany”) (BM, ML), 19 8 7 2 Switzerland (BM, ML, ZMA), 14 8 4 2 Austria

(BM, ML, RJ), 57 g° 4 2 Italy (BM, ML, ZMA, RJ), 15 & 7 2 Sicily (BM), 6 dg

6 2 Corsica (BM), 3 ¢& Istria (RJ), 12 § Croatia (BM), 14 G 13 9 Hungary

(BM).

Distribution. — European part of the range of the armoricanus subspecies group, east-

ward to C Germany.

Subspecific characters. — A variable subspecies. 11.3—14.2 mm. Above spots

variable, usually smaller than in the subspecies maroccanus and disjunctus, suffusion

with light scales usually not very pronounced.

Variation. — The first generation, and in the north (from C France northward) also

the second generation, have usually a dark upperside, with rather small spots upf and

indistinct spots uph (submarginal spots often partly absent). This probably ecophenotypic

form occurs at the type locality of subspecies armoricanus. In southern regions specimens

of the second brood are often suffused with yellow above and more frequently than in

R. DE JONG: Pyrgus in the Palaearctic region 83

the first brood with a tawny unh (fulvoinspersus Verity, 1919 = siciliae Oberthür,

1913). According to Verity (1940) this form occurs in relatively dry localities. The first

and the second brood under relatively humid conditions are characterized by a rather

large size, a black ground colour unf and a dark greenish or brownish ground colour!

unh (cacaoticus Verity, 1929). In the second brood also specimens occur with a vivid

reddish ground colour unh (rzfosaturus Verity, 1925 = corsicus Oberthür, 1919, see

Picard, 1949d). In southern regions the specimens are often better marked above,

especially uph, than in the north, but much variation occurs. I regard the above forms

as ecophenotypic variations of the subspecies.

According to Alberti (1940), the Hungarian populations can be referred to subspecies

disjunctus but the Hungarian specimens examined by me may equally well be listed as

subspecies armoricanus. It may be recalled that of the ten males from Hungary of which

I examined the genitalia, one has the persicus type of cuiller and one is intermediate

between armoricanus and persicus. This means that the influence of persicus is per-

ceptible as far north-west as Hungary. The Hungarian populations can be listed as ssp.

armoricanus, as the influence of persicus appears to be small.

The subspecific status of S German populations is unclear, probably as a result of lack

of material, but also of the slight degree of differentiation between the subspecies

armoricanus and disjunctus. Alberti (1940) regarded the S German populations as ssp.

armoricanus, but Kauffmann (1951) listed the populations of S Bavaria as ssp. dis-

janctus.

Pyrgus armoricanus disjunctus Alberti

Alberti, 1940 — Mitt. Münch. Ent. Ges. 30: 252; pl. 2 figs. 4-c, 7-a.

Type-locality: Halle.

Material examined. — 5 d 2 9:1 g Giessen (ML), 2 d 2 £ Halle, 1 G Praha,

1 g Bohemia (all BM).

Distribution. — The northern part of C Europe; from C Germany and Czechoslovakia

northward. Alberti (1940) also assumed that the Hungarian population and even the

Algerian belonged to this subspecies.

Subspecific characters. — Small, $ 11—12 mm; above with large and conspicuous

spots, also uph, and strong suffusion with light scales; below also with large spots,

ground colour unh greyer than in ssp. armoricanus.

Note. — Small specimens can be found throughout the range of the species. Some of

these are indistinguishable from specimens from C Germany. Specimens in W Germany

(Kreuznach; see above, under ssp. armoricanus), Belgium, N France and Styria are often

not larger than ssp. disjunctus but have smaller spots.

Ssp. disjunctus is probably rather constant in its distribution range. The differentiation

is slight with regard to the variability of ssp. armoricanus.

persicus subspecies group

Characterized by the blunt apex of the cuiller. It is impossible to distinguish this sub-

species group on the basis of external characters.

Distribution. — The eastern part of the range of the species, east of the northern

Balkans.

I include the populations of Bosnia, Dalmatia and Bulgaria in this subspecies group,

although these could be listed as ssp. armoricanus as well (see above).

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pyrgus armoricanus persicus Reverdin

Reverdin, 1913 — Bull. Soc. Lép. Genève 2: 218—224; pl. 21 fig. 4; pl. 22 fig. 9.

Type-locality: Kuldsar (Iran).

Material examined. — 151 4 52 9:2 & Bosnia (RJ, ZMA), 7 g 1 ® Dalmatia

(BM, RJ), 1 8 Montenegro (BM), 3 4 Yugoslavian Macedonia (RJ), 49 8 21 9

Greece (BM, ML, ZMA), 8 3 1 9 Bulgaria (BM, ML), 1 & Rumania (BM), 35 &

9 Bosphorus (BM), 33 & 8 ® Turkey (BM, RJ, ML), 1 g Armenia, 1 & Georgia,

5 d 2 9 S Russia (Zymljanskaja, Novorossiisk), 1 & 3 9 N Syria, 18 3 9

W Iran, 1 g 4 9 Elburz Mts., 2 & Achal Tekke (all BM).

Distribution. — As that of the persicus subspecies group, except the Lebanon.

Subspecific characters. — Very variable. & 12—14 mm; usually well spotted above;

ground colour unh ochreous to greenish.

Variation. — Pfeiffer (1927) described the Anatolian population as prostanae, dif-

fering from the Iranian form in the larger spots above and the greenish ground colour

unh. Neither of these characters, however, is confined to Anatolia, nor is the Anatolian

population uniform in this respect, although the greenish ground colour unh may occur

more frequently in C Anatolia than elsewhere. The few specimens examined from Iran

cannot be decisive for a subspecific distinction of the Anatolian population. The form

lecerfi Verity, 1928 (= reverdini Le Cerf, 1914), described from Iran (Bazouft and

Lagherit) appears to be an extreme variation with very well-developed and sharply

contrasting spots uph (even with white basal spots). I have not seen such extreme

specimens, but some specimens from Turkey (near Ankara) resemble these.

Pyrgus armoricanus philonides Hemming

Hemming, 1931 — Ann. Mag. Nat. Hist. (10) 8: 534—535.

Type-locality: Khan Ghaffa (Lebanon).

Material examined. — 5 ¢ Lebanon (BM).

Distribution. — Confined to the Lebanon.

Subspecific characters. — Small, & 11.6—12.7 mm. Differs from ssp. persicus mainly

in the pale yellowish ground colour unh. Ssp. philonides is a little differentiated form

in comparison with the great variability of persicus.

Note. — According to Hemming (1931: 535): “The genitalia have been examined and

differ in no way from nominotypical armoricanus.’’ The genitalia of four males (incl.

two paratypes) of ssp. philonides examined by me turned out to be of the persicus type.

Geographic history. — Undoubtedly the armoricanus and the persicus subspecies

groups arose in a West Mediterranean and an East Mediterranean glacial refugium

respectively. There is no reason to suppose that the present differentiation within these

groups is older than of postglacial age. The two subspecies groups may have originated

during the Last Glacial.

The northern localities of this warmth-loving species (Denmark, S Sweden) seem to

be relics of a wider distribution in a postglacial period which was warmer than the

present (the Atlantic climatic optimum?). If ssp. disjunctus is not a nongenetic

ecophenotypic variety, a recent colonization of Denmark and S Sweden from N Germany

R. DE Jong: Pyrgus in the Palaearctic region 85

can also be taken into account, or else a long-lasting similar selection pressure in Den-

mark and C Germany can have brought about the characters of ssp. disjunctus (in this

case disjunctus cannot be regarded as a subspecies).

In the Balkans the two subspecies groups evidently met postglacially. The fact that the

armoricanus type of genitalia has been found in Turkey and N Iran does not necessitate

the assumption of any other contact between the groups (cf. Alberti, 1940), as this may

be a part of the normal variation of the persicus group.

18. Pyrgus alveus Hübner

Hiibner, 1803 — Samml. Eur. Schmett., 1: 70; pl. 92 figs 461—463.

Type-locality: Germany (Verity, 1940).

Distribution (Map 16). — Palaearctic. From NW Africa and Spain to the Amur

region. In Scandinavia north to approximately 63° N Lat. The northern limit further

eastward is not known. The records of Pyrgus serratulae from Vologda and Kotlas

(European Russia) by Krulikowsky (1909) presumably refer to P. alveus. The same may

be the case with the record of P. serratulae from the River Vilui (NE Siberia) by Herz

(1898). The distibution in W Europe is rather well known. Detailed distribution maps

can be found in Agenjo (1963 - Spain), and Nordstrém, Opheim & Valle (1955 -

Scandinavia). P. alveus has a restricted range in Peninsular Italy, viz. the larger mountain

massives of Marche and Abruzzi (as P. serratulae and P. bellieri) (Verity, 1940).

In E Europe presumably widely distributed, but many records are unreliable as they

may be based on misidentifications. In Asia possibly continuously distributed through

Siberia to Vladivostock. In China only found in the Kuku-Nor region, Szechwan and SE

Tibet (see below, Material examined).

Rare in warm or lowland districts.

Habitat. — Flowery meadows, waysides, often in large numbers in damp places, e.g.

on muddy shores along streams. Most common in hilly and mountainous districts. In the

Alps up to 2700 m (Warren, 1928; Verity, 1940), in Szechwan and SE Tibet still

found at 4700 m (BM), but also just above sea level, e.g. in N France (Picard, 1948d)

and the northern part of E Germany (Urbahn, 1939; Friese, 1956).

Biology. — Monovoltine, but bivoltine in some warm and low localities in the

southern Alps (Kauffmann, 1948). It may even produce a second generation at high

altitudes (2200 m) in extremely hot years (Warren, 1953); other records of a second

generation are unreliable, as confusion with P. armoricanus cannot be excluded. The

flight period usually is in July and August, but particularly for the accretus subspecies

group (SW and W Europe) the emergence may begin at the end of May. Kauffmann

(1953) recorded accretus from SW Germany from the end of May to the end of June,

and the specimens I saw from N France mainly date from the end of May to the middle

of June. From W France (Charente) I also saw specimens from September which may

indicate a second generation.

According to Higgins & Riley (1970) in NW Africa there are two broods a year,

but neither Warren (1953) nor I have found any indication for a second generation of

P. alveus (ssp. numidus) in NW Africa. The specimens examined dated mainly from

the end of May (25.V) until the middle of July.

Various food plants have been recorded, but reliable records are rather scarce. Most

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

frequently various species of Helianthemum (Cistaceae) and Potentilla (Rosaceae)

have been mentioned, Rehfous (77 Reverdin, 1912) and Kauffmann (1948) observed

depositing of eggs on Helianthemum, Bergmann (1952) on Potentilla; in addition, the

last person recorded Polygala (Polygalaceae) and Agrimonia (Rosaceae) as food plants.

Geographic variation. — An important part of the geographic variation has been

outlined in Chapter 2.b, but it will be now discussed in more detail. P. alveus has the

most variable male genitalia of all Palaearctic species of Pyrgus. This variation is partly

geographic, partly individual. It is particularly apparent in the development of the

cuiller, but also other parts are affected; style, stylifer, antistyle, uncus and lateral

apophyses. I will only mention the most obvious variations. The geographic variation in

the genitalia is partly connected with a geographic variation in the facies, but generally

there is a large individual variation in the facies, presumably due to a high sensit-

iveness to differences in temperature and moisture. Kauffmann (1954c) recorded a

breeding experiment that indicated that pupae of P. alveus are very sensitive to frost:

pupae exposed to temperatures below 0° C produced butterflies which were above

totally black. This is of course important for the evaluation of the observed variability.

The geographic variation may be described as follows.

Pyrgus alveus numidus Oberthiir

Pl. 6 Fig. 91, 100

Oberthiir, 1910 — Et. Lép. Comp. 4: 404; pl. 50 figs. 484—486.

Type-locality: Algeria.

Material examined. — 130 8 33 9:78 & 9 Q Morocco (Azrou, Daiet-Achlef,

Timhadit, Ifrane, Assila, Djebel Hebbri, Bekrit, Tarzeft, Sidi Ali, Anngour) (BM, MP,

RJ), 52 8 24 © Algeria (Lambèse, Djebel Aurès, Batna) (BM).

Distribution. — Confined to NW Africa.

Subspecific characters. — ¢ genitalia: cuiller high and narrow, apex rising well above

the dorsal ridge of the harpe; harpe of a more or less even width throughout. Facies:

a very conspicuously spotted form, also uph. Central spot unh usually with a projection

towards the base of the wing and towards the termen along vein 6, absent in 10 % of

the specimens examined. Size: & 13.8—15.6 mm.

Variation. — I could not establish a difference between the Moroccan and the Al-

gerian specimens.

accretus subspecies group

The cuiller in specimens from W and SW Europe is characterized by the short free

part of the proximal edge of the cuiller, which is one quarter or less of the total

depth of the proximal edge and sloping towards the harpe, sometimes almost

touching it. The shape of the cuiller is also characteristic, varying from the alvezs type

to the bellieri type (see Chapter 2.b).

Some Pyrenean specimens are nearly (?entirely) indistinguishable from P. bellieri in

the shape of the cuiller. It may be recalled that there are indications in the southern

Alps of a rare hybridization between P. bellieri and P. alveus accretus. Specimens with

a less developed cuiller can always be distinguished from ssp. alveus by the proximal

slant of the free part of the proximal edge of the cuiller, but as stated in Chapter 2.b

R. DE JONG: Pyrgus in the Palaearctic region 87

specimens have been found in the French Alps which are intermediate between ssp.

accretus and ssp. alveus (also in other characters). The ventral part of the lateral

apophyses is rather small in comparison with the type found in ssp. alvews, and shorter

than the shaft which connects it with the dorsal part.

In the northwestern part of the range (NE France/SW Germany) ssp. accretus possibly

emerges much earlier than ssp. alvezs in the adjacent regions (cf. Kauffmann, 1953;

see also above, under Biology). The situation in SE France is not very well known, but

at any rate the flight periods of ssp. accretus and ssp. alveus overlap.

The taxonomic position of the accretzs group is still uncertain. Warren (1953) gave

it specific rank, as mentioned in Chapter 2.b. If one gives the accretus group specific

rank, it is reasonable to consider numidus also specifically distinct from ssp. alveus, as

the differences with ssp. alveus are as obvious as between ssp. accretus and ssp. alveus.

For the time being the classification proposed here seems to be practical for the present

zoogeographic study.

Agenjo (1963) described the variation in the cuiller in the accretus group and part

of the a/vews group as clinal, particularly the variation in the direction of the proximal

edge. He based his conclusion on photographs of the genitalia of males from Bavaria, the

French Alps, the Pyrenees and C Spain. If he had also presented a picture of the genitalia

of a Scandinavian male, the variation would have been still more obvious, but it is

not correct to study clinal variation on a selection of individual specimens. Actually, the

alveus type is, at least in the character mentioned, rather constant throughout C Europe

and the accretus type is rather constant throughout France and a large part of Spain.

This does not suggest a gradual change of the character mentioned by Agenjo.

Within the accretus group nine ‘‘races’’ have been distinguished but as some appear

ecophenotypic variations or are difficult to define geographically, I will reduce the

number of subspecies to four and perhaps even this number should be reduced.

Pyrgus alveus insigniamiscens Verity

Pl. 6 Fig. 92, 101

Verity, 1929 — Trab. Mus. Cienc. nat. Barcelona 11: 14.

Type-locality: Sierra Nevada.

Material examined. — 2 d 2 9:2 $ 1 © Sierra Nevada (BM, RJ), 1 @ Sierra de

Alfacar (BM).

Distribution. — Only known from the Sierra Nevada and the adjacent Sierra de

Alfacar.

Subspecific characters. — Large, & 15—15.6 mm, 9 15.7—17.1 mm. Externally

almost indistinguishable from ssp. r4midus, particularly unh. In the male genitalia,

however, the free proximal edge of the high and rather narrow cuiller slopes distinctly

towards the harpe.

Pyrgus alveus centralhispaniae Verity

Verity, 1925 — Ent. Rec. 37: 56.

Type-locality: C Spain (Montes Universales).

Material examined. — 8 8 6 9:3 & 3 2 New Castille (BM), 3 2 Old Castille

(BM), 5 & Aragon (Albarracin, Tragacete) (BM, RJ).

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Distribution. — C Spain.

Subspecific characters. — Male genitalia: free proximal edge of the cuiller usually

closer to the harpe than in any form other than ssp. insigniamiscens. Above usually

very conspicuously spotted, also uph. Below resembling the Spanish form of Pyrgzs

carthami, but varying towards ssp. accretus.

Pyrgus alveus accretus Verity

Pl. 6 Fig. 93—94, 102—103

Verity, 1925 — Ent. Rec. 37: 55.

Type-locality: Gédre (Hautes Pyrénées).

Material examined. — 317 d 91 2:7 8 2 9 Asturias (BM, RJ), 34 g 1 9

Spanish Pyrenees - prov. Huesca (ZMA, RJ), 63 4 22 @ Spanish Pyrenees - prov.

Gerona (BM, MP, ZMA, HC, RJ), 13 & Andorra (BM, ZMA), 1 4 1 9 Basses

Pyrénées (RJ), 95 S 29 ® Hautes Pyrénées (BM, MP, ZMA, RJ), 2 8 Ariège (MP,

RJ), 54 g 12 ® Pyrénées Orientales (BM, MP, ZMA), 1 g' Aude (ZMA),1g 1 9

Hérault, 1 9 Gers, 3 4 Aveyron, 1 4 Charente Mar. 5 & 2 9 Charente (all BM),

2 d 2 2 Puy-de-Dôme (MP), 8 g 4 9 Haute-Loire (BM), 1 & 1 9 Loire (BM),

2 4 1 Q Saône-et-Loire (BM, RJ), 12 g 8 9 Seine-et-Oise (BM, MP), 6 g 4 9

Eure (BM), 8 & Alpes Mar. (BM).

I examined the genitalia of 75 males from the Pyrenees and of 25 males from C and

N France. A reliable identification of specimens from the French Alps is only possible

by a genital examination. Therefore, I have enumerated only the specimens from the

French Alps of which I could examine the male genitalia of this subspecies as well as of

ssp. alvens.

Distribution. — From N Spain to N France and SW Germany. The eastern limit is not

exactly known, but ssp. accretus seems to be absent east of the Rhine and of the Swiss

Jura. According to Warren (1953) populations in NW Italy, in the Cottian and in the

Ligurian Alps should also be referred to this subspecies.

Subspecific characters. — With regard to the male genitalia ssp. accretus differs from

ssp. insigniamiscens and ssp. centralhispaniae in the free proximal edge of the cuiller

usually being less close to the harpe. Some specimens, however, are indistinguishable

from ssp. centralhispaniae in this respect, e.g. a male from Aude.

Usually less spotted than the two subspecies just mentioned, but there is much

variation. For difference with ssp. jurassicus, see below.

Variation. — In the Pyrenees there is much variation in size (4 13—15 mm), in

spotting above and below and in ground colour unh (bright yellow to greenish yellow).

The variation is continuous and certainly not geographically defined. Therefore, I

consider the distinction of pyreneialpium and necaccreta, both described by Verity

(1929), superfluous and their denomination as subspecies unjustified. A form living in

the departments Eure and Seine-et-Oise has been named /utetianus by Picard (1948d),

mainly differing from ssp. accretus in the larger spots and the stronger superscaling with

light scales on the upperside (Pl. 6 Fig. 94). However, the only difference I could

establish is the absence of darker specimens in the north, which may be due to the

absence of high mountains. Many Pyrenean specimens cannot be separated from

specimens from N France.

Specimens with very strong light suffusion above and with a pale yellow ground

R. DE JONG: Pyrgus in the Palaearctic region 89

colour unh from the department Gard have been named swbdecoloratus by Picard (in

Gaillard, 1952). Presumably it is an extreme variation, due to some particular ecological

conditions.

Large specimens from the French Alps have been distinguished by Verity (1925)

as grandis.

In the Pyrenees there exists much variation in the male genitalia. Some specimens may

be nearly or entirely indistinguishable from P. bellieri. Agenjo even distinguished the

subspecies bellieri hospitalis from Gerona. I could not find the original description, but

Agenjo (1963) used this name for his figure 1 (male genitalia) on plate 2. I have

seen comparable genitalia in four males taken in the eastern Pyrenees and in one male

taken on the Col du Pourtalet (Basses Pyrénées). For the present it seems advisable to

consider such specimens extreme variations of ssp. accretus, instead of representatives of

western populations of P. bellieri. The localities of P. bellieri nearest to the eastern

Pyrenees are in the south of the department Aveyron, more than 100 miles away.

Pyrgus alveus jurassicus Warren

y J

Warren, 1926 — Trans. Ent. Soc. Lond. 74: 121; pl. 42 figs. 9—12.

Type-locality: Jura.

Material examined. — 3 g 4 9: 1 9 Weissenstein (Swiss Jura), 3 g 3 9

Eclépens (Vaud) (BM).

Distribution. — French and Swiss Jura; Mont Saléve (cf. Warren, 1953).

Subspecific characters. — Differs from ssp. accretus mainly in being smaller (¢ to

about 13 mm) and in having very prominent nervures unh which are outlined in bright

yellow-orange. The subspecies is somewhat intermediate between ssp. accretus and ssp.

alveus externally; the male genitalia are typically accretus.

As it has a rather large distribution it may be more than an ecophenotypic variation,

as e.g. subdecoloratus. I did not see enough specimens to get a good impression of the

variation.

alveus subspecies group

From C Europe to China the cuiller is characterized by its free proximal edge which

is about one third the total depth of the proximal edge (varying to nearly one quarter)

and, with slight variation, is directed vertically to the ventral plane of the cuiller. The

ventral part of the apophyses is longer than the shaft. The apex of the cuiller is almost

level with the dorsal edge of the harpe, but in some specimens it rises well above this

edge.

There is very much individual variation in the male genitalia, particularly in the shape

of the cuiller. The variation is partly geographically defined. This has been shown by

Kauffmann (1954a) in a nice study on the variation in the southern Alps. Using

characters of 300 male genitalia he demonstrated that the percentage of specimens with

a deformed cuiller (usually a restriction in the horizontal depth) increases from nearly

50 % in the Central Alps to more than 80 % in the pre-alpine zone in the lowlands.

In the same way the proportion of specimens with a concavity in the proximal edge ot

the stylifer (once thought to be a characteristic of the Chinese reverdini, see Reverdin,

1916, and Warren, 1952) increases from about 50 % to nearly 90 %. Unfortunately,

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Kauffmann compared his rich material from the southern Alps with five males from

N Turkestan (Ili), from where Reverdin (1912) described the “subspecies” zliensis.

As he found a good resemblance between zliensis and the lowland alvezs of the

southern Alps, he concluded that they were conspecific and specifically distinct from

alveus, and named “iliensis’ of the southern Alps ssp. colurnus. This is quite incom-

prehensible, as alvens and colurnus, in this way distinguished, imperceptibly pass into

each other in the southern Alps. Moreover, five males from Turkestan are entirely in-

sufficient for a reliable impression of the local variation. Both types of genitalia, with

transitions, occur throughout the Central Alps, as correctly stated by Warren (1957)

and as can be seen from the fifty-fifty proportion given by Kauffmann himself. Both

types also occur in S Bavaria, C Italy and the Balkans in varying proportions. Thus, it

is even impossible to define colwrnus geographically. Perhaps a statistical study on much

more material from E Europe and Asia than available at present, would give interesting

results. At the moment we can only conclude that the co/wrnus type predominates in

some areas.

This variable alvezs type of genitalia is widely distributed in Asia. I have stated this

type in specimens from the Ili region (Boro Choro), the Altai, Abakan (north of the

Altai), the Kuku-Nor region and W Szechwan and SE Tibet. The photographs by

Reverdin (1912; pl. 18 fig. 1; the same has been reproduced by Warren, 1926; pl. 9

fig. 5) and by Kauffmann (1954a; pl. 2 fig. 7 and 8), representing iliensis from the

Ili region, also show the alvezs type of genitalia. However, the alveus type is accom-

panied in all these regions by the scandinavicus type (cuiller dorsally more developed

than ventrally, free proximal edge sloping away from the harpe, broad stylifer and

antistyle). This is most evident in the Altai and north of it (from W Siberia I saw

only the scandinavicus type). In the Kuku-Nor region and further south it may be due

to intergradation with ssp. speyeri. It is impossible to define exactly the geographic

boundary between the scandinavicus and the alveus types in Asia, at least so long we do

not have hundreds of specimens available from Siberia; and even then it may be im-

possible. Provisionally I shall list all Asiatic forms, except speyeri, in the alveus sub-

species group. The male genitalia are in spite of their variability much more constant

than the facies, and while it is difficult or impossible to define the variation in the

male genitalia geographically, the variation in the facies can, at least in part, be

geographically defined. It may be recalled that the species is very susceptible to

temperature influences, low temperatures causing a reduction of the white spots above.

Knowing this, it is difficult to define subspecies on the basis of external characters, but

the following classification may be useful.

Pyrgus alveus alveus Hiibner

Pl. 6 Fig. 95—96, 104—105

Material examined. — 554 8 190 2:4 g Alpes Maritimes (MP), 31 g 28 2

Basses Alpes (BM, ZMA), 31 9 9 @ Hautes Alpes (BM, MP, RJ), 38 4 13 2

Savoie (BM), 24 g' Haute Savoie (BM), 134 & 37 Q Italian Alps (BM, ML, ZMA,

RJ), 123 8 64 2 Switzerland (Vaud, Valais, Ticino, Berne, Uri, Unterwalden,

Grisons) (BM, ML, ZMA, RJ), 23 & 13 9 W Germany (Baden-Württemberg,

Bavaria) (ML, RJ), 67 g° 12 @ Austria (N Tyrol, Gross Glockner, Carinthia, Styria

Inf.) (BM, ML, ZMA, HC, RJ), 12 g 5 9 Hungary (BM, ML, ZMA), 4g 19

R. DE JoNG: Pyrgus in the Palaearctic region 91

Rumania (BM, ML, MW), 10 G 2 9 Bulgaria (BM), 31 4 4 9 Yugoslavia (Bosnia,

Herzegovina, Serbia, Macedonia) (BM, ML, ZMA, ZSM, RJ), 5 8 Albania (BM,

MW), 7 g Caucasus (MP, BA, RJ), 9 g° 2 2 Transcaucasia (BM, MP, ZMHB),

1 & Hyrcania (BM).

I examined the male genitalia of specimens from the various regions in the following

numbers: Alpes Mar. 4, Basses Alpes 16, Hautes Alpes 14, Savoie 12, Haute Savoie

10, Italian Alps 37, Switzerland 25, Germany 15, Austria 22, Rumania 3, Yugoslavia

26, Albania 4, Caucasus 7, Transcaucasia 6, N Iran 1.

Distribution. — C, E and SE Europe, Caucasus to N Iran. The distribution in

Russia north of the Caucasus is entirely unknown.

Subspecific characters. — Very variable; not every specimen can be distinguished

from specimens of other subspecies of the group (particularly from specimens of

centralitaliae and sifanicus). See the following paragraph.

Variation. — (1). C Europe. Some authors have given subspecific rank to forms

which are regarded here as individual or ecophenotypic variations. Apart from colurnus

(see above), these are:

(a). alticolus Rebel, 1910 (PI. 6 Fig. 96, 105). Small, but larger than Pyrgus warrenen-

sis, with reduced spots. A form of high altitude, usually flying with larger and more

distinctly spotted specimens. There is a confusing synonymy about this form. Evans

(1949) used the name for specimens belonging to Pyrgus warrenensis, which latter

name he regarded as a synonym of a/vews. Other names used for this form are ryffelensis

Oberthür, 1910, and scandinavicus Strand, 1903, cf. Warren (1953).

(b). claralveus Verity, 1934. Characterized by rather conspicuous and sharp markings

uph; ground colour unh greenish or greenish-yellowish, not dark. According to Kauff-

mann (1951), this form is more common at Jow than at high altitudes, just as can be

excepted from a well-marked form.

(€). prabornius Kauffmann, 1951. Above black, with small spots upf and inconspicuous

white streaks uph; unf very pale; unh ground colour rather homogenous pale yellow,

spots large and coherent; of average size, g° 13—13.7 mm. This is, briefly, the original

description of prabornius. Kauffmann states that it occurs in the “Oberes Nikolaital”

in Valais. I have seen such specimens also from other localities, particularly the Gross

Glockner region.

(2). SE Europe. Warren (1926) proposed the name trebevicensis to replace reverdini

Schawerda, 1918, the latter being a homonym of reverdini Oberthür, 1912 (see below).

However, Warren had not seen specimens of this form and relied entirely on the original

description of Schawerda and the statement by Reverdin that it is a very striking variety.

I do not know whether Warren saw this form in 1952, but at that time he stated that

it occurs “from Turin to the Altai Mountains, and the Carpathians to Macedonia”

(p. 95). The original description reads (Schawerda, 1918: 23): “Etwas grösser, oberseits

heller braun, nicht so dunkel wie die Nennform, normal stark weiss gefleckt. Unterseits

sind die weissen Flecke grösser, das Braun lichter, Breitere Flügelform. Flügelspannung

meist 29 mm. 5 g', 2 9, Trebevic, Vucijabara. Von mir im Juli 1907 und 1912 erbeu-

tet.” This has little value, in view of the great variability of ssp. alveus and, indeed,

I have not found any reason to separate Yugoslavian specimens subspecifically from

specimens from elsewhere within the range of ssp. alveus, as outlined above. By giving

trebevicensis the rank of subspecies one would reduce the nominate form almost to the

rank of an aberration.

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

(3). Caucasus - Transcaucasia - N Iran. The populations of this region have long been

known under the name of sifanicus, originally given to Chinese specimens, on account of

the white borders unf and unh and the large spots unh in the few specimens known

(mainly from Transcaucasia). However, Alberti (1967, 1969) showed by his own

captures that the usual form of the Caucasus is indistinguishable from specimens from

S Germany. Indeed, six males taken in the Caucasus and sent to me by Alberti appeared

to be similar to specimens from S Germany. I saw only one specimen from the Caucasus

(Kuban district, MP) with sifanicus characters. South of the Caucasus there appear to

be proportionally more specimens of the sifanicus type. Alberti (1967) found among

seven males and five females, five males of the alveus type and among the material

examined by me (9 & 2 Q) there were also five males of the alvews type, the other

specimens resembled in varying degrees ssp. sifanicus. The single specimen from N Iran

(“Hyrcania”) is small (12.6 mm), but otherwise like ssp. stfanicus. It is possible that

there exists a genetic-historical relation between the Transcaucasian population and the

Chinese ssp. sifanicus. As a subspecies is a population (or a group of populations) and

not a character, it seems at present desirable to refer the populations of the whole

region, from the Caucasus to N Iran, to ssp. alveus, of course without forgetting the

variation in this region.

I have found white borders at the underside also (rarely) in ssp. numidus, in the

accretus subspecies group and (very rarely) in Central European ssp. alveus.

Pyrgus alveus centralitaliae Verity

Verity, 1920 — Ent. Rec. 32: 4.

Type-locality: Sibillini Mts.

Material examined. — 74 & 28 9 C Italy (Marche, Sibillini Mts.) (BM, RJ).

Distribution. — Confined to C Italy.

Subspecific characters. — Usually small, $ 13—14 mm, but as noted by Verity

(1940), sometimes reaching 17 mm. Verity believes that the larger specimens are the

result of humid weather. Many specimens have clear white markings uph, but as far

as I could ascertain, they do not dominate. Ground colour unh usually yellower than in

ssp. alveus.

Pyrgus alveus iliensis Reverdin

Reverdin, 1912 — Bull. Soc. Lép. Genéve 2: 155—157; pl. 16 figs. 7—8; pl. 18 figs. 1—2.

Type-locality: Ili region.

Material examined. — 16 & 8 9:1 g 1 9 Kansk (W Siberia) (BM), 1 &

Boro Choro (BM), 3 & Abakan (MO), 11 g' 7 2 Altai (BM, ML). I examined the

genitalia of all males, except six from the Altai.

Distribution. — W Siberia to the Altai and the Ili region.

Subspecific characters. — Characterized by the very large and square spots above. This

appears to be the usual form of the Ili region (original description; Alberti, in litt.).

The specimens from Kansk and Boro Choro agree very well with it. The other specimens

have smaller spots, but they are better spotted than most European specimens of ssp.

alveus. The size varies from 12.4 to 14.4 mm.

Note. — The union of all populations from the Ili region and W Siberia into a

single subspecies is provisional, as is the place in the alvews subspecies group. As

mentioned above, the scandinavicus type of genitalia also occurs in this region. I found

it in a specimen from Kansk, one from the Altai and two from Abakan.

R. DE JONG: Pyrgus in the Palaearctic region 93

Pyrgus alveus sifanicus Groum-Grshimailo

Pl. 6 Fig. 97, 106

Groum-Grshimailo, 1891 — Hor. Soc. Ent. Ross. 25: 459.

Type-locality: Kuku-Nor.

Material examined. — 9 g 1 9 Kuku-Nor region (Kuku-Nor, Amdo, Sining)

(BM, ZMHB). I examined the genitalia of four males.

Distribution. — Only known from the Kuku-Nor region.

Subspecific characters. — Typically distinguished by the white borders unf and unh.

However, four males out of nine examined were nearly indistinguishable from European

alveus.

Note. — (1). A male in the BM, labelled “E Siberia”, is externally and on account

of the genitalia a typical s7fanicus. Unfortunately, the locality is rather vague.

(2). The male genitalia are very suggestive of European alveus, those in one specimen

being indistinguishable from the European structure, the others having the proximal

edge of the cuiller rather oblique and the apex of the cuiller more or less massive, in

the manner of ssp. speyer?.

Pyrgus alveus reverdini Oberthiir

Pl. 6 Fig. 98, 107

Oberthür, 1912 — Et. Lép. Comp. 6: 72, pl. 137 fig. 1217.

Type-locality: Ta-Tsien-Lou (Szechwan).

Material examined. — 20 & 4 9:14 & 2 9 Ta-Tsien-Lou, 2 $ Chengmengka,

1 8 Ying-kuan-chiai, 1 9 Kwan-chiai (all Szechwan), 1 & Shiuden Gompa (SE

Tibet). 2 g 1 9 How Kow (SE Tibet) (all BM). I examined the genitalia of

four males.

Distribution. — SE Tibet and W Szechwan.

Subspecific characters. — Above very much like ssp. speyerz but spots never as

reduced as sometimes in that form. Unh white spots very extended and with

a conspicuous white band along termen. Ground colour unh of a particular yellow-green

tinge. Central spot unh usually with a projection towards the base of the wing and

towards the termen along vein 6.

Note. — (1). Of the male genitalia examined none has the shape of the genitalia

figured by Warren (1926; pl. 6 fig. 2). They are suggestive of ssp. stfanicus and of

ssp. alveus. Undoubtedly the male genitalia of ssp. reverdini are rather variable, as in

all subspecies of P. alvens.

(2). Ssp. reverdini is possibly confined to very high altitudes. It has been taken

from 10.000 ft (How Kow) to 14.000 ft (Shiuden Gompa), but the specimens from

Ta-Tsien-Lou lack exact data.

There remain two subspecies to be discussed, viz. ssp. scandinavicus and ssp. speyeri.

I have left them out of the alveus subspecies group because their male genitalia are

somewhat different. As stated above, there may exist a large-scale intergradation in

W Siberia (between ssp. scandinavicus and ssp. iliensis and possibly even ssp. alveus)

and in E Siberia and N China (between ssp. speyeri and ssp. sifaricus). Another

interesting region seems N and C Russia, but material is lacking. On the records of

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Pyrgus serratulae from Vologda and Kotlas (Krulikowsky, 1909) and of alveus with

the basal spot in space 7 unh roundish, from Estonia (Petersen, 1924) one may assume

that ssp. scandinavicus is widespread in N Russia. More material, however, is needed

for certainty about this.

The fact that ssp. scandinavicus and ssp. speyeri are discussed here after the alveus

subspecies group does not mean that they are more closely related to each other than to

that group.

Pyrgus alveus scandinavicus Strand

Strand, 1903 — Arch. Naturv. Christian. 25: 6.

Type-locality: Dovre. :

Material examined. — 43 g 10 9: 39 & 9 9 Norway (BM, MO), 4 g 1 @

Sweden (BM, MO). I examined the genitalia of 34 males from Norway and 4 males

from Sweden; of 28 g' from Norway and 1 & from Sweden I saw only the genitalia.

Distribution. — Norway, Sweden, Finland and possibly N Russia.

Subspecific characters. — Rather small, $ 12.4—14.2 mm. Upf spots usually small;

they are rather square when they are well-developed. Uph nearly plain brown. Unh

spots rather large and sharply contrasting with the deep greyish-yellow or greenish

ground colour.

The male genitalia have been discussed above (Chapter 2.b.). In brief it may be

recalled that they are characterized by:

(a) broad stylifer and antistyle; (b) dorsally greater development of the cuiller than

ventrally; (c) backward slope of the free proximal edge of the cuiller, which is one

third to half of the total depth of the proximal edge. The genitalia vary towards the

alveus type (cf. Opheim, 1956).

Pyrgus alveus speyeri Staudinger

Pl. 6 Fig. 99, 108

Staudinger, 1887 — in Romanoff, Mém. de Lép. 3: 153; pl. 8 fig. 5.

Type-locality: Amur.

Material examined. — 45 & 21 9:1 g E Altai, 1g 1 9 E Sajan, 1 & Irkutsk,

3 & 1 9 Kentei Mts, 1 9 Viluisk, 1 g 1 9 Chita, 16 d 8 2 Jablonowyj Khrebet

(all BM), 18 $ 8 9 Amur (BM, ML), 3 & 1 @ Little Chingan (BM); 1 8 Kuldja

(ML). I examined the genitalia of 15 males.

Distribution. — From the E Altai to the Amur region. The male examined from

Kuldja (upper course of the Ili River) is a normal speyer? without any trace of zliensis.

If correctly labelled, this specimen may indicate a distributional overlap with zlienszs.

Subspecific characters. — Externally easily distinguishable by the reduced spots upf

which are often partly lost, by the greenish-yellowish superscaling above, by the uniform

deep dirty yellow ground colour unh and by the rather reduced spots and the usually

roundish or oval basal spot in space 7 unh.

As stated above (Chapter 2.b) the male genitalia are very variable. The cuiller is

ventro-distally often flattened, causing a scandinavicus-like appearance. The proximal

edge of the cuiller is sometimes perpendicular on the ventral line of the valve, but

R. DE JONG: Pyrgus in the Palaearctic region 95

usually it is distinctly oblique; the free part of the proximal edge is more than one

third the total depth of the proximal edge in all specimens examined. The apex of the

cuiller is massive and strongly dentate; it usually rises well above the dorsal ridge of

the harpe, and in some specimens it is very conspicuous, being highly raised. The

harpe is usually rather narrow and of more or less even width.

Geographic history. — A part of the geographic history of P. alveus has been outlined

in Chapter 2.d, in particular the origin of the subspecies numidus and speyeri and the

accretus group. Only additional notes are given here on the accretus and alveus groups

and on ssp. scandinavicus.

(a) accretus subspecies group. — The differentiation of ssp. jurassicus is undoubtedly

of a postglacial age as it seems impossible to refer this form to a glacial refugium other

than that of ssp. accretus (presumably N Spain). The fact that ssp. accretus looks more

like Pyrgus bellieri than ssp. centralhispaniae may be due to a contact between ssp.

accretus (or rather its precursor) and P. bellieri during the Third Interglacial. The

bellieri-like specimens sometimes found in the Pyrenees may be extremes of the accretus

variation or may indicate a postglacial contact between ssp. accretus and P. bellieri in

the Pyrenees.

During the Last Glacial ssp. centralhispaniae apparently originated in a Central

Spanish refugial area or at least in an area south of and more or less isolated from where

ssp. accretus originated.

The glacial history of ssp. insigniamiscens (Sierra Nevada) is uncertain, just as its

origin. At least its morphologic origin appears rather clear, as its male genitalia seem

intermediate between the subspecies numidus (NW Africa) and centralhispaniae (C

Spain).

(b) alveus subspecies group. — This may have originated from the form of the

Second Interglacial which also gave rise to the precursor of the accretzs group and to

Pyrgus bellieri during the Third Glacial. If this is true, the precursor of the a/vews group

must have originated in an eastern refugium during the Third Glacial, but the location

of this refugium is obscure (E Europe?, Asia?). The fact that both the Chinese ssp.

sifanicus and the Transcaucasian alveus often have white borders unf and unh may

indicate a close relationship. It is not impossible that they are the remnants of an eastern

form (the ancestor of the a/veus group) of the Third Interglacial which have been

driven away to border regions. Ssp. alveus postglacially largely extended its range from

its refugium in SE Europe to the Altai and to the Caucasus region, where it nearly

entirely replaced the s/fanicus-like form in the northern range (the Caucasus proper);

ssp. alveus replaced that form in the southern range only partially.

The occurrence of a slightly differentiated form in C Italy (ssp. centralitaliae) seems

to date from a colder period than the present one. Likely, there was a more or less

continuous refugial area for ssp. a/vews during the Last Glacial from Italy to the Balkans.

Ssp. centralitaliae may be a postglacial geographic isolate of ssp. a/veus.

The origin of ssp. iliensis and its intergradation with other forms must remain

obscure as long as the material available from W Siberia and N Turkestan is so extremely

scarce.

The geographic origin and history of ssp. reverdini is also uncertain. Its distribution

suggests that it is an ancient form, but it may also be regarded as a rather recent des-

cendant of ssp. sifanicus.

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

dinavia as in W Siberia. It is therefore unlikely that ssp. scandinavicus is merely a post-

glacial form of ssp. alveus. As the greater part (if not the whole) of its present distribu-

tion area was covered by land ice during the Last Glacial and as the southern European

refugial areas had been taken by other forms (or by their ancestor), ssp. scandinavicus

must have colonized Scandinavia postglacially from the east (E Russia or W Siberia).

Little can be said about the history of ssp. scandinavicus, because of the scarcity of

specimens available from W Siberia, but the general shape of the cuiller seems to indicate

an influence of ssp. speyeri. It would not be surprising if ssp. scandinavicus turned out

to be a hybrid between the subspecies speyeri and alveus (or iliensis). In this connection

it is interesting to note that Warren (1953) found a remarkable variation in five out of

twenty-seven specimens of ssp. scandinavicus, viz. the loss of the apex of the cuiller, “as

if a triangular piece had been cut off, from a point half-way down the free edge to one

a similar distance from the apex on the dorsal ridge, leaving a straight line connecting

two sharp angles.” (p. 101).

Indeed, I saw this remarkable form of cuiller in ssp. scandinavicus only, but it is

impossible to give a percentage of specimens affected, as the cuiller is highly variable.

This “lost apex” appears to be the massively sclerotized apex of ssp. speyert.

19. Pyrgus bellieri Oberthür

Oberthür, 1910 — Et. Lep. Comp. 4: 404; pl. 56 fig. 490.

Type-locality: Larche (Basses Alpes).

Distribution (Map 17). — SE France, NW Italy, C Italy. In France it has been taken

in the following departments: Aveyron, Lozere, Gard, Ardeche, Dröme, Vaucluse, Bou-

ches-du-Rhöne, Var, Alpes Maritimes, Basses Alpes, Hautes Alpes (Picard, 1949b). In

N Italy confined to the Cottian and Maritime Alps, in C Italy in the mountains of

Emilia, Marche, Abruzzi and Latium (Verity, 1940; Picard, l.c.).

As stated above (under P. alveus accretus), some specimens with the bellieri type of

genitalia have been taken in the Pyrenees, although apparently flying within the range

of Pyrgus alveus accretus which resembles P. bellieri very much. As long as the existence

of P. bellieri populations in the Pyrenees has not been established, I prefer to regard such

bellieri-like specimens from the Pyrenees as extreme variations of Pyrgus alveus accretus.

Habitat. — Dry, flowery slopes, from 100 to 1700 m (Verity, 1940; Guillaumin,

1964). At Digne (Basses Alpes) I found P. bellieri in rather large numbers on mud

along a little stream, together with innumerable specimens of the Lycaenid Lysandra

coridon.

Biology. — Monovoltine; flight period July-August. Larvae and food plants not

known.

Geographic variations. — As expected, the geographically isolated C Italian population

is somewhat different from the other populations and there is nothing against its

separation as a subspecies.

In its restricted range in SE France and NW Italy bellieri is rather variable and it has

been divided into five “subspecies’” by various authors (see Picard, 1949b). I regard

them as ecophenotypic variations, as these forms are mainly confined to particular

altitudes and very restricted ranges.

R. DE JONG: Pyrgus in the Palaearctic region 97

Pyrgus bellieri bellieri Oberthür

Material examined. — 221 d 109 9:2 g 2 Q Aveyron (BM, ML), 9 & 2 2

Lozère (BM), 3 g' 2 9 Vaucluse (ML), 23 & 6 © Bouches-du-Rhône (BM, ML),

53 & 28 Q Var (BM, ML), 24 & 15 9 Alpes Maritimes (BM, RJ), 102 & 51 9

Basses Alpes (Digne, Larche) (BM, RJ, HC), 5 & 3 9 Hautes Alpes (BM).

Distribution. — As that of the species, except C Italy.

Subspecific characters. — A variable subspecies, & 12.5—15.5 mm; usually larger

than ssp. picenus, with smaller white spots and less strongly white suffusion above. Much

variation, however, occurs.

Variation. — Two “races” have been described which are confined to higher altitudes

(above 1100 m), viz. nigropictus Verity, 1926 (type-locality: Cottian Alps) and bellieri

Oberthiir, 1910 (type-locality: Larche). The latter is characterized by its rather small

size, the very strong suffusion of white scales above and the large white spots unh. It

occurs as well in the French as in the Italian Alps. According to Picard (1949b) it is

the “race” of the highest altitudes. It is the only form recorded from Larche. Some

specimens taken at Larche are difficult to distinguish from the lowland f. foulguiert and,

vice versa, some specimens from the lowlands (Septèmes, St. Zacharie) resemble f. be/-

lieri very closely.

The form nigropictus is still smaller than f. bellieri, above with a rather strong white

suffusion, unh with a strong suffusion of dark scales and with smaller white spots than

in f. bellieri. According to Picard (1.c.) it is confined to the Alps (French and Italian)

where it occurs at altitudes higher than 1100 m, but at lower altitudes than f. bellieri.

Especially the darkened ground colour unh occurs (very infrequently) also elsewhere

(Var, Bouches-du-Rhöne, Vaucluse).

The other forms, described as races, are: foulguieri Oberthür, 1910, adkini Picard,

1948, and gaillardi Picard, 1948. The last-mentioned form, of which I have seen no

specimens, is pale and small and is only known from a few dry and hot localities in

Gard and Ardeche.

The form adkini is mainly characterized by the very large white spots upf and the

bright spots uph. According to Picard (l.c.) it is the race of the ‘‘Causses’’ (Lozère,

Aveyron). The specimens I have seen from that region do not allow a subspecific

separation.

The form foulguieri is variable in size (4 12.5—15.5 mm), extension of spots and

white suffusion above and extension of spots and ground colour unh. It is difficult (if

at all possible) to distinguish this form from the other forms of the subspecies except

from extreme f. bellieri. Originally bellieri and foulquieri were described as separate

species.

Pyrgus bellieri picenus Verity

Verity, 1920 — Ent. Rec. 32 4.

Type-locality: Sibillini Mts.

Material examined. — 32 & 25 9: 26 g 24 9 Sibillini Mts. (BM), 1 & Gran

Sasso (ML), 3 & Aquila (ZMA), 1 g' Sabini Mts, 1 & Roccaraso, 1 Q Sorrento

(BM).

Distribution. — C Italy.

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

Subspecific characters. — Small, $ 12-14 mm. With regard to the upperside and

underside more or less intermediate between the forms foulguieri and bellieri of sub-

species bellieri: upf with stronger white suffusion than in f. foulguieri, in

some specimens as strong as in f. bellieri, uph white spots larger than in f. foulguieri

and unh white spots varying from the size found in f. foulguteri to the very broad spots

typical of f. bellieri.

Geographic history. — As the present morphologic differentiation of P. bellieri

appears to be of a postglacial age, nothing can be added to what has been said on the

history of this species in Chapter 2.d.

20. Pyrgus warrenensis Verity

Pl. 4 Fig. 62, 71

Verity, 1928 — Bull. Soc. Ent. France 1928: 140—141.

Type-locality: Lenzerheide (Grisons) (Warren, 1929).

Distribution (Map 18). — Widely distributed in the Alps, but apparently a local

species. As fas as I know only taken at the following localities: France: Larche, St. Veran,

Mt. Cenis, Pralognan (Warren, 1953; Bretherton, 1966; Material examined); Switzer-

land: Tasch, Zermatt, Ryffel Alps, Lenzerheide (Warren, l.c.; Material examined); Italy:

Oulx, Valtournache, Ortler Alps (Warren, l.c.; Material examined); Austria; Samnaun

Gruppe, Vent, Brenner Pass, Gross Glockner, Mallnitz (Warren, l.c.; Prose, 1955b;

Material examined): S Germany: Allgäu, Berchtesgadener Alpen (Pröse, l.c.). Many

other localities of this high altitude species can undoubtedly be found.

Habitat. — Alpine meadows; presumably not below 1700 m; the upper limit is not

known, but specimens have been taken at least at 2500 m.

Biology. — Monovoltine; flight period July—August, according to local conditions.

Larvae and food plants not known.

Geographic variation. — Geographic variation has not been established.

Material examined. — 108 8 62 9: 37 8 15 @ Basses Alpes (Larche), 1 &

Savoie, 2 & Mt. Cenis, 5 & 2 9 Ryffel Alps, 5 g 2 9 Grisons, 3 4 Piedmont,

15 4 8 9 Valtournache (all BM), 5 4 1 9 Ortler Alps (Franzenhöhe), 2 g° Ötztal

(Vent) (ZMA), 33 & 34 9 Gross Glockner (BM, ML, RJ).

Geographic history. — See Chapter 2.d.

21. Pyrgus jupei Alberti

Pl. 4 Fig. 63, 72

Alberti, 1967 — Deutsche Ent. Zeitschr. 14: 464—465; pl. 1 figs. 1 a—b; pl. 2 figs. 1 a—d;

pl. 3 figs. 2 a—f.

Type-locality: Kasbek (C Caucasus).

Distribution (Map 18). — Caucasus and Transcaucasia. Very few localities are known

at present, but undoubtedly P. jupei is more widely distributed in this rather poorly

explored region.

The localities known at present are: Caucasus: Tschutschur Pass, Kasbek, Karaugom,

Passanauri; Transcaucasia: Achalzich, “Chotschalj”, Borshom (Alberti, 1967, 1969, in

litt.; Material examined).

Habitat. — Alpine meadows, 1700—2700 m.

R. DE JONG: Pyrgus in the Palaearctic region 99

Biology. — Unknown. Flight period presumably mainly in July.

Geographic variation. — Unknown.

Material examined. — 6 d 1 9:1 & Kasbek (BA), 4 g' 1 9 Passanauri (BA,

RJ), 1 & Borshom (BM).

Geographic history. — See Chapter 2.d.

22. Pyrgus schansiensis Reverdin

Reverdin, 1915 — Bull. Soc. Lép. Genéve 3: 109—112; pl. 5 fig. 3, pl. 6 fig. 5.

Type-locality: Ta-tsing-schan (Shansi).

Distribution. — Apart from the type-locality only known from Kinschou (Fengtien,

Manchuria) (Alberti, 1940) and the Stanovoy Khrebet (RJ).

Habitat, biology and geographic variation. — Unknown. It should be born in mind

that the specific rank of schansiensis is disputable.

Material examined. — 1 G' Stanovoy Khrebet (RJ).

4. DISCUSSION OF SOME ZOOGEOGRAPHIC PROBLEMS

4.a. The geographic origin of species and groups of species

Any attempt to reconstruct the geographic history of a species eventually leads to the

question: where did this species come into being. If we define speciation as the process

by which a population (or group of populations) acquires reproductive isolation, the

geographic origin of a species evidently is the region where the speciation took place,

irrespective of subsequent extensions of the range or further differentiation. Matters

become complicated when through geographic isolation a further differentiation has

led to species multiplication. Virtually one of the new species will resemble the

parental species more closely than the other new species and can be regarded as the

temporal representative of the parental species. Alberti (1955b) describes this as

follows: “Vom theoretischen Grenzfall abgesehen, wird praktisch immer mehr oder

weniger lange die Mutterart neben der Tochterart existieren.” (p. 218). The problem

whether one considers the parental species to have become extinct or still to exist after

species multiplication, is mainly of phylogenetic importance( cf. also the discussion on

the species in time dimension in Hennig, 1966), but it has also a bearing on the

denomination of the geographic origin of a species. If we consider the parental species

still to exist after its multiplication, there is no obvious reason to suppose that it still

exists in the region where it originated. This region may be populated by what we

regard as a new species, instead of the continuation of the parental species. In this

case the parental species and the new species have the same geographic origin. If

fossils are absent, we can make a distinction between a parental species and a new

species only on the basis of a difference in the frequency of characters which are sup-

posed to be primitive, as the parental species must have retained more primitive

characters than the new species.

The above considerations also apply to the geographic origin of subspecies (if we

define the subspecies as an actually or formerly geographically isolated group of

populations, see Chapter 1.b).

In the reconstruction of the geographic history of the centanreae species group

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

(Chapter 2.d) it was supposed that a repeated speciation had taken place in Siberia.

This led to the statement, that all three species of the group had a Siberian origin,

although two of them (P. andromedae and P. cacaliae) at present are restricted to

European mountains. Of the third species of the group (P. centaureae) the European

and American subspecies seem to have originated in Siberia, just as the Siberian sub-

species did in a more recent time. These considerations led to the statement, that

from the phylogenetic point of view the difference between a European and a Siberian

origin in cold-adapted species is gradual: it depends on the number of primitive

characters, retained in the European and the Siberian populations, respectively. This

is only another way of stating that the assignment of a species as parental species depends

on the number of supposedly primitive characters, but it shows the bearing of phylo-

genetic considerations on zoogeographic classifications (see Chapter 5).

With respect to the total distribution area of the centaureae group, the distributional

areas of P. andromedae and P. cacaliae are peripheral. In the second part of this chapter

it will be shown that the occurrence of primitive characters in peripheral populations

does not appear exceptional. It may suffice here to call attention to the possibility that

the geographic origin and the present distribution area of a species or subspecies lay far

apart.

In systematic studies one can often find such general statements as “genus A-us is of

European origin” or “family B-idae is of Palaearctic origin”. For example, while

discussing the Pyrgus group of genera, Evans (1953) stated: “The group is of Palaearctic

origin with a considerable development in the Nearctic and Neotropical region” (p. 2).

Evidently the authors of such statements do not bother about the meaning of the

expression “the geographic origin of a genus” or of whatever group of species. Unlike

the origin of a species, the origin of a genus has no time dimension. Although a genus

may be a monophyletic, quite natural group of species, one cannot indicate a moment or

period in which the genus became generically distinct from other genera. Virtually a

genus originates in the brain of an author, as it is a subjectively delimited group of

species. Apparently the brain of an author is not meant by “the geographic origin of

a genus”. The geographic origin of a group of species can be defined as the region where

the differentiation took place which led to this particular group of species. This is a very

vague definition, but one should realize that the geographic origin of a group of species

is something quite different from the geographic origin of a species.

Possibly some authors mean by the geographic origin of a group of species the

geographic origin of the ancestor of the group, whatever this may be. I can understand

the statement of Evans mentioned above only in this way. The way a definition reads, is

less important, so long a definition is provided.

4.b. The geographic history of the Palaearctic Pyrgus species in relation

to the general history of the Palaearctic fauna

The occurrence of species and subspecies with primitive characters in border regions,

as indicated in the first part of this chapter, appears to be a widespread phenomenon in

the Palaearctic fauna. The fact of a character being primitive often is very uncertain.

In some cases it is concluded from morphologic data, as in the centaureae species group

and in superspecies malvae (Chapter 3). In other cases it is concluded from the distri-

butional discontinuity of the character, if this character is supposed to have no adaptive

R. DE JONG: Pyrgus in the Palaearctic region 101

value (e.g. characters of the genitalia). Consequently, the occurrence of primitive

characters in border regions (in a broad sense) can be accepted in Pyrgzs in the following

cases:

(1). The centaureae species group. Widely distributed in the colder parts of the Holarc-

tic; primitive characters in European mountains (P. cacaliae and P. andromedae).

(2). Superspecies malvae. Distributed throughout the greater part of the Palaearctic;

most primitive characters in the W Mediterranean region (P. malvoides), less primitive

characters in the E Mediterranean region (P. melotis) and E Asia (P. malvae kauff-

manni), most recent characters in the intervenient area (P. malvae malvae).

(3). The alveus species complex. Distributed throughout the greater part of the Palaearc-

tic; primitive characters still very uncertain, but a possible distribution of such characters

can be described as follows: NW Africa (P. alveus numidus), Alps (P.warrenensis),

Caucasus (P. jupei) and E Siberia (P. alveus speyeri).

It is difficult to trace this phenomenon in the literature concerning other families of

Lepidoptera, as this literature usually is merely taxonomic and phylogenetic reflections

are left out of consideration. Possibly the following distributional patterns concern the

occurrence of primitive characters in border regions:

(4). Papilio machaon (Papilionidae; Eller, 1936—1938) is distributed throughout the

Palaearctic; P.machaon saharae (NW Africa) and the closely related P. hospiton

(Corsica and Sardinia) are supposed to be primitive forms.

(5). The Melitaea phoebe group of species (Nymphalidae; Higgins, 1941) is widespread

in the Palaearctic; M. aetherie (S Spain and NW Africa) and M. collina (E Mediter-

ranean) are possibly more primitive than the other species of the group.

(6). The Exphydryas aurinia group of species (Nymphalidae; Higgins, 1950) is

widespread in the Palaearctic; E. desfontainii (W Mediterranean) and E. orientalis (E

Mediterranean) may be more primitive than the other species of the group.

In addition, three instances quoted from zoogeographic studies on forest birds are:

(7). The Woodpeckers of the Dendrocopos major group are distributed throughout the

Palaearctic. The presumably most primitive members of this group, D. catphartus and

D. darjellensis, are restricted to SE Palaearctic border regions: chiefly the Himalayas to

SW China (Voous, 1947).

(8). The genus Pyrrhula (Bullfinches) occurs in the Palaearctic region and in the

Oriental region in the Himalayas and to the southeast as far as Formosa, the Philippines

and the Malayan peninsula. P. nipalensis is considered the most primitive form; it occurs

from the W Himalayas to NW Yunnan and in a few isolated ranges further south and

southeast (Voous, 1949).

(9). The Nuthatches of the Sitta europaea group, distributed throughout the Palaearctic

and part of the Oriental region, are supposed to have descended from the S. canadensis

group, which has a highly discontinuous distribution: Corsica, Asia Minor and Caucasia,

E Asia and N America. S. canadensis probably invaded America from E Asia. S. cana-

densis yunnanensis (Yunnan, W Szechwan) is the most conservative of the living forms

of the S. canadensis group (Voous & van Marle, 1953).

The above instances concern groups with a very wide distribution in the Palaearctic.

At least in Pyrgus the occurrence in peripheral regions of primitive forms of species or

species groups with a restricted range (e.g. European) is not apparent (see also below).

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

As one would expect to find primitive forms mainly in regions which have remained

rather undisturbed by ecological changes since long, it is not surprising that most of the

primitive forms mentioned above occur in regions which are known to have been refugia

for the Palaearctic fauna of temperate regions during the Pleistocene glaciations

(Reinig, 1937; De Lattin, 1951-52, 1967). Evidently the forms, which were most

successful in reoccupying regions, which were uninhabitable during glacial times, are the

most advanced forms. Species and species groups with a restricted range and without

primitive forms in border regions may be primitive themselves, at least when they can be

regarded as rather close relatives of species with a Palaearctic distribution. In Chapter

2.d, Pyrgus cinarae and P. armoricanus, both restricted to the SW Palaearctic, have

been considered to be the most primitive members of the alvexs species complex, which

has a Palaearctic range, on the ground of their morphologic characters and habitat

preference. On similar grounds P. carthami and P. sidae, both restricted to the SW

Palaearctic, have been considered to be older than the widely distributed P. centaureae

group. Further there is morphologic evidence for the supposition that the Pyrgus species

which are restricted to the SE Palaearctic (e.g. P. maculatus, P. oberthuri) are the most

primitive of the Palaearctic Pyrgus species.

In conclusion, it appears to be a widespread phenomenon in the Palaearctic fauna

that in groups with a Palaearctic distribution the most primitive members have restricted,

more or less peripheral ranges. The most important factors leading to this distributional

pattern possibly are:

(1). The Pleistocene glaciations. During glacial periods all species in the Palaearctic,

which were not adapted to a cold climate, were confined to refugial regions, situated

south of the land ice. These refugia were situated more peripherally than centrally in

the Palaearctic.

(2). Differentiation during geographic isolation. Formerly connected populations could

differentiate during geographic isolation in refugia. Some populations retained more

primitive characters than others.

(3). The most advanced forms were most successful in re-occupying lost territories.

Possibly the glacial refugia for these forms were situated more closely to the land ice

than the refugia of the more primitive forms. This would explain the loss of primitive

characters (by high selection pressure) and give the advanced forms a geographically

(and perhaps ecologically) favourable starting position for the re-occupation of the

Palaearctic.

The particular distributional pattern of primitive and advanced forms in the Palaearc-

tic appears to contradict the statement of Mayr (1963): “the “original” phenotype of a

species is usually found in the main body or central part of a species range, while the

peripheral populations, particularly the peripherally isolated populations, may deviate

secondarily in various ways’ (p. 386). Mayr did not take the geographic history of

the species into account in this case. In the instances mentioned above the “original”

phenotype of the species in the central part of the range has become extinct by the

glaciations and the peripheral populations which deviated most from the “original”

phenotype, re-occupied the lost territories after the retreat of the land ice. These

expending populations may have attained reproductive isolation with regard to formerly

conspecific, at present more primitive populations. In many cases no populations were

available for the re-occupation of the Palaearctic in postglacial times, possibly as a result

of specialized ecological demands or of the deteriorating effect of the succession of four

R. DE JONG: Pyrgus in the Palaearctic region 103

glacial periods. In these cases the species are relics in their present, restricted ranges and,

of course, the present area of distribution is no indication of the geographic origin.

After this general discussion some aspects of the history of two types of fauna will

be treated, as far as it may be useful for the understanding of the history of the Pyrgus

species.

Cold-adapted fauna. — In his interesting work on the history of the cold-adapted

Holarctic fauna, Petersen (1954) pointed to the absence of endemic genera or even

subgenera in the Arctic butterfly fauna. He attributed this absence to the great exchange

of forms during the Pleistocene between the Arctic and mountains of the temperate

zone. Apart from this exchange, the Arctic fauna may be too young for the

development of endemic genera. The typical tundra fauna appeared in C Europe first at

the end of the long Second Interglacial, although such cold-adapted animals as Tichorhi-

nus, Rangifer and Ovibos have been found as early as the Second Glacial (Petersen, p.

274).

Both the exchange between the Arctic and mountains of the temperate zone and the

late appearance of cold-adapted species in Europe are indicated by the distribution and

supposed history of the species of the Pyrgus centaureae group (P. centaureae, P. cacaliae,

P. andromedae). This history has been discussed extensively in Chapters 2.d and 3,

and can be summarized here as a series of three invasions from Siberia into Europe. The

most ancient invasion, which can be traced at present, took place during the Second

Glacial (P. andromedae). The P. centaureae group has a boreoalpine distribution in

Europe; P. centaureae is possibly continuously distributed in Asia from the S Siberian

mountains (Altai, Sajan) north to the Arctic region. A list of Lepidoptera with a

boreoalpine distribution can be found in Warnecke (1959).

The other cold-adapted Pyrgus species in Europe, viz. P. carlinae, P. warrenensis and

P. jupei, are supposed to have originated during the Pleistocene from more temperate

or eurytherm species in Europe. Many cold-adapted species, which are at present

restricted to European mountains, may have originated in this way, or else, they may be

descendants of such species. Kostrowicki (1969), discussing the Papilionoidea of the

European mountains stated that “the tundra species did not take part in the formation

of the fauna of the Alpine tiers in European mountains, and species now living in that

tier came into being from autochtonous original forms.” (p. 207). Although he appears

to have overlooked the exchange of cold-adapted species between the European and

Asiatic mountains through the Arctic, he may be correct in supposing a European origin

for by far the greater part of the cold-adapted species of European mountains. I have

not found any indication that these species are older than of Pleistocene age. Warnecke

(1958) suggested that the species of the butterfly genus Erebia are much older:

“During the eocene, with the formation of the last high mountains (especially those of

the Alps in Europe), there has obviously been another transformation and evolution of

alpine species, for instance those of Erebia.” (p. 724). More than half of the about

77 species of Erebia (Warren, 1936; Higgins & Riley, 1970) are confined to European

mountains. Warren classified the Erebia species into 14 species groups, of which only

one, viz. the pronoe group (10 species), is restricted to European mountains. As far

as the other groups have representatives in Europe, they have also representatives in

Siberian mountains, thus indicating the exchange of forms between European and

Asiatic mountains. This exchange has been possible only during the Pleistocene and

hence European species belonging to groups with representatives in Siberia possibly

originated during the Pleistocene.

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

As the present European cold-adapted species appear to be of a rather recent,

Pleistocene age, the question arises whether there has been a European cold-adapted

fauna in the Tertiary. In Pyrgus there are no indications of such a fauna. If there have

been Pyrgus species in the alpine zone of European mountains in the Tertiary, these

were probably eurytherm, also occurring at lower altitudes. The present cold-adapted

Asiatic Pyrgus species, all belonging to the subgenus Sce/otrix, on the other hand, may

be of a Tertiary age or at least, they have retained more primitive characters than the

cold-adapted species which originated in Europe (see also Chapter 2.d).

It is improbable that the European alpine fauna in the Tertiary consisted of eurytherm

species only. The ancestors of some Erebia species groups possibly were cold-adapted

European Tertiary species, but indications are still vague. In other genera of butterflies

there are no indications of a cold-adapted European fauna in the Tertiary, but in the

geometrid genus Psodos the Tertiary ancestors of the six species groups, which all

except one are limited to European mountains (cf. Petersen, 1954), may have been

cold-adapted European species.

Temperate fauna. — Some problems concerning the history of the temperate fauna

of two regions, viz. the E Palaearctic and Europe, will be discussed now.

(a). East Palaearctic. The wealth of species, many of which are considered old

(‘‘arctotertiary’’) has tempted some authors to consider this region a centre of origin

of a large part of the Eurasiatic fauna (e.g. De Lattin, 1951-52). In my opinion this

conception is incorrect. As stated above, a large number of ancient species in a particular

region indicates that this region has been little disturbed by ecological changes since

long, but can tell us little about the geographic origin of the species. If the preglacial,

temperate fauna was rather homogenous throughout the Palaearctic (De Lattin, 1.c.), the

large number of old, temperate species in the E Palaearctic region indicates the extinction

of many species in the W Palaearctic. This extinction appears to have been most im-

portant among species living on trees. While discussing the edaphic structure of the

species of Papilionoidea in the faunas of the northern Mediterranean coast, Kostrowicki

(1969) stated: “The share of species living only on trees and shrubs is insignificant

in the above faunas; it is almost identical with the remaining faunas of the province.

So the problem arises what happened to the old Tertiary sylvatic fauna, the traces of

which are so distinct in the Far East?.... it may be assumed that the sylvatic Tertiary

species became extinct and only those have been preserved which live on herbs (and

also ubiquitous species)” (p. 214). As far as known, the Pyrgus species live on herbs

(and shrubs?) only. If there has been any extinction in the Palaearctic of a part of

Pyrgus, it has been largely surpassed by differentiation, as there are at present more

temperate Pyrgus species in the W Palaearctic than in the E Palaearctic. Virtually, only

one temperate Pyrgus species is restricted to the E Palaearctic, viz. P. maculatus. As

this may be one of the most ancient Palaearctic Pyrgus species, the present distribution

is hardly an indication of the geographic origin. So we can only state, that P. maculatus

has a Palaearctic origin.

(b). Europe. According to Kostrowicki (1969), 372 species of Papilionoidea have

been recorded from the region covered by N China, the Amur district, Korea, Sakhalin

and Japan; these species, as a whole, have an outstanding temperate character. From the

W Palaearctic (west of the Russian frontier and the Bosphorus) 338 species of

Papilionoidea have been recorded, of which less than 200 can be called temperate

(cf. Higgins & Riley, 1970). Therefore, the number of eight temperate Pyrgus species

R. DE JONG: Pyrgus in the Palaearctic region 105

in Europe (carthami, superspecies malvae, onopordi, semispecies cirsii, serratulae, armori-

canus, alveus and bellieri) appears to be exceptionally large, as there are only three or

four temperate Pyrgus species in the E Palaearctic (maculatus, superspecies malvae, alveus

and possibly, schansiensis). The absence of six temperate Pyrgus species in the E

Palaearctic does not appear to be the result of extinction in that region. Apart from

P. carthami, these species are supposed to be of a recent, possibly early Pleistocene

age. Their thermophilous character may have prevented these species from extending

the distributional area far to the east during the Pleistocene. Only species which can live

in a cool temperate climate (in Pyrgus : alveus and superspecies malvae), extended

their range during the Pleistocene over the western and eastern part of the Palaearctic.

P. carthami, however, may be an ancient species (cf. Chapter 2.d) and its European

origin 1s not quite certain.

Although the temperate Pyrgus species differ much with regard to their habitat

preference, it is difficult to make a sharp distinction between warm temperate species

of lowlands and rather cool temperate species of mountainous regions, as there is much

overlap in the habitats. It is also difficult to make a sharp distinction between warm

temperate and Mediterranean species. The thermophilous P. onopordi, P. armoricanus

and P. cirsii occur in regions with Mediterranean maquis and steppe vegetation, as well

as in less dry regions. There is no objection against their denomination as Mediterranean

species, but I prefer to use that term only for species which are mainly or entirely

confined to maquis or steppe vegetation. Therefore, of the Pyrgus species, only P. cinarae

and P. sidae are termed Mediterranean species here.

All temperate Pyrgus species in Europe occur in Spain, except P. bellieri, which has

a restricted range in Italy and France. However, only three of these species, viz. P.

onopordi, P. armoricanus and P. alveus, also occur in NW Africa. This is in accordance

with the general view that there was no land bridge over the Mediterranean at any time

during the Pleistocene and that mainly flying animals were successful in crossing the

Strait of Gibraltar (Moreau, 1955). Evidently the chance of a successful crossing of the

Strait of Gibraltar has not been small for butterflies. Of the about 140 non-cold-adapted

species of Papilionoidea in Spain, 81 are represented in NW Africa; they outnumber

the tropical species in that region largely (cf. Higgins & Riley, 1970). Of the 19 species

of Hesperioidea, which occur in NW Africa, only three, (Gegenes nostrodamus, G.

pumilio and Borbo borbonica) can be called tropical and nine temperate; the remaining

seven can be regarded Mediterranean species.

5. ZOOGEOGRAPHIC CLASSIFICATION OF THE PALAEARCTIC Pyrgus SPECIES

Many lepidopterists have been engaged in a zoogeographic classification of Palaearctic

(mainly European) species. They have often stated, that a zoogeographic classification

must be founded on the present distribution of species alone (e.g. Rebel, 1932; Amsel,

1939; Wiltshire, 1945). This idea has led to the distinction of many distributional types

and the division of the Palaearctic region into zoogeographic regions. Recently,

Kostrowicki (1965, 1969) divided the Palaearctic area into 43 units (subrealm, province,

region, transition zone) on the basis of an analysis of the distribution of the Macro-

lepidoptera and distinguished 115 “‘areographic elements and subelements”, i.e. distri-

butional types.

As Voous (1963), pointed out this geographic method of classification is a part of

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

the classic zoogeography of P.L. Sclater and A. R. Wallace. It is a merely descriptive,

static method, although often used as a starting point for historical considerations (e.g.

Friese, 1958; De Lattin, 1964; Kostrowicki, 1969).

As the present distribution of species is the result of historical events, it is not clear to

me, why these events should not be taken into account in a zoogeographic classification.

Stegmann (1938) strongly advocated such a dynamic approach. This led him to the

recognition of faunal types, instead of faunal regions. His classification was refined by

Voous (1960), in his zoogeographic classification of the European birds. I will follow

this system in the classification of the Palaearctic Pyrgus species.

The term “faunal type” refers to the characters of the regional fauna indicated. The

characters of a regional fauna result from the particular history of that fauna, its

geographic distribution and the habitats afforded in the range covered (Voous, 1960,

p. 6). A species can be referred to a particular faunal type, if it, by its origin and

history, is a characteristic element of the regional fauna indicated, i.e. if it has taken

part in, or has been subjected to, the geographic and climatological history of that

fauna.

The Palaearctic Pyrgus species can be referred to the following faunal types (the

descriptions of the faunal types, according to Voous (1960), have been added in

parentheses).

Siberian-Canadian (belonging to the fauna of the boreal climatic zone in the Holarctic,

notably in the coniferous forest belt) — P. centaureae.

Palaearctic (belonging to the fauna of the cold, temperate, and subtropical regions of

the northern half of the Old World) — P. maculatus, P. malvae, P. alveus.

European (belonging to the fauna of the temperate and Mediterranean regions of

Europe) — P. carthami (?), P. malvoides, P. serratulae, P. onopordi, P. cirsii, P. armo-

ricanus, P. bellieri.

Mediterranean (belonging to the fauna of the Mediterranean region) — P. sidae, P.

cinarae.

Palaeomontane (belonging to the fauna of the alpine or nival zones of the high

mountains of the Palaearctic) — P. badachschanus, superspecies P. alpinus, P. andro-

medae, P. cacaliae, P. carlinae, P. warrenensis, P. jupei.

The following notes may be added to this survey.

(1). The species P. bieti, P. oberthuri and P. dejeani cannot be classified zoogeo-

graphically, because of lack of data on distribution and habitat preference. Possibly they

can be referred to the Tibetan faunal type (belonging to the fauna of the Tibetan

highlands in the tundra climatic zone; some members of this group have spread into

the alpine zones of central and western Palaearctic high mountains).

(2). P. melotis cannot be assigned to one of the faunal types proposed by Voous.

By its distributional area this species can be called Mediterranean, but with regard to

habitat preference only P. m. melotis can be assigned to the Mediterranean faunal type.

(3). It is possible that P. sidae and P. cinarae could better be listed as elements of

the palaeo-xeromontane fauna (belonging to the fauna of the arid slopes of the low

mountains of the southern Palaearctic; some palaeo-xeromontane elements have, after

the latest glacial period, spread into the dry, sunny slopes of the high mountains).

The difference between the concept of faunal types and that of distributional types

is comparable with the difference between the biological and the typological species

R. DE JONG: Pyrgus in the Palaearctic region 107

concept: it is the difference between a dynamic and a static approach. This does not in

all cases find expression in the denomination, just as the application of the biological

species concept does not necessarily result in an other delimitation of the species than

the application of the typological species concept.

The distributional types are often explained as the result of the influence of the

Last Glacial on the distribution of the Palaearctic species (Friese, 1958; De Lattin,

1964). This gives the distributional types a more dynamic character, but as it only takes

the recent history into account, the faunal type and the distributional type, to which a

particular species may be assigned, does not need to coincide. This is apparent, if we

compare the zoogeographic classification of the Pyrgus species given above with a

classification based on distributional types. Friese (1958) classified the Rhopalocera

(Hesperiidae inclusive) of C Europe into twelve distributional types, to which the

C European Pyrgus species are assigned as follows:

“boreo-altaisch-alpiner Verbreitungstyp”: P. andromedae.

“altaisch-alpiner Verbreitungstyp”: P. cacaliae, P. cirsii.

“atlanto-mediterraner Verbreitungstyp”: P. malvoides, P. onopordi, P. cirsit.

“ponto-mediterraner Verbreitungstyp'': P. sidae.

“euromediterran-vorderasiatischer Verbreitungstyp': P. armoricanus.

“europäisch-westasiatischer Verbreitungstyp': P. carthami.

“euro-asiatischer Verbreitungstyp”: P. malvae, P. serratulae.

“palaearktischer Verbreitungstyp”: P. alvens.

To the four remaining distributional types (‘europäisch, holomediterran, europäisch-

vorderasiatisch, circumpolar”) no Pyrgus species have been assigned.

6. SUMMARY

1. Twenty-two species (including three superspecies) of Pyrgus have been recognized

in the Palaearctic region.

2. An attempt has been made to distinguish between subspecific, clinal and nongenetic

ecophenotypic variation. As a result, many subspecies of earlier authors have been

assigned to clinal and ecophenotypic variation.

3. Clinal variation is presumably widespread in Pyrgzs species, but appears most

noticeable in the semispecies P. c/rsit and P. carlinae.

4. Three superspecies have been recognized, viz. P. alpinus, P. malvae and P. carlinae.

5. The dynamic character of the species and the subjective character of the distinction

of species is particularly apparent in the a/vews species complex.

6. An attempt has been made to reconstruct the geographic history of the genus as a

whole and that of the separate species. This reconstruction bears clearly evidence of

the influence of the Pleistocene glaciations on the distribution and geographic history

of the species.

7. On the basis of the supposed history of the P. centaureae group of species attention

has been drawn to the possibility that the geographic origin and the present distribution

of a species lay far apart.

8. The occurrence of primitive forms in Palaearctic border regions has been discussed

and a tentative explanation of this distributional pattern has been given.

9. The geographic history of the Pyrgus species has been discussed in relation to the

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

general history of the Palaearctic fauna. Pyrgus is noticeable for its relatively large

number of temperate species in the W Palaearctic.

10. A zoogeographic classification into faunal types has been applied to the Palaearctic

Pyrgus species. These species can be assigned to five faunal types. A comparison has been

made with the more static method of classification into distributional types.

R. DE JoNG: Pyrgus in the Palaearctic region 109

Vn |

Su Ay

à om dl PY

dI N Map 1 - Pyrgus medal

re nn au

| FR = - 3 de

(i) Hi

\ p WC, È

N Y | x

dI 1 Map 2 - Pyrgus bieti

Map 3 - Pyrgus oberthuri

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

ul a

a 3 gi

4 > À :

AF” AM m

EN ;

DIA È

RM i Dy

1 fai z

> i 4 “

Pi ta Ss

= e

ù i Map 5 - Pyrgus carthami

A Lo - TARR MOL VA \

A 5 Ge LS =

i N im NN | Gr rer

a, LD

\ NS

Map 6 - Pyrgus sidae

R. DE JONG: Pyrgus in the Palaearctic region 111

VILA À

UE a

LAN a)

gar i Map 7

=“ |

\ NND Superspecies Pyrgus alpinus

N SAT

È \ aq Pyrgus alpinus

& Pyrgus cashmirensis

d @® Pyrgus darwazicus

MB Pyrgus cacaliae

& Pyrgus centaureae

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

p SUS EG x + >

Arme

it Kp 5 Da til Il Im | TT EA TT ‘i

ll) Mi [A

9 ng il) \ f+ i 2 x N IN 4) Hl)

ls f

\ TR Be e di

i Map 10

D N Superspecies Pyrgus malvae

D Pyrgus malvae

& Pyrgus malvoides

& Pyrgus melotis

ji]

Pi)

& FA i el ii > | \ |

Ze pa x

"==

Map 12 - Pyrgus onopordi

R. DE JONG: Pyrgus in the Palaearctic region 113

27 R Ts

À à II x

LI (à 2 A

È 0 u) È

En I,

SAS OSE 2

0” Map 13

‘Superspécies Pyrgus carlinae

& Pyrgus carlinae

@ e Pyrgus cirsii

IN OP fons

| ISIS

“A se %

Peak a Gs

22% È va Pm U N |

mia m

RU || 62

DINE

Map 15- Pyrgus armoricanus

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

) fs => pap = nea oy

a) > EN x > Im i

‘I FO à In i

Je {N LI in

AT Pyrgus warrenensis

ù E qD Pyrgus jupei

R. DE JONG: Pyrgus in the Palaearctic region 115

7. LITERATURE

Agenjo, R., 1952. — Fäunula Lepidopterolögica Almeriense. — Cons. Sup. Invest. Cient., Madrid.

1963. — Distribution geografica y morfologica del Pyrgus alveus (Hb., 1802) en Espana.

— Eos 39: 7—22.

Alberti, B., 1927. — Bemerkungen zu B.C.S. Warrens “Monograph of the tribe Hesperiidi (Euro-

pean species)". — Zeitschr. f. wiss. Insektenbiol. 22: 113—124.

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—, 1939. — Hesperia badachschana nov. sp. — Entom. Rundschau 56: 107—109.

—, 1940. — Ueber einige neue oder wenig bekannte palaearktische Hesperiden nebst Revision

der Hesperia staudingeri-Gruppe (Hesperiden-Studien, 4. Mitteilung). — Mitt. Münch.

Ent. Ges. 30: 235—254.

-——, 1952a. — Das Variationsbild der Pyrgus alpina Ersch.-Gruppe (Hesperiidae). — Zeitschr.

Lepid. 2: 77—87.

—, 1952b. — Wie weit geht Pyrgus malvoides Elw. und Edw. im Inntal abwärts? (Lep.

Hesperiidae). — Entom. Zeitschr. 61: 187—188.

—, 1952c. — Die Deutung der Rösel'schen Pyrgus-Figur von 1746. — Nachr.bl. Bayer. Entom.

1: 65—67.

——, 1953a. — Eine Lepidopteren-Ausbeute aus Südost-Polen. — Zeitschr. Lepid. 3: 51—58.

—, 1953b. — Die Deutung der Urabbildungen von Pyrgus carthami Hübner (Lep. Hesperii-

dae). — Nachr.bl. Bayer. Entom. 2: 37—40.

—, 1955a. — Eine neue Vikariante der Pyrgus malvae L.-Gruppe (Lep. Hesperiidae). — Zeit-

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N.F. 2: 211—224.

—, 1956. — Zur Frage der Bastard-Bildung zwischen Pyrgus malvae L. und malvoides Elw. &

Edw. (Lep. Hesperiidae). — Zeitschr. Wiener Entom. Ges. 41: 301—306.

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—, 1967. — Ueber die Hesperia alveus Hbn.-Gruppe im Kaukasus-Raum nebst Beschreibung

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Alberti, B. & J. Soffner, 1962. — Zur Kenntnis der Lepidopteren-Fauna Siid- und Siidostrusslands.

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Amsel, H. G., 1933. — Die Lepidopteren Palästinas. — Zoogeographica 2: 1—146.

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116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

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R. DE JONG: Pyrgus in the Palaearctic region 117

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R. DE JoNG: Pyrgus in the Palaearctic region 119

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120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972

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R. DE JONG: Pyrgus in the Palaearctic region

INDEX

of names of species, subspecies and forms of Pyrgus

accretus 17, 19, 88

adkini 97

albistrigus 31

alpinus 13, 47, 48

alticolus 91

alveoides 71

alveus 17, 19, 85, 90

analogus 41

andromedae 13, 51

armoricanus 17, 80

arvernensis 70

atratus 77

badachschanus 13, 46

balcanicus 41, 70

bellieri 17, 96

bieti 13, 33

bocki 32

cacaliae 13, 52

cacaoticus 83

caecus (cacaliae) 53

caecus (serratulae) 69

carlinae 24, 75

carthami 13, 36, 42

cashmirensis 13, 47, 49

celsimontius 51

centaureae 13, 54

centralhispaniae 19, 87

centralitaliae 92

chapmani 56

cinarae 17, 80

cirsii 77

claralveus 91

clorinda 79

colurnus 21, 90

conyzae 75

coreanus 63

corsicus 83

cottianus 77

darwazicus 13, 47, 50

deficiens 39

dejeani 13, 35

delavayi 35

diniensis 70

disjunctus 83

elegantior 62

evansi 46

fabressei 78

foulquieri 97

freija 57

fritillarius 36

fritillum 77

fulvoinspersus 83

gaillardi 97

grandis 89

grisescens 72

herrichii 78

hospitalis 89

ibericus 78

iliensis 17, 21, 92

infraobscuratus 71

insigniamiscens 87

jordana 67

jupei 17, 18, 98

jurassicus 89

kansuensis 17

kauffmanni 63

kotzschi 71

lecerfi 84

lilliput 51

loki 57

lucasi 39

luciarius 63

lutetianus 88

maculatus 13, 31

magnagallicus 70

major Fabricius (carthami) 70

major Rebel (carthami) 41

major (serratulae) 70, 71

malvae 15, 58, 61

malvoides 15, 63

maroccanus 82

melotiformis 65

melotis 15, 66

microcarthami 38

modestior 65

moeschleri 42

mustagatae 48

necaccreta 88

nemausensis 39

nevadensis 38

nigrocarens 78

nigropictus 97

numidus 17, 19, 86

oberthuri 13, 34

occidentalis 70

References indicate descriptions of characters.

occiduus 44

ochroides 77

onopordi 24, 74

parafabressei 78

persicus 81, 84

philonides 84

picenus 97

planorum 69

ponticus 15, 67

prabornius 91

prosensis 53

prostanae 84

pseudoalpinus 49

pyrenaeus 53

pyrenaicus (carthami) 38

pyrenaicus (malvoides) 38, 65

pyreneialpium 88

reverdini (alveus) 17, 21, 93

reverdini (armoricanus) 84

rufosaturus 83

ryffelensis 91

scandinavicus 17, 21, 91, 94

schansiensis 17, 22, 99

schoyeni 55

scotti 21

semiconfluens 65

septentrionalis 42

serratulae 23, 67

sibiricus 55

siciliae 83

sidae 13, 43, 45

sifanicus 17, 19, 92, 93

speyeri 17, 22, 94

subdecoloratus 89

taras 63

thibetanus 33

tramelensis 78

trebevicensis 91

tripuncta 55

turcivola 78

uralensis 72

valesiacus 41

warrenensis 17, 18, 98

wyandot 57

yunnanus 34

zona 31

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT 1

Plate 1. Fig. 1—3. Pyrgus maculatus maculatus. 1. Spring form, &, Amdo; 2. Spring form, à,

Shikoku, Japan, 27.111.1954; 3. Summer form, &, Tai-shan, Shantung, 1.VII.1934. Fig. 4—6. Pyr-

gus maculatus bocki. 4. Spring form, 4, Tien-Mu-Shan, Chekiang, middle of April, 1936; 5. @

Chang Yang; 6. Summer form, &, Shaowu, Fukien, 25.V.1937. Fig. 7—8. Pyrgus maculatus thibe-

tanus. 7. &, Tsekou; 8. 2, Ta Tsien Lou. Fig. 9. Pyrgus bieti bieti. &, Ta Tsien Lou. Fig. 10—11.

Pyrgus bieti yunnanus. 10. &, Li-kiang, Yunnan; 11. 9, N Yunnan. Fig. 12. Pyrgus bieti bieti

Q, Ta Tsien Lou. Fig. 13—14. Pyrgus oberthuri oberthuri. 13. &, Ta Tsien Lou; 14. 9, Wa-Ssu-

Kow. Fig. 15. Pyrgus badachschanus. 2, Upper Panjshir Valley, Hindu-Kush. Fig. 16—17. Pyrgus

dejeani. 16. &, Nan Shan; 17. ®, Nan Shan. Fig. 18. Pyrgus darwazicus lilliput. 3, Bala Quran,

Anjuman Valley, NE Afghanistan; paratype. (All specimens natural size)

R. DE JONG: Pyrgus in the Palaearctic region

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT 2

Plate 2. Fig. 19—36. Underside of specimens on Plate 1 Fig. 1—18. Fig. 19—21. Pyrgus maculatus

maculatus. Fig. 22—24. Pyrgus maculatus bocki. Fig. 25—26. Pyrgus maculatus thibetanus. Fig. 27.

Pyrgus bieti bieti. Fig. 28—29. Pyrgus bieti yunnanus. Fig. 30. Pyrgus bieti bieti. Fig. 31—32.

Pyrgus oberthuri oberthuri. Fig. 33. Pyrgus badachschanus. Fig. 34—35. Pyrgus dejeani. Fig. 36

Pyrgus darwazicus lilliput. Further data, see Plate 1

R. DE JONG: Pyrgus in the Palaearctic region

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT

Plate 3. Fig. 37—54. Pyrgus carthami. 37. Ssp. nevadensis, &, Sierra Nevada. 38. Ssp. pyrenarcus,

4, Panticosa, prov. Huesca. 39. Ssp. analogus, &, Trebevic, Bosnia. 40. Ssp. valesiacus, 2,

Gomagoi, Ortler, N Italy. 41. Ssp. valesiacus, &, Vintschgau, S Tirol. 42. Ssp. septentrionalis, 8,

Hennigsdorf near Berlin. 43. Ssp. carthami, &, Marchfeld, Lower Austria. 44. Ssp. carthami, &,

Budapest. 45. Ssp. moeschleri, ©, Kisilskaja, E Russia. Fig. 46—54. Underside of specimens of

Fig. 37—45. (All specimens natural size)

R. DE JONG: Pyrgus in the Palaearctic region

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT 4

Plate 4. Fig. 55—60. Pyrgus sidae. 55. Ssp. occiduns, &, Mouan, S France. 56. Ssp. occiduus, 2,

St. Barnabé, S France. 57. Ssp. evansi, &, Askhabad, Transcaspian (paratype). 58—60. Ssp. sidae.

58. 4, Kotor, Dalmatia. 59. &, Aksehir, Anatolia. 60. 4, Sliven, Bulgaria. Fig. 61. Pyrgus malvae

f. taras, 3, Wahner Heide, W Germany. Fig. 62. Pyrgus warrenensis, 4, Ortler, N Italy. Fig. 63.

Pyrgus jupei, &, Passanauri, C Caucasus. Fig. 64—72. Underside of specimens of Fig. 55—63.

(All specimens natural size)

R. DE JONG: Pyrgus in the Palaearctic region

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115,

AFL. 1, 1972

Plate 5. Fig. 73—90. Pyrgus serratulae. 73. Ssp. serratulae f. caecus, &, Val di Campo, Switzerland

74. Ssp. serratulae f. serratulae, &, Vintschgau, S Tirol. 75. Ssp. serratulae f. magnagallicus, ®,

Vendée. 76. Ssp. serratulae, &

î, Sierra de Gredos, C Spain. 77. Ssp. balcanicus, &, Treska, Mace-

donia. 78. Ssp. major, 4, Aksehir, Anatolia. 79. Ssp kotzschi, &, Kuh-i-Mirabi, NE Iran (para-

type). 80. Ssp. grisescens, 3, Passanauri, C Caucasus. 81. Ssp uralensis, 4, Uralsk. Fig. 82—90.

Underside of specimens of Fig. 73—81. (All specimens natural size)

R. DE JONG: Pyrgus in the Palaearctic region

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT 6

106 © 107 108

Plate 6. Fig. 91—108. Pyrgus alveus. 91. Ssp. numidus, &, Djebel Anngour, Morocco. 92. Ssp.

insigniamiscens, 2, Sierra Nevada. 93. Ssp. accretus, &, Gerona, E Pyrenees. 94. Ssp. accretus, &,

Forêt St. Germain, Seine-et-Oise. 95. Ssp. alveus, 4, Hersbrucker Jura, Bavaria. 96. Ssp. alveus

f. alticolus, &, Venter Tal, Ötztaler Alpen. 97. Ssp. sifanicus, &, Sining. 98. Ssp. reverdini, &, Ta

Tsien Lou. 99. Ssp. speyeri, &, Kuldja. Fig. 100—108. Underside of specimens of Fig. 91—99.

(All specimens natural size)

R. DE JONG: Pyrgus in the Palaearctic region

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No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov

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R. T. Simon THOMAS & A. M. J. SIMON THOMAS. — Some observations on the be-

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Philanthus triangulum (F.) (Hymenoptera, Sphecidae), pp. 141—152, Fig. 1—3.

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SOME OBSERVATIONS ON THE BEHAVIOUR OF FEMALES OF

PHILANTHUS TRIANGULUM (F.)

(HYMENOPTERA, SPHECIDAE) ')

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS

Pharmacological Laboratory of the University of Amsterdam

In connection with research on the mechanism of action and the chemical structure of

paralysing wasp venoms, females of Philanthus triangulum (F.) (Sphecidae, Hym.) have

been collected in the southwestern part of France since 1965. Since these animals can

only be caught during the summer, attempts have been made to breed them in the

laboratory (Simon Thomas 1966), but failure to bring about mating under laboratory

conditions has made continuous breeding impossible. In the search for a better breeding

method, the behaviour of P. triangulum in the field was investigated.

Ordinarily, Philanthus triangulum preys on honey bees and, according to Beekhuis van

Till (1935), occasionally on solitary bees (Andrena, Dasypoda, Megachile, and Halic-

tus). The females of P. triangulum feed on some of the paralysed bees, sometimes by

pressing the honey from the bee’s stomach by means of rhythmic abdominal move-

ments, sometimes by biting the prey just behind the first pair of legs and sucking the

haemolymph (Hirschfelder 1952). The majority of the captured, paralysed bees is used

for food for the larvae.

The behaviour of Philanthus triangulum has been described by Tinbergen (1932) and

van Beusekom (1946), who investigated orientation, and by Beekhuis van Till (1935),

who gave a detailed description of the reproductive behaviour. Other data have been

collected by Bouwman (1927), Bouvier (1916), Picard (1903), Roth (1917), and

Vergne (1935). However, these studies do not provide a complete picture of the

behaviour of this insect and we therefore undertook further research between 1968 and

1970. Since P. triangulum had disappeared from The Netherlands in 1960, the observa-

tions were made in southwestern France near Linxe (Department of Landes) in an

almost flat, sandy area, an excellent biotope for this wasp. In 1968 a colony near Linxe

was studied and in 1969 and 1970 another colony near Naboude (Fig. 1). Most of the

observations were made in 1969 on the colony near Naboude, situated on a sandy cart

track running more or less East-West. The nests were located between the ruts in the

untravelled middle part of this track, and also on both verges. The colony received

sunlight throughout most of the day. On both sides of the track, there was a stand of

pines (Pinus pinaster Ait.), about 5 m high, with a ground cover of heather (Erica

cinerea L.) and furze (Ulex namis Forst. and U. europaeus R.).

1) Supported in part by the European Research Office, United States Army, London, United King-

dom of Great Britain.

123

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

METHODS

During the observation periods, the temperature and humidity were registered with a

thermo-hygrograph mounted 2 m above the ground. Unfortunately, these measurements

had to be made at a distance of 8 km from the research area. Data obtained from the

meteorological stations of Cazaux and Mont-de-Marsan were also taken into account. The

heat content (the sum of the air temperatures in degrees Centigrade above 25°, added

per hour between 10 a.m. and 6 p.m.) was calculated for all the observation days. On

the basis of the heat content the days were classified as hot (heat content > 50),

moderately hot (heat content between 50 and 0), and cold days (heat content 0, i.e.

maximum temperature < 25° C).

BORDEAUX

GIRONDE

A tl an t tq ule

Oc Te ann

Mont-de-

Marsan

O Bayonne

Fig. 1. Location of the study area, near Naboude in southwestern France

For the study of the behaviour of individual wasps, the thorax was marked (Simon

Thomas 1970). If two lines were used, the colour of the left line is mentioned first; if

the marking consisted of a dot in a circle, the notation sequence is as follows, C (circle),

colour of the circle, colour of the dot (e.g. CYR). Nests of individual females were

marked with one or more coloured marking pins.

Casts were made of six nests, by filling them with prevulcanized latex, a product

which can be mixed with water in any given proportion to obtain the desired viscosity.

After the material had hardened the casts were dug out carefully.

In 1969 the observations were made continuously from July 7—28 and from August

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

12.”

17? -19

14 16

August

21. 23 25 UA 3775 ed

a EE CS CE u) a

B1 B1 CYR

S5——S5

G7 G7

BY ?

O2-O2.CYR CYR ?

AG——AG

GP GP

—R

Y—— Y

\Y—— nn YR2—— R2

CG —CG

K MR

CG—CG.CW —CW

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus 2 127

1—5, as well as on August 11, starting in the morning before the first females opened

their nests until the evening, after all the nests had been closed for the night. In 1970

the observations were made continuously from June 17th through July 17th. To find out

whether the nests were opened during the night, white sand was strewn on the outer

closure.

The research mainly concerned the behaviour of females; males were observed only

when they returned to their sleeping holes, in the evening or at the approach of a shower.

RESULTS

Table 1 shows the climatological data for the periods of observation in 1969 and 1970,

including the heat content and the incidence of rain. No data on climate conditions are

available for 1968.

In 1968 the first male was seen on June 12th and the first female on June 19th. For

1969, these dates were June 22nd and 24th and for 1970, June 19th and 29th, res-

pectively.

General description of the behaviour on a moderately hot day

The females spend the night in their own nests, which are kept closed (Fig. 2). In

the morning the nest is opened. The female emerges, usually starts digging the sand

away from the entrance, and then flies off, either to suck nectar or to capture a bee. All

day long, paralysed bees are carried into the nests. At the end of the day the female digs

for a long time and removes the sand lying in front of the entrance. Finally, the nest is

closed with a plug of sand pushed outwards. Fig. 3 shows the location of the nests at the

observation site, the duration of the periods during which the nests were occupied, and

the colour code of the female then living in a particular nest (1969). It is evident from

Fig. 3 that many females occupied the same nest for a long period of time.

Every form of activity proved to be influenced by temperature. Rain strongly reduces

any activity. On rainy days no bees are caught.

Detailed description of a number of behaviour patterns

a. Awakening.

Under awaking is understood the first perceptible activity of the female of Philanthus

triangulum, which consists of removing the outer plug of sand from the entrance of the

nest. After the opening of the nest, the female’s head appears in the entrance and after

some time the female emerges. In cold weather it stays in the entrance for a long time

Table 1. Climatological data in Naboude (France) during the observation periods of 1969 and 1970;

heat content is the sum of the air temperatures in degrees Centigrade above 25, added per hour

between 10 a.m. and 6 p.m.

Heat content 1969 1970

Number of observation days 24 31

Number of hot days 150 3 (12.5 %) 2 (6%)

Number of moderatiely hot days <50, > 0 15 (62.5 %) 12 (39%)

Number of cold days <0 6 (25%) 17 (55 %)

Number of rainy days 3 (12.5 %) 9 (27%)

Fig. 3. A. Location of the 19 nests between the ruts in a cart-track near Naboude. At the left, a

cross-section of the track. Arrows point in the direction of the burrow. B. Periods during which the

females occupied the 19 nests

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

after having removed the outer plug. The time of awakening varies from specimen to

specimen and per specimen, from day to day. Although no clear relationship with tem-

perature was demonstrable, the impression was gained that the appearance is delayed

after a cold night. Also, the animals appear to awake earlier on hot days than on cold

days.

b. Digging.

The nest dug by Philanthus triangulum can be recognized by the shape of the mound

(the cone of sand situated in front of the entrance). This mound roughly resembles that

of Dasypoda hirtipes L. (Apidae, Hym.), although in the case of the latter the entrance

lies within the mound.

As soon as a female has hatched, it starts to look for a convenient nesting spot in the

immediate vicinity. Olberg (1953) mentions as the origin of what he calls “pseudo

colonies” the wasps’ preference for the best-situated localities, which are of course

limited in number. The female flies and walks about, and tries to dig in every open

sandy spot. Sometimes it attempts to take possession of an existing nest or a male’s

sleeping hole. After a trial period lasting from a few hours to half a day at most, the

female starts digging. Sometimes a sleeping hole may be dug for the night only, and is

then dug out to a definite nest next day. The total digging period lasts from one to three

days (Table 2). When a female takes possession of an existing nest, it also digs, but for

a shorter time. When a nest is usurped, the owner is usually chased away (Table 2, CYR

and R2). Once an intruder has taken possession of a nest, the original owner hardly ever

succeeds in recovering it, althrough it often keeps wandering about in the neighbourhood

a whole day, repeatedly trying to re-enter the nest.

The nests are generally dug rather far apart. When the distance between two nests is

less than 10 cm the females tend to fight, seizing each other with their jaws, although

this never leads to injury (Fig. 3A, nests 17 and 18). Only once was a female seen

bringing a bee to a neighbouring nest.

The tarsae of the first pair of legs are equipped with a row of spines forming a comb

with which the sand is moved. When digging, the female usually stands high on the

Table 2. Time needed by a female of Philanthus triangulum (F.) to prepare a nest

wasp start of digging first bee brought in ,

no. date time date time

new nest S6 25/6/68 18.10 28/6/68 16.45

new nest R9 25/6/68 18.20 28/6/68 14.45

new nest G15 27/6/68 16.30 29/6/68 13.16

new nest O19 28/6/68 + 18.00 30/6/68 17.21

new nest $20 28/6/68 18.49 30/6/68 18.25

new nest R22 29/6/68 18.04 30/6/68 18.29

new nest B23 29/6/68 evening 4/7/68 12.25

new nest O2 18/7/69 18.20 19/7/69 15.13

new nest S5 1/8/69 9.40 3/8/69 13.41

new nest O 10/7/70 19.05 11/7/70 18.35

occupation of abandoned nest B1 23/7/69 10.34 23/7/69 13.05

occupation of abandoned nest G 15/7/69 9.45 15/7/69 12.26

occupation of abandoned nest B 3/7/70 16.58 4/7/70 16.27

usurpation CYR 16/7/69 18.56 17/7/69 18.05

usurpation R2 25/7/69 18.16 26/7/69 16.25

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus ® 129

entrance

outer closure

Pr +

main burrow

cocoon x

2 1

LT + ESS !

a) Base + cell

ROSE. DI

C Se

Fig. 4. Generalized nest of Philanthus triangulum (F.); A. side view; B. seen from above; C. cross

section through horizontal part of main burrow

middle and hind legs and thrusts the sand backwards under the abdomen. Hard obstacles

are dislodged with the jaws and dragged away by walking backwards. When this is not

possible the insect digs around the obstacle. In removing loose sand the female first

walks backwards away from the entrance as far as the edge of the mound and then starts

digging furiously while walking forward in the direction of the entrance. The mound

obtains its specific form because in walking backwards from the nest, the animal chooses

different directions. The outer closure at the entrance is formed of sand pushed by the

abdomen from the inside of the nest towards the entrance.

The nest consists of a main burrow running downward with a slope of 30° and

varying in length from 15 to 25 cm (Fig. 4). Occasionally, after a short steep descent,

the tunnel becomes horizontal. The brood cells lie on both sides of the horizontal

segment. When a cell is filled with bees and an egg has been deposited, the cell is closed

(inner closure, see Fig. 4). A second horizontal tunnel is sometimes dug from the end

of the slanting section, usually about 10 cm deeper in the sand.

In 1969 the total digging activity (in minutes per 15-minute period) was determined

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

CO ai GONE

fas

= 8

5 4

[=]

8 am. 10 12 2 4 6 8 p.m.

Fig. 5. Digging activity (in minutes) of a number of females in the course of the day, divided into

classes of 15 minutes

for a number of females (Fig. 5). By total digging activity is meant: (1) the removal

of sand in front of the entrance of the nest, (2) the throwing out of loose sand from

inside the nest, and (3) the time spent by the wasp inside the nest apparently busy

digging underground (regular expulsion of sand). Fig. 5 shows that the main digging

activity occurred between 7 and 11 a.m. (31.7 % of the total) and between 5.30 and

9.00 p.m. (65.8 % of the total). Between noon and 5.15 p.m. only 2.5 % of the digging

activity took place. This activity is lower on cold than on hot days and varies between

individuals. The abnormally high digging activity shown by the females occupying nests

17 and 18 (Fig. 3A) is explained by the fact that their nests faced each other at a

distance of about 6 cm, so that the sand removed by one female continually blocked the

entrance to the nest of the other.

c. Foraging.

After awaking and digging, the females fly off. The average duration of this first

flight was 31 + 2 (SD) minutes (n = 175). Out of 175 cases the wasp came back in

32 without a bee after the first flight (see e.g. Fig. 2A) and in 143 with a bee (see e.g.

Fig. 2B and C). The average duration of later flights was 20 + 1 (SD) minutes (n =

269). These later flights are thus considerably shorter than the first one (P < 0.005).

Evidently, the females also use the first flight to provide for their own food supply; they

are often found on blackberry flowers in search of nectar.

d. Orientation.

When a female flies out, it often performs an orientation flight in the shape of a

figure eight with the point of intersection of the loops just over the entrance of the nest.

While flying over the nest, the wasp’s head is directed towards it. The loops gradually

become wider and successively higher. Fig. 6 shows the incidence of the orientation

flights (in percent), plotted against the total number of flights for all females. When

the number of flights is low, the percentage of orientation flights is about 100. When

the number of flights increases, the percentage of orientation flights drops to about 20.

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus 9 131

100

orientation flights as percentage of total flights

(o) 50 100

number of flights

Fig. 6. Percentage of orientation flights plotted against total number of flights, for 19 females

For eight females together a homogeneity test demonstrated that the distribution of the

percentage of orientation flights over the day was not homogeneous, x? — 48.929 (21

degrees of freedom), P < 0.001 (cf. Table 3). Four females (R, Y, BY, and AG)

form a homogeneous group, x? = 4.791 (9 d.f.), 0.75 < P < 0.9. In this group there

is no significant difference between the percentage of orientation flights for the morning

Table 3. Number of flights, with and without orientation flights, made by eight females in the

morning and afternoon

wasp morning afternoon total no. of

no. with without with without flights

R 4 (29 %) 10 17 (37 %) 29 60

dg 7 (30 %) 16 19 (29 %) 47 89

BY 11 (37%) 19 25 (41 %) 36 91

AG 6 (27 %) 16 19 (35 %) 36 77

GP 8 (29 %) 20 12 (16 %) 63 103

YW 10 (40 %) 15 13 (15 %) 74 112

RG 5 (45 %) 6 17 (22%) 61 89

CYR 13 (72%) 5 23 (37%) 39 80

64 107 145 385 701

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

5 15 A

o 10

—

mR ©

£ 0 €

= 60 120 240 360 min

2 40

Ro)

2 20

30 60 90 min

Fig. 7. Intervals between successive homing of Philanthus triangulum with prey; A. nest closed;

B. nest left open

and the afternoon. The other four females cannot be placed in one group, x? = 28.476

(9 d.f.), P < 0.005, though they all showed a higher percentage of orientation flights

for the morning than for the afternoon.

The percentage of orientation flights does not seem to be related to the age of the

animal or to the heat content of the day in question. In the case of one female that dug a

new nest 10 cm from the old one, the percentage of orientation flights did not change.

e. Hunting for bees.

In 1969 the females under study each brought in an average of four bees a day. A

female coming home with a bee either immediately closes the entrance behind it (Fig. 2A

and B), or leaves it open (Fig. 2C, 2nd and 3rd bee). The time at which bees were

brought in was recorded. Fig. 7 shows the duration of the interval between two such

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus @ 133

—

number of units of watching

—

(0) 50 100

heat content above 20 °C

Fig. 8. Number of units “watching” per female per day plotted against heat content above 20° C

(encircled points are: one rainy and one cold day)

events. Fig. 7A shows the distribution of the intervals after which the nest was closed,

Fig. 7B that for instances when the nest was left open. For closed nests, the average

interval is 148 + 7 (SD) minutes (n = 243). For nests left open, the average interval

is only 29 + 1 (SD) minutes (n = 293). Fig. 7A also shows that sometimes the female

only stays inside briefly, even after closing the nest. This is quite usual for females that

have just dug a new nest (Fig. 2A) and possibly also when a female is alarmed while

entering. When the female leaves the nest open, a series of 2, 3, or 4 bees is brought in

successively at short intervals. Table 4 shows that there is a preference for a series of

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

Table 4. Distribution of bees captured in series by Philanthus triangulum (F.) compared with the

theoretical distribution

number of actual distribution of captured bees distribution according to equal chances

bees captured

in one series number of series number of bees number of series number of bees

1 284 284 343 343

2 131 262 142 284

3 104 312 57 171

4 13 52 28 112

910 910

three bees, and that a single bee is brought in far less often than could be expected on

the basis of equal chances. This behaviour creates the impression that the female “plans”

to collect a certain number of bees and leaves its nest open while bringing in this series.

In 1970 the average was two bees a day per female, possibly as a result of the cold

weather that summer.

f. Closing the nest on leaving.

When leaving the nest, the female rarely closes it. On some occasions, however, it is

closed so carefully that the entrance is practically invisible, and on others sand is hastily

scratched over the entrance (see e.g. Fig. 2B). During the 1969 observations, seven

females never closed their nests, five females did so only in 3 % of the cases, and one

female made a closure in 40 % of the cases. In 1970 closing of the nest from the outside

was observed only three times. Closure of the nest offers a defence against parasitic flies,

but the cuckoo wasp Hedychrum intermedium Dahlbom (Chrysididae, Hym.) is able to

open closed nests.

g. Dragging out paralysed bees.

Paralysed bees are frequently dragged out of the nest and dumped outside. During the

period of observation in 1969 these bees always were entirely fresh, with a full honey

stomach. Nineteen females ejected a total of 103 bees, having brought in 1,045 bees in

the same period. The removal of bees was observed 73 times. In 72 cases it occurred

during the digging period. About one-third of the removed bees proved to have been

parasitized by a fly.

h. “Watching”.

Especially on cold days the females spend a long time in the entrance. This is called

“watching”. Fig. 8 shows the frequency of this behaviour per day, plotted against the

heat content of the observation days. Each period of watching shorter than five minutes

was recorded as one unit. If a period of watching exceeded five minutes, it was divided

into periods of five minutes and each such period (eventually together with an additional

shorter period) was counted as one separate unit of watching. Fig. 8 shows that watching

increases at low temperatures whereas at higher temperatures the value asymptotically

approaches 2. The two encircled deviating points coincide with a rainy day and an abnor-

mally cold day, respectively. The average for all the females was 5.6 a day. The daily

average per female ranged from 2.1 to 9.8.

In 1970 the impression was gained that watching increased whenever cuckoo wasps

(Hedychrum intermedium) were present near the nest entrance.

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus ® 135

number of g/day

o >

number

of d/day

19 30 10 20 271 5

1969 June July August

Fig. 9. Numbers of females and males of Hedychrum intermedium Dahlbom (Chrysididae, Hym.)

collected in the observation area (Naboude, France), in 1969

Life span of the females.

In 1969, the first female was caught on June 25th. On June 26th, one female (R)

was marked. This female could be observed continuously. On July 26th and 27th, the

activities of this female decreased sharply. Digging activities no longer occurred. On

August Ist it flew out twice without bringing in a single bee. On August 2nd this female

was found in the heather near the nest. Upon being returned to the nest, it went in

quickly and was not seen again. Another female was marked on July 8th, and was still

active on August 11th. Thus, the lifespan of these females amounted to at least 37 and

34 days, respectively.

In 1970 two females were marked while they were looking for a place for a new nest,

one on July 2nd and the other on the 3rd. On August 12th both were recaptured, at

which time they were 41 and 40 days old, respectively.

Second generation.

During the collection performed between 1966 and 1968, it was observed that the

population clearly increased in August and that at the time the females looked less worn

out and appeared to be larger.

In 1969 a few marked nests, the owners of which were also marked, were kept under

observation. On August 2nd a male emerged from one nest and on August 4th an

unmarked female from another. On August 11th two unmarked females emerged from a

marked nest; a male and an unmarked female from another nest; two unmarked females

from a third nest, and one unmarked female from a fourth. The males of the second

generation as well as the females are not immediately chased away by the owner of the

nest. Once they have flown out, however, they are chased away when they return. On

the other hand, a young female looking for a nest occasionally chases away the old owner.

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

Parasites.

In 1969 there were many cuckoo wasps (Hedychrum intermedium Dahlbom, Chrysidi-

dae, Hym.) in the vicinity. They usually lie in wait in front of the other species’ nests

until its owner returns with a bee, to which the cuckoo wasps quickly attempt to attach

an egg. They may also open a nest and go inside. Fig. 9 shows the data on captures of

cuckoo wasps in the different periods. The male cuckoo wasp appears to hatch earlier

than the female, and the numbers of both sexes caught per day show a rapid initial

increase and then stay more or less constant. Since captured specimens were killed,

captures always concerned new individuals from outside the area of observation.

Observation of marked cuckoo wasps showed that they explore a rather large area and

visit nests continually. Marked cuckoo wasps were found at a distance of at least 100 m

from the release point. The marked cuckoo wasps were seen near the observed nests

several times a day. Apparently, new nests are found accidentally. When a Philanthus

triangulum comes flying slowly along with a bee, a waiting cuckoo wasp starts walking

quickly to and fro in front of the nest and often goes inside for a short time. The P.

triangulum is alarmed by this, often flying away and sitting on the branch of a pine tree

nearby, before attempting to enter the nest again. As long as the cuckoo wasp is there,

these attempts may be repeated several times. Finally, the P. triangulum quickly enters its

nest, followed by one or more cuckoo wasps. Occasionally, it drops the bee and goes

inside alone. When a P. triangulum wants to dig and sees cuckoo wasps in front of the

entrance, it drives them away. A fight may occur, although the intruder is never stung

or injured, because it rolls itself up. Cuckoo wasps stay equally frequently near closed

and open nests.

In 1970 an average of 2.6 cuckoo wasps kept watch throughout the day near each of

the nests under observation. The presence of so many parasites may cause a P. triangulum

population to die out almost completely in one year’s time, as was observed in the Forét

de Campet (Lot-et-Garonne): in 1965, P. triangulum was common in this area and there

were only a few cuckoo wasps; in 1966, P. triangulum was still common in July, but in

August showed no numerical increase even though the second generation had hatched.

The cuckoo wasps were much more abundant. In 1967, only a few P. triangulum were

observed and there was a very large number of cuckoo wasps.

Discussion.

In the present paper certain features of the behaviour of the female Philanthus trian-

gulum in and around its nest have been described in some detail. The observations con-

cerned awaking, “watching”, digging, bee hunting and orientation flights.

According to Tinbergen (1932), Philanthus triangulum may sometimes search for as

much as ten days for a suitable spot to dig its nest. In southern France this trial period

usually lasted half a day to one and a half days, though in cold weather it may take three

days. The much longer period reported for The Netherlands may be explained by con-

tinuous cold weather. This whole trial period is spent in searching and occasional

digging. Although the female does fly off to feed, it never returns with a bee.

Opinions concerning the number of bees deposited per cell vary considerably. Bouw-

man (1927) found 5 or 6 bees per cell, Tinbergen (1932) mentions 3 to 6, and Fabre

(1886) 1 or 2 bees for a male larva and 3, 4, 5, and 6 for a female larva. Vergne

(1935), who dug out many cells, always found two bees with a male larva and five bees

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus 9 137

with a female larva in a bigger cell. Olberg (1953) mentions 2, 3, or 4 bees for a male

larva and 3 to 6, but usually 5, for a female larva.

In our laboratory breeding experiments with P. triangulum (Simon Thomas 1966),

614 nests were dug out. The numbers of cells containing from 1 to 6 bees were 11, 254,

135, 60, 41, and 6 respectively (19 %, 41 %, 22 %, 10 %, 7 %, and 1%). Nests with

two bees are therefore the most common. Since all P. triangulum had already hatched,

it was impossible to determine whether the cells had contained males or females.

Evans (1964) reports that the female of the American Philanthus lepidus Cresson

buries her bees just inside the entrance and only fills the cells after enough bees have

been collected. Similar behaviour is not known for Philanthus triangulum. As shown in

Table 4, females prefer bringing three bees in succession and as a rule leave the nest

open in the meantime. In our laboratory experiments in which, as mentioned above, the

cells were mainly filled with two bees, the progeny showed a male-to-female ratio of

5—1. Thus, the supply of three bees in succession does not tally with the number of

bees per cell. Conceivably, Philanthus triangulum too is in the habit of burying its bees

inside the entrance temporarily, in which case it may not use all of them afterwards. This

is consistent with the fact that 10 % of the captured bees are dragged out again during

the digging period, (page 00), possibly representing the surplus left after the cells have

been filled. As already mentioned, the rejected bees are completely fresh, and the honey

stomach is often still full. Picard (1903), on the other hand, refers to dead bees he

observed outside the nests, but later he says that the females squeeze these bees dry,

which means that they were not dead at all but only paralysed.

A remarkable feature is that the females often sit “watching” in the entrance of the

nest for very long periods of time. Two possible explanations can be offered for this

behaviour. Firstly, this “watching” could be a reaction to disturbance, in this case the

observer's presence. Disturbance in the form of cuckoo wasps in front of the nest may

also induce “watching”. But a more likely explanation seems to be that females are

roused to activity by a sufficiently high ground-temperature but that the air-temperature

at the entrance turns out to be lower than the ground-temperature. This would cause a

conflict situation. We indeed observed that on cold mornings females did open their nest

but did not fly away. When the temperature remained low they sometimes sat “watching”

for hours.

Philanthus triangulum has very few predators. During the observations, only two

females (CG and CW) failed to return to their nests. The two oldest females in our

material lived for at least 40 or 41 days. The cause of death could not be verified.

Toward the end of the lifespan the wasps’ activity declined sharply within a few days,

after which they did not emerge from their nests again. This decline in activity showed a

strong resemblance to the syndrome caused by parasitism by Physocephala chrysorrhoea

Mg. (Conopidae, Dipt.). This fly lays its eggs on the abdomen of the adult wasp. The

larva bores into the wasp and eats the organs, starting with the least essential parts. We

have bred this fly four times from captured Philanthus triangulum females.

Beekhuis van Till (1935) found that in The Netherlands Philanthus triangulum has

two generations a year. Eggs laid in May give rise to adult animals at the end of July.

When we were collecting Philanthus triangulum in the southwestern part of France, it

struck us that there was generally a strong increase of the populations in August and that

the females then seemed larger and less worn out by flying. Vergne (1935), who made

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

observations near Clermont-Ferrand, however, found only one generation a year. Bouvier

(1916) mentions having seen several males and females emerging from the same nest

in the dunes of Lus-sur-Mer on July 28th, 1901. Only one of these females returned

with a bee. Bouvier concluded that Philanthus shows a transition to colony formation,

but this seems highly unlikely. He probably saw a second generation hatch, especially

since he stated that Philanthus is solitary before and after this period. According to our

observations, a second generation undoubtedly hatches at the end of July or the beginning

of august in southern France.

SUMMARY

To investigate conditions for breeding Philanthus triangulum (F.) (Sphecidae, Hym.)

successfully in the laboratory, an ethological field study was performed in southwestern

France. The results provide the following picture.

Each female digs a nest close to the spot where it has hatched. During the period

of reproduction, they stay in the same nest. Occasionally, owners are chased away by

other females searching for a good spot to dig a nest. The wasps spend the night in their

closed nests, and emerge at varying times in the morning. Foraging occurs mainly in the

morning during the first flight. Bee hunting (up to 10/9 /day) only occurs when the air

temperature is higher than 22° C. Bees are brought to the nest either singly or in series

of 2, 3, or 4. While a series is being brought in, the entrance is left open and between

two series it is closed. Series of three bees are preferred. The nest is seldom closed when

the wasp leaves. When they fly off, the females often perform an orientation flight. The

percentage of orientation flights is 100 when very few flights are made, and drops to 20

when there are many flights. Of the bees brought in, 10 % are dragged out again. The

females often sit “watching” in the entrance. These periods of “watching” are longer

when there are disturbances or during cold weather. The digging inside the nest and

pushing out of the sand takes place daily, occurring mainly at the end of the day but also

in the morning. The longest stated age of a female in this material was 41 days.

The main parasite is the cuckoo wasp Hedychrum intermedium Dahlbom (Chrysididae,

Hym.) which is probably able to extinguish a population almost completely in a few

seasons. In the southwestern part of France the second generation hatches some time

around the beginning of August depending on the summer temperature.

ACKNOWLEDGEMENTS

The authors’ warmest thanks are extended to Prof. C. van der Meer, Director of the

Pharmacological Laboratory of the University of Amsterdam, not only for his unflagging

interest, encouragement, and valuable criticism from the very beginning of the study

through the completion of the manuscript, but also for obtaining the funds necessary for

repeated research in France.

REFERENCES

Beekhuis van Till, F. W., 1935, Enige beknopte aantekeningen over het leven van de bijenwolf

(Philanthus triangulum F.). Maandschrift v. Bijenteelt 38: 258—269.

Beusekom, G. van, 1946, Over de oriëntatie van de bijenwolf (Philanthus triangulum Fabr.).

Eduard IJdo N.V., Leiden, 162 pp.

R. T. SIMON THOMAS & A. M. J. SIMON THOMAS : Behaviour of Philanthus ® 139

Bouvier, M. E. L., 1916, Quelques observations sur les philanthes. Ann. Institut Pasteur 30: 205—

208.

Bouwman, B. E., 1927, Graafwespen van Nederland. De Levende Natuur 32: 110—116.

Evans, H. E., 1964, Notes on the nesting behaviour of Philanthus lepidus Cresson (Hymenoptera

Sphecidae). Psyche 71: 142—149.

Fabre, J. H., 1886, Souvenirs Entomologiques IV. p. 191—218. Ch. Delagrave, Paris.

Hirschfelder, H., 1952, Zur Biologie des Bienenwolfes (Philanthus triangulum F.). Zeitschr. f.

Bienenf. I: 1—3.

Olberg, G., 1953, Der Bienenfeind Philanthus (Bienenwolf). Die Neue Brehm Biicherei, Akade-

mische Verlagsgesellschaft Geest & Portig K.-G., Leipzig, 85 pp.

Picard, F., 1903, Note sur l'instinct de philanthe apivore. La Feuille des Jeunes Naturalistes IV, 34e

Année, no. 397, p. 17.

Roth, P., 1917, Notes sur l'instinct et les moeurs du Philanthus Abdelkader Lep. St. Fargeau. Bull.

Soc. d'Hist. Nat. Afrique du Nord 8: 63—71.

Simon Thomas, R. T., 1966, A method of breeding Philanthus triangulum F. (Sphecidae, Hymeno-

ptera). Entom. Ber. 26: 114—116.

, 1970, Het individueel merken van Philanthus triangulum (F.). Entom. Ber. 30: 149—152.

Tinbergen, N., 1932, Über die Orientierung von Philanthus triangulum Fabr. Z. f. vergl. Physiolo-

gie 16: 305—334.

Vergne, M., 1935, Contribution a l’&thologie et au développement post-embryonnaire de quelques

Hymenopteres prédateurs (Sphégiens) en particulier de Philanthus Triangulum F. Cler-

mont-Ferrand, Imprimerie Moderne, 141 pp.

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NOTES ON THE BEHAVIOUR OF MALES OF PHILANTHUS

TRIANGULUM (F.) (HYMENOPTERA, SPHECIDAE)

R. T. SIMON THOMAS!) & E. P. R. POORTER

Pharmacological Laboratory of the University of Amsterdam

INTRODUCTION

Fertilized females of Philanthus triangulum (F.) (Sphecidae, Hym.) collected in the

field can produce normal offspring in the laboratory (Simon Thomas, 1966). However,

the females of the first laboratory generation gave rise to a second generation consisting

exclusively of males, thus indicating that fertilization had not taken place under the con-

ditions used.

In 1968, 1969, and 1970 the behaviour of females was studied in the field (Simon

Thomas and Simon Thomas 1972). Since this investigation provided no information

about where and how copulation takes place, the behaviour of the males was also studied.

This was done in the summer of 1969 in Les Landes (SW France), in an area where P.

triangulum occurred in large numbers. In 1970 some additional observations were made

by the first author in the same area.

The study area — about one hectare — was situated along a main road (route D 5)

near Naboude (see Simon Thomas and Simon Thomas 1972, Fig. 1). A cart track

running East-West offered the only bare sand in this area. The vegetation consisted

mainly of young pine trees (Pinus pinaster Ait.) mingled with various shrubs. Along

the main road there were several species of flowers on which the wasps were feeding.

The soil consisted of calciferous fine-grained sand mixed with some humus, and was

blackened by resin dust. The nests of the females were only found in the bare sand of

the cart rut (Fig. 1). Males were rarely seen in the vicinity; only a few had their

sleeping holes in the area. They were seen mainly on flowers along the road and among

the bordering vegetation.

To study the males individually, they were marked with a coloured spot of plastic

paint on the thorax (Simon Thomas 1970).

Table 1 gives some relevant data on the various observation periods. According to

these data, the observations concerned first-generation males only.

Both sexes of P. triangulum stay underground at night in burrows dug by themselves

or in available holes. The depth of the sleeping holes of the males ranges from 2 to 10

cm, and the entrance is entirely or partly closed from the inside. During the day, the

holes of the males are left open.

Most males return day after day to the same sleeping hole and reappear in the morning

at varying times. Feeding usually takes place for about an hour in the morning and late

in the afternoon and also during additional short periods during the middle of the day

(Fig. 2).

1) Supported in part by the European Research Office, United States Army, London, United King-

dom of Great Britain.

141

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

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Table 1. Miscellaneous data concerning the three observation periods in Naboude

1968 1969 1970

Observation periods: males 25/6—17/7 17/6—17/7

Observation periods: females 25/6—22/7 25/6—5/8 17/6—17/7

First male seen in the field 12/6 22/6 19/6

First female seen in the field 19/6 24/6 29/6

First male, second generation 2/8

First female, second generation 4/8

Lifespan male (in days) 33

Lifespan female (in days) 38

Many males occupy a territorial station and prefer a specific feeding site, which may

be located at considerable distance from this station. These observations indicate that

individual territorial males are familiar with the location of at least three sites in the

field: the territorial station, the feeding site, and the sleeping hole. The locations of the

respective sites of seven males are indicated on the map in Fig. 1. The area regularly

visited by the animal is a multiple of the size of the territorial area.

Several males return day after day to the same territorial station in the field, usually

some small pine tree. They defend these sites against conspecific males and react to other

moving insects by flying in their direction. The territorial males confine their attendance

and their defensive behaviour to the small tree and its immediate surroundings up to

about 50 cm beyond the outer twigs of the three.

RESULTS

Position and distribution of territorial stations.

The observation area had a vegetation of grass clumps and shrubs with a scattering of

young pine trees. The territorial stations were situated on small pine trees ranging in

height from 0.5 to 4 m, except two that were situated in a vegetation of heather and

bracken. The territorial stations measured about half a metre. In the choice of a territorial

station no preference for a particular type or site of pine tree could be distinguished.

Behaviour of territorial males in the presence of conspecific males.

The first reaction of the resident male to an intruder is to rush at it. When the intruder

turns away and flees, he may be chased by the resident male over a short distance. The

latter returns within a few seconds.

When the intruder is unwilling to leave, a complicated conflict may arise in which the

males circle around each other in an irregular and jerky manner. The resident male

repeatedly takes up a higher position in the air by flying upwards. The upward flight of

the resident male is followed by a downward zigzagging swinging flight by which the

intruder is re-encountered. The amplitude of this zigzagging decreases gradually. How-

ever, the intruder may follow the resident male in the upward flight and in that case the

wasps may start circling again. This makes it difficult for the observer to describe the

conflict in more detail, because it is impossible to tell the two wasps apart.

When the intruder does not follow the resident male in the upward flight the former

either keeps circling or starts hovering below, or it may alight on the substrate of the

territorial station. The intruder is usually re-encountered by the resident male when the

R. T. SIMON & E. P. R. POORTER : Behaviour of Philanthus & 145

% ber of observations

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latter comes zigzagging down. When the intruder keeps circling or starts hovering while

the resident male flies upward, the former dashes up at the descending resident male.

When the intruder has alighted on the substrate of the territorial station it may either

be chased away by the descending resident male or dash upward at it. The difference

is not always clear to the observer. We observed many zigzagging flights ending in

perching of the resident male while the intruder was perched nearby. The repeatedly

zigzagging ending in perching was also observed when the intruder had left in the

meantime. Physical contact of skirmishing males was never observed.

In a number of cases in which more than two males were present at the same territorial

station, small groups of skirmishing males were observed.

Attendance at the territorial stations.

Table 2 (data from 1969) shows the highest scores of the total numbers of males

recorded simultaneously during the day at the various territorial stations. If one or two of

the males were marked, this is also indicated. It is evident that the same territorial

stations were often visited by marked males for several days in succession. One male

(CW, station 6) attended its territory for a period of at least 14 days. The individually

marked males were seldom seen at other territorial stations than their own.

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

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R. T. SIMON & E. P. R. POORTER : Behaviour of Philanthus & 147

In many cases more than one male was present at the same territorial station. Fig. 3

(data from 1969) shows the relative frequencies of the numbers of males at different

hours in the course of the day. For each hour, the data are based on all observations of

“no male present”, “one male present”, etc., at all known territorial stations during the

period from June 29th up to and including July 6th, except for June 30th. This was a

period without rain and with moderate temperatures (maximum 20°—30° C). Fig. 3

shows that the largest number of simultaneously present males occurred between noon

and 4 p.m. In approximately 10 % of the observations during this period, no males were

seen.

Table 3 shows the number of times that visiting males were observed at 11 territorial

stations between noon and 4 p.m. in the presence or absence of the marked resident

males. The numbers in the last column show that in 107 cases the resident male was

present without visitors and in 104 cases with one or more visitors. Therefore, visitors

were present during 49 % of the observations in which resident males were present.

Table 3. Number of observations at which males were seen in the presence or absence of the marked

resident male (M = marked resident male; X — unmarked male) between noon and 4 p.m.

Territorial station 2 5 6 7 9710 12 15 16 17730

total

Marked male S YR CW G YS YB WG RY PG BR P “1

M 8 26 9 6 3 8 15 O12 4 7000107

MX ZUNE 120 ES TS ARI, 77

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no male 3 5 9 1 3 4 2 3 5; 0 6 41

262

When the resident male was absent, there were visiting males in 10 cases and no visitors

in 41 cases. Therefore, visitors were present during 20 % of the observations in which

the resident males were absent. According to a x? test, this difference is statistically

significant (P < 0.0005), and it may therefore be concluded that resident males attract

visiting males.

To find out whether residential and visiting males can be distinguished at the stations,

25 unmarked males were caught at 12 territorial stations in the afternoon of July 4th,

1969. The wasps were kept in a cage and supplied with honey and distilled water. The

males were marked blue (B*). One of these received a blue circle on its thorax and was

therefore individually recognizable (CB). In the afternoon of July 5th the blue-marked

wasps were released. Fourteen of these were seen on July 6th at 13 territorial stations

(cf. Table 2). Blue-marked males were seen for a number of consecutive days at 7

Table 2. Attendance of males of Philanthus triangulum at territorial stations. The code-letters in-

dicate the presence of individually marked males staying for at least one hour. Code-letters with an

asterisk indicate marked males that were not recognizable individually. The figures to the left of the

oblique bar indicate the number of observations made between noon and 4 p.m. The figures to the

right of the oblique bar indicate the maximum number of males (including marked males) observed

at one time. The figures in parentheses (July 4th) are the numbers of unmarked males caught and

released as blue males (B and CB) on the afternoon of July 5th.

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

territorial stations (nos. 9, 10, 17, 26, 27, 28, and 29). It seems likely that the blue-

marked males observed at a given territorial station, were always one and the same

individual. Since only unmarked males were present at stations 26, 27, 28, and 29 before

July 4th, and all unmarked males were captured at these stations, it is likely that the blue

males observed afterward were the original territorial males.

At stations 9, 10, and 17, marked territorial males were already present before the

blue-marked males were released. These territorial stations were visited during consecutive

days by blue-marked males, which eventually replaced the original territorial males.

Station 10, at which no male was caught originally, was occupied by a blue-marked male

on consecutive days. Before the afternoon of July 4th it was repeatedly noted that an

unmarked male flew to and fro between stations 9 and 10. On July 4th, two unmarked

males were caught at station 9. Therefore, one of the males caught at station 9 might

have been the future blue-marked male of station 10. At station 17 the original male

(BR) was replaced by a blue-marked male on July 6th. On July 8th two blue-marked

males were seen here.

These observations indicate that there are two kinds of males, territorial and visiting

males. There may be more than one residential male at the same territorial station, but

only for a short time.

The fact that only 7 out of the 24 males marked blue were seen again for more than

one day, means that most of the blue males were occasional intruders when they were

caught. This might also have been the case for several other individually marked males

caught in the period from June 27th through July 2nd and which did not return to the

station at which they were caught. A few of the latter were later seen occasionally on

flowers or near their sleeping holes. Other males appeared to have subsequently estab-

lished their own territory.

According to the data in Table 2, some territorial stations (11, 14, 24, 30, and 31)

were seldom visited by more than one male at the same time, whereas other territorial

stations (5, 9, 15, 17, 27, and 34) were visited frequently by several males simultane-

ously. The preference for certain territorial stations is illustrated by the data obtained at

station 27. On July 4th, six unmarked males were caught at this station and since no

more males were seen during the rest of the day, these seemed to be all the males present.

These six animals belonged to the group of blue-marked males released in the afternoon

of July sth. On July 5th, no males were observed at territorial station 27. On July 6th,

two blue males were seen, and on July 7, 8, 9, and 10, one blue male was seen. This

latter male may well have been the original resident male. New unmarked males kept

being attracted to territorial station 27, and in the period from July 6th through the

10th, this station had the highest score of males simultaneously present of all the territorial

stations under study (see Table 2).

The total number of males seen in 1970 in the field was much lower than the figure

for the year before. The only territorial station found was closely observed. It was

occupied for eleven days (June 24th—July 4th). All visitors were caught and marked

individually. During six of these eleven days, one male (OY, marked at this station on

June 24th) was seen at this territorial station either alone or with some other males. Six

visitors came to this station on July 2nd. Throughout the day, the owner OY was present.

On July 3rd another male (caught on blackberry bushes nearby and marked OP) took

possession of this territorial station for one day only. On the last day (July 4th), only

R. T. SIMON & E. P. R. POORTER : Behaviour of Philanthus & 149

two unmarked males were seen at this site. One of the marked visitors (BO) established

a new territorial station nearby.

These observations suggest that a male chooses its territorial station more or less at

random. Once established, this resident male attracts other males, and these may start

new territorial stations nearby.

Behaviour of territorial males in the absence of conspecific males.

A territorial station in the pine-clad area usually comprises one or more sites where the

animal sits or walks as well as an area around the station used for flying.

When a male is alone at its territorial station it is often perched on a grass blade, pine

needle, or some other part of the vegetation but never on the ground. Males may remain

perched for periods ranging from a few seconds to several minutes. When a male is

perched, his antennae are raised slightly. Perching is regularly alternated with walking

and flying. When a male is walking it moves hurriedly up and down grass blades, pine

needles, or branches, meanwhile flickering its antennae and tipping the substrate with

them. When a male is flying, it moves either in circles around its perch, or from perch

to perch, or between different walking sites.

When perching is alternated with flying, the duration of a period of locomotion is

relatively short, i.e. 10 seconds or less in 88 % of the observations, 10 to 20 seconds in

8%, more than 20 seconds in 4% (50 observations). The maximum duration of one

period of locomotion consisting exclusively of flying was 48 seconds.

When locomotion consists of flying as well as walking, the duration of that period is

relatively long, less than 10 seconds in 3 % of the observations, 10 to 20 seconds in 9 %,

20 to 50 seconds in 52 %, and more than 50 seconds in 36 % (33 observations). The

maximum duration of one period of locomotion was 91 seconds.

We never observed locomotion consisting exclusively of walking. Walking is always

interrupted by flying. The duration of periods of walking during this combined type of

locomotion was less than 10 seconds in 63 % of the observations, 10 to 20 seconds in

23 %, and more than 20 seconds in 15 % (87 observations), with a maximum of 48

seconds.

Mating behaviour.

The first mating was observed on June 29th, 1969 at 4.05 p.m. (31° C). It was

carried out by a marked territorial male (YR) at its territorial station (no. 5). During a

period of at least 14 minutes before the mating started, no other males were seen at this

station. At that time we were recording the activities of the male. Just before the mating

occurred, YR had been perched for 56 seconds. It then flew up, and had been circling

around the branches of its tree for 9 seconds when a female alighted on a grass blade at

a distance of about 75 cm from the flying male. Immediately after the female had landed,

YR flew straight at it, made a short turn behind it, and alighted on its back. A few

seconds later the male dismounted from the female and turned around. It then became

apparent that the male was attached by its genitals to those of the female. The copulating

pair faced in opposite directions (almost in a straight line). After 3 minutes and 25

seconds an unidentified fly dashed against the pair, which then flew up in copulating

position and was out of sight within a second. Ten minutes later YR was back at its

station, but without the female.

The second copulation observed by us occurred on July 2nd at 3.03 p.m. (26° C).

150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972

This copulation was also carried out by a marked territorial male (BR) at its territorial

station (no. 17). Copulation had already started when we discovered the animals. Dis-

turbed by our arrival, the pair flew off and alighted on a grass blade at the territorial

station. After we had taken a photograph, an unmarked conspecific male pounced upon

the pair, which caused it to tumble down. The female got grip on a grass blade, but the

male hung unsupported from the female with floundering legs. When the female shifted

its position, the male obtained support. Shortly afterward, the female wrenched itself free

and flew away.

The third copulation was seen outside the main study area on July 11th at 3.30 p.m. |

(23° C). This pair was disturbed by the observer and alighted on a grass blade still in

the copulating position. This copulation too took place at a known territorial station.

In 1970 no mating was observed.

Discussion

The observations described in the present paper show that males of Philanthus triangu-

lum (F.) may possess at least three orientation points at which they are present regularly,

viz. a sleeping hole, a feeding site, and a territorial station at which mating was observed.

These three sites may be separated by considerable distances.

The behaviour of the males at the territorial stations is of particular interest. In many

cases territorial stations were occupied by one and the same male for a considerable period

of time, after which another male might become resident at this station.

Besides the resident males, visiting males — often more than one at a time — were

present at many territorial stations. This resulted in skirmishes between resident and

visiting males, sometimes occurring in small groups.

On three occasions a female was seen to visit a territorial station, and copulation was

observed. In two of these cases it could be shown that copulation took place with the

resident male.

In analogy with various species of birds, e.g. Black Grouse (Lyrwrus tetrix L.) and Ruff

(Philomachus pugnax L.), the males of which display in groups, the territorial stations

of Philanthus triangulum could be considered ‘‘/e£s”, which means display-grounds or

arenas. Like those of the above-mentioned birds, the wasps’ leks are surrounded by inter-

territorial areas.

Our observations of displaying males agree with those of Vergne (1935), who noted

groups of male Philanthus triangulum on elder bushes in the Botanical Garden of

Clermont-Ferrand (France). Beekhuis van Till (1935), on the other hand, saw males

dancing in front of the nests of the females in Het Gooîs Natuurreservaat in The

Netherlands, and also observed copulations near the nests. During extensive observations

in Landes (France) from 1968 to 1970 no dancing males, let alone copulations, were

ever seen near the nests (Simon Thomas and Simon Thomas 1972).

In the summer of 1970, Philanthus triangulum was found again in The Netherlands

(Zonderwijk 1971) after an absence of about seven years. This was in Tienraij, in the

province of Limburg. On the whole, the behaviour of the males was similar to that

observed in France. However, in Tienraij one female nest was found inside a male

territorial station. The male frequently tried to copulate when the female passed by in

going to or coming from its nest. It is conceivable that Beekhuis van Till saw something

similar.

R. T. SIMON & E. P. R. POORTER : Behaviour of Philanthus & 151

SUMMARY

To investigate the circumstances under which the copulation of Philanthus triangulum

(F.) (Sphecidae, Hym.) occurs in the field, the behaviour of the males was studied near

Naboude (France).

The males dig individual shallow sleeping holes to which they return every night.

They also have special feeding sites. The warmest part of the day is spent at certain

other places unrelated to either sleeping or feeding. Observation of marked specimens

showed that males occupy these particular spots for a considerable number of days.

These site-attached males show territorial behaviour, and they attract other males (visi-

tors), which results in skirmishing between two or more males.

On three occasions females were seen to visit these territorial stations, and copulation

followed. In two of these cases it could be shown that copulation took place with the

resident male.

ACKNOWLEDGEMENTS

The authors are greatly indebted to Prof. C. van der Meer, Director of the Pharma-

cological Laboratory of the University of Amsterdam, for his valuable criticism during

the study and for his time-consuming critical reading of the manuscript.

REFERENCES

Beekhuis van Till, F. W., 1935, Enige beknopte aantekeningen over het leven van de bijenwolf

(Philanthus triangulum F.). Maandschrift v. Bijenteelt 38: 258—269.

Simon Thomas, R. T., 1966, A method of breeding Philanthus triangulum F. (Sphecidae, Hymeno-

ptera). Entom. Ber. 26: 114—116.

, 1970, Het individueel merken van Philanthus triangulum (F.). Entom. Ber. 30: 149—152.

, and A. M. J. Simon Thomas, 1972, Some observations on the behaviour of females of

Philanthus triangulum (F.). Tijdschr. Ent..115: 123—139.

Vergne, M., 1935, Contribution à l’éthologie et au développement post-embryonnaire de quelques

Hyménoptères prédateurs (Sphégiens) en particulier de Philanthus Triangulum F. Cler-

mont-Ferrand, Imprimerie Moderne, 141 pp.

Zonderwijk, P., 1971, Philanthus triangulum Fabricius (Hym., Sphecidae). Entom. Ber. 31: 79.

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No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende

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